1.

INSECTS AS HUMAN FOOD

1. INSECT CONSUMPTION FROM PRE-HISTORY

TO THE PRESENT DAY.

It has been known for many centuries that insects are eaten as
delicacies in many parts of the world. Reports have come down
from antiquity of insects, especially locusts, being eaten by primitive
peoples. Honey has been known as a prized food from time imme-
morial. Yet entomophagy, apart from honey consumption, has
always been regarded as a curiosity or as barbarism. Although a
number of recent authors such as NETOLITZKY (1918/20), BEQUAERT
(1921), HARDY et RICHET (1933), GOUROU (1947) and others have
hinted at the actual and potential nutritive value- of insects for
primitive man, the present writer began his study a few years ago
largely out of curiosity. It is actually astonishing that the real and
basic importance of insects as food for early and primitive man has
been !!O long ignored. One of the main reasons is that a fuller under-
standing of the requirements of a well-balanced diet and of its
necessary vitamin content, over and above the mere calorific value
of food, has only been gained in our days. The French Colonial
Service has played a leading role in investigating the actual diet of
tropical peoples, followed by those of the British and Dutch em-
pires. These studies revealed that very many of the primitive peoples
of Africa, Asia and America are underfed or live on unbalanced,
entirely unsatisfactory diets, with a serious shortage of either animal
fats, animal proteins or carbohydrates. This deficient or improperly
composed diet is today regarded as the main reason for the low
standard of life and vitality, for the lack of energy which so often
reduces the vital standard of men in hot climates, and also for
the lack of resistance to many diseases. The gathering of insects
has often helped to supplement grave dietary deficiencies either at
certain regular seasons of the year or in times of emergency such

7
F. S. Bodenheimer, Insects as Human Food
© Springer Science+Business Media Dordrecht 1951
as recurrent droughts. Their utilization as food was the proper
instinctive response to the physiological need for animal proteins,
fats or other substances which large quantities of insect food could
provide. We have insisted upon the words 'in emergencies or in
certain seasons.' The reason is simple. Most insects abound even in
the tropics only in certain seasons. Very few are available through-
out the year, and even if this is the case, as with ants or termites,
the desired form appears only during a short season. This fact is
important, as neither insects nor any other food is ever stored by
the food-gatherers or by the primitive hunters. The fact that some
insects may be available in the tropics at every season makes the
group as a whole still more important as a source of food, since
every individual insect species is usually of limited seasonal in-
cidence.
It is rather doubtful whether primitive man ever felt an instinctive
aversion against the eating of insects. Scores of writers have ex-
plained at great length how most of the vegetarian insects in them-
selves, by their environment and by their food habits, belong to the
cleanest of animals, actually being much cleaner than most other
animals which are served at our tables. There is no possible reason
to explain why insects should be more repulsive to man than dead
mice, snakes, snails or mussels, toads, shrimps or fish and many
other titbits of human gourmets of every race. No evidence suggests
that there is anything basically repellent about insects. If this were
so, the almost worldwide incidence of insect consumption by men
of all races would be difficult to explain. It is certainly a falsification
of facts, as the later chapters of this book will clearly show, to assume
that only extreme famine has reduced man here and there to such
depravity that he was forced to overcome his natural aversion and
to still the acute pangs of his hunger with insects. Even the stench
of fermenting insects described so vividly by ROESEL and FRISCH
in connection with decaying locusts would not suffice to explain
their exclusion from our daily food. Man, and even highly civilized
man, has overcome his aversion to unpleasant and repellent smells
in many other types of animal or vegetable food. The proper pre-
paration by roasting, cooking or drying will easily transform any
insect into a dish which at its best is often acclaimed as a rare dainty,
or recognized at its worst as an insipid, yet nutritive food.

8
 Since the times of the Hellenistic writers many students have been
convinced that insects eaten in large quantities or as the main
source of food would bring about lowered vitality, diseases and a
shortening of life. This attitude has continued down to modern
times. Even men like RILEY and HOWARD felt the need to invoke
special precautions, such as complete sterilization of the insects,
before eating them. And even in 1939 MILLS and PEPPER felt the
need to prove experimentally that the consumption of a few flour
beetles (Yribolium conJusum Duv.) did not cause the slightest
physiological disturbance in the four experimenting heroes. We
should, however, be careful not to ridicule lightly the conception
that certain special articles of food may induce certain diseases.
Nevertheless, at the Xth International Congress of Medicine at
Berlin, ]. HUTCHINSON could state, for instance, his opinion that
a heavy fish diet may very probably be the main cause of the local
spread of leprosy (vide C. H. ROBINSON 1900, p. 149.) This theory
was, of course, soon shown by special research on the question to be
entirely without foundation.
Certain insects are poisonous and are traditionally avoided. In
some of the aposematic species avoidance is perhaps instinctive, as
could be concluded from certain experiments of CARPENTER (192 I)
in which monkeys hesitated to pick up aposematic insect larvae
offered to them or even never touched them at all, whilst all other
insects were readily taken and crunched. Frequent reference is also
made to poisonous honey. Another rather curious case of a sickness
which may prove fatal after large-scale consumption of insects is
mentioned by G. BOUVIER (1945). He reports that locusts are
hunted at Lomani in the Belgian Congo at regular times by bow
and arrow, the special arrows having four divergent points; and
that they are caught by tons during locust invasions. On such occa-
sions the quantity of these insects which are devoured whole is
such that the legs with the sharp bristles on the tibias may cause
a serious stoppage of the intestines which proves fatal if no surgical
intervention is made. After some time these accidents grow more
rare, as the Negroes then pull off the legs and wings before frying
the insects in palm oil. Greediness may also prove fatal to monkeys.
BOUVIER saw several dead monkeys, the autopsy of which revealed
intestinal occlusion caused by masses of locust legs.

9
 And yet there cannot be the slightest doubt that, under normal
circumstances, the average white inhabitant of Europe or America
will refuse with horror any offer or temptation to taste insects. Or,
if he eventually agrees or decides to do so, he looks upon himself
as a martyr or hero of science. This conspicuous aversion is a pre-
judice acquired incidentally to the progress of civilization. Also our
aversion to eat raw meat, to eat the flesh of man, to eat raw fish,
etc. is by no means a primary aversion. It falls into the same cate-
gory as the refusal to eat insects. Equally, the eating of frog-legs,
snails, oysters, shrimps, frutti di mare and similar animal food is
just as repulsive to the great majority of people who are not ac-
customed to eat them, as would be the eating of insects. All over
Europe, however, we find children sucking the sweet crops from
the bodies of bumble-bees, honey-bees or certain ants, crunching
the meaty legs or rumps of grasshoppers or beetles, or eating with
delight the sweet honeydew excretions of aphids and other Homop-
tera, etc.
No, the aversion to insect food in Western civilization, though
an established fact, is nevertheless not based on a hereditary instinct.
It is established by custom and prejudice. But, of course, important
contributory factors must have helped to develop and maintain
this prejudice. In the conditions of modern European and North
American agriculture, man has not only, as a rule, sufficient to eat
without being driven to secure regular additions to his normal diet,
but his food is also well-balanced and adapted to the physiological
exigencies of his environment. Hand in hand with this establish-
ment of a high quality and adequate diet bq.sed upon intensive
cultivation, goes an enormous and progressive reduction in the
number of primary articles of food. A primitive food gatherer
cannot dispense with any gift which nature doles out to him; many
hundred~ of species of plants and animals make up his 'daily bread',
which exclud~s nothing edible and easily available. But modern
man is reduced to a very few species of cereals, tubers, fruits and
vegetables, in addition to the various products derived from a
limited number of domestic animals. More substantial food, such
as game or fish, may be occasionally added, and in these, as in
other fields, some rare dainties may be occasionally welcome. Yet,
the smaller game and the less stahle objects of food gathering have

10
disappeared and with them insects, for two reasons: the well-
balanced, normal diet eliminated any dietary deficiencies which
led to the desirability of insect eating. And further, food-gathering
becomes uneconomic when there is neither a market nor a favoured
place on the table for the special crop to be gathered. Finally, in
the moderate climates of Central Europe and North America the
insect component of food has certainly become extremely reduced
since historical times and perhaps never was developed to the same
extent as in tropical and sub-tropical regions.
In order to become an established article of food the object in
question has to be, at least seasonally or periodically, of sufficient
volume to form an important part of the diet. If, as will be shown
below, insects provide not only a sufficient bulk, but in addition
make good precisely those basic substances in which the normal diet
either at a certain season or in general is locally poor or deficient,
their importance becomes relatively even greater than that of their
bulk alone.
In all primitive human societies, from the early food-gatherers
and hunters to the early agriculturists and animal-breeding nomads,
special rituals are devoted to every important article of food to
ascertain, for instance, its abundance and the fertility of seeds or
to ensure a good supply of animals and game. Such rituals or magic
ceremonies are of worldwide distribution. They have found their
earliest documented expression in the famous pictures in caves or
on rocks of hunters from the Palaeolithic Age onwards to those of
the present day Bushmen of South Africa. But the ceremonial
paintings of the Australian aborigines represent a still more primi-
tive stage, that of the food-gatherers. In all primitive societies we
find totem groups. These are groups within the tribe which regard
a certain object, usually an animal, as their ancestor. It must be
stressed that this belief of transformation is quite material. The
traditions of the Australian totems leave no doubt that these an-
cestors were originally individuals of the totem animal, which at a
certain date in the remote past transformed themselves by their
own resolution into the first :men-ancestors of that totem. This
explains how the eating of the totem animal or plant is forbidden
to the members of each totem, or reduced to a very moderate con-
sumption in connection with certain totem ceremonies. Yet the

I I
totem rituals for bringing about fertility and abundance of the
totem anima or plant are regarded as of vital importance to the
whole tribe, whose other totem groups will thus be certain to enjoy
ample food. Every totem, while being extremely restricted in the
consumption of its own totem animal, contributes thus to the total
food production of the tribe as a whole. In all food-gatherers the
drawings of the totem animals are rather rough, primitive and
unnaturalistic, in contrast to the wonderful drawings of game
animals of the Palaeolithic and even of more recent hunters. In the
latter, the naturalistic achievements of the pictures were apparently
in direct correlation with the magic effect of these drawings on the
yield of the hunt.
It is of the utmost importance for the understanding of our
problem that the knowledge not only of a number of totem names
referring to insects have been preserved from many Australian
tribes, but that mainly through the devotion of SIR BALDWIN
SPENCER, who has written' an imposing array of books on the topic,
many of the rituals, the ceremonies and the traditions of a number
of these insect totems have been recorded and preserved. These
totem ceremonies are by far the best criterion for judging the role
of any food in the regular diet of primitive man. Only basically
important articles of food have been accorded the rank of totem
animals or plants. To illustrate the great role which insects played
in Australian totemism and hence in Australian nutrition, we have
had to devote some space to the ceremonies of the insect totems of
Australia, as well as to those of other continents, about which, un-
fortunately, far too little is known. The work of SPENCER has
definitely established the basic role of insects as food, mainly in
the arid interior of Central Australia, while in the fertile tropical
North of that continent honey-bags still form the main important
insect totems.
NOYES (1937, pp. 226 ff.) states that it is safe to assume that
throughout the ages primitive people have eaten termites in tropical
countries. This agrees with much current experience. He extends
this experience into the undocumented past: 'Though we have no
means of proving it, Pithecanthropus, shambling through the jungle,
must surely have drawn largely on the termitary for his food sup-
plies. It is only reasonable to suppose that since Pleistocene times

12
there has been no diminution of the practice on the part of those
who live, rawly, in the immediate neighbourhood of termites; apart
from what we can observe, all the evidence points to this conclu-
sion .... Whoever contrives to break open a termite hillock and
abandon it to the tender mercies of attentive monkeys or baboons
lurking in the neighborhood, may witness their interest. Chattering
at the prospect of a feast, snatching, when the time arrives at the
unfortunate workers and alates, careless of the soldiers' grip upon
their marauding paws, they cram the insects indiscriminately into
their greedy mouths. But, scrabble as they may, they never reach
the sanctuary of the queen, the chiefest delicacy of all. That
privilege is reserved for man and the ;:tnt-bear'. NOYES even suggests
that the idea of tools first occurred to man through the difficulty
his ancestors found in opening termite hills: when finding their
talons unequal to the task the utilized sticks to force an entrance,
from which the evolution of the lever would have been a simple
matter.
This latter theory is, however, too far fetched and probably in-
correct. Termite eating is rare in Australia, where the most primi-
tive man was studied and where termites abound in the drier areas.
Termite hills are only very rarely broken into and this in times of
famine. In tropical Africa, Asia and America, termites are eaten
in large quantities when the winged sexuals swarm out of the nests.
This restriction is physiologically well founded, as the swollen ab-
domen, especially of the females, is rich in fats and proteins, in
contrast to the much poorer body composition of the neutral castes.
These swarming sexuals are easily caught without breaking the
hard crust of the termite hills. And the thorough breaking deep into
these hills in quest of the large and much prized queens is doubtless
a very late development in search of delicacies or of medicine.
Yet, just as monkeys, together with scores of other mammals,
birds, lizards, and other animals are very fond of locusts, swarming
termites or ants, caterpillars and grubs, early man was doubtless
fond of all available insects with no repellent taste, especially if he
could gather them in quantities. We have no reason to assume that
Pithecanthropus differed in this respect from the Australian aborigines
or the inhabitants of primeval forests. There is no reason to doubt
that locusts were readily eaten by prehistoric man, whenever they
 .. .;. ."

<$-

Fib' 2. Total view of the Palaeolithic cave drawing at Arana of honey-hunting.

Above: The honey-hunter and the bees enlarged. From F. HERNANDEZ-PACHECO
(19 21 ).
were available in numbers. We may even suppose that he ate them
roasted, as a rule, as all his animal food was roasted, before he took
at a relatively late stage of his development to fishing. Fish were
apparently the first animals eaten raw apart from occasional insects
and molluscs. For honey-hunting we are by no means forced to
base our conclusions upon parallels from recent ethnology. A Pa-
laeolithic cave drawing from Arana in Spain gives a vivid picture
of a honey-hunting Palaeolithic man who climbs along a ladder
down a rock to collect the honey from a nest of wild bees (H.
PACHECO, 1921.) This picture is the more convincing as it is almost
identical with a recent sketch which SPITTEL (1924) has made of
a Yedda taking honey from a rock in the forests of Ceylon (see
fig. 33).

In view of the general importance of the discovery of honey-hunting

as portrayed in the Palaeolithic art first described by F. HERNANDEZ-
PACHECO (1921) in the caves of Arana (Spain), we give here a full
description of these drawings. Many figures of men and animals,
such as deer, wild goats and bulls, horses and a few extinct carni-
vores are drawn on the walls. Superimposed upon them are other,
clumsier figures from a later period. Some of these latter designs
illustrate scenes from the life of Palaeolithic man. They are brick-
red on the yellow-reddish rock.
From two heavy horizontal lines above, three fine parallel lines
descend over the length of the entire wall, here and there con-
nected by short horizontal lines. Close to the upper lines there is
a small natural cavity in the rock, over which the vertical lines pass.
At the level of this cavity there is the figure of a man, surrounded
by various small figures on the wall. Much farther below another
man is portrayed in contact with the vertical lines.
In the neighbouring ravines close to this site, bees are still common,
making their nests in crevices in the rocks. And even today, at the
approach of the cold weather of winter, the local peasants collect
the honey from any nests which are easy of access. This makes it a
simple matter to interpret the picture: two men climb up one of
the ropes which hang down from above, fixed to a pole, to collect
the honey from a bee's nest. The three ropes make possible the
ascent, forming a primitive ladder by means of the occasional hori-
zontal connections. The man above, close to the entrance of the
bee's nest, holds himself to the ladder with his legs and with one
of his arms. The other, outstretched arm holds a vessel, perhaps a
skin bag, with a distinct handle. The 16 small surrounding figures
represent flying bees, and on some of them the head, the abdomen,
the legs and the extended wings are recognizable. The man below
carries a similar vessel on his shoulder, thus keeping his arms and
legs free for the ascent. Both men are naked and apparently without
protection against the stings of the bees; but it is well known, writes
HERNANDEZ-PACHECO, that in winter the bees are lethargic. from
the cold and not so inclined to sting as on warm summer days.
In the neighbouring cave of Alpera there is a contemporaneous,
similar design of a climbing man, whom BREUIL (192 I) describes
as climbing in the manner of the Australians, i.e. without making
use of his knees. Espartograss still abounds in that region, offering
an ideal material for the construction of such ropes. This first Pa-
laeolithic insect illustration in a region where honey-hunting is still
practised shows the general similarity which must exist between the
local climate at that time and today.
In the Magdalenian grotte of Trois-Freres (Ariege) the graving
of a cave-grasshopper on animal bones was found, wich L. CHOPARD
could determine generically (BERLAND 1942 p. I 12). The highly
exact and artistic execution indicates the interest of the artist, and
quite possibly it was also connected with food magics.
To return to our cousins the monkeys, D. and R. KATZ (1930),
when studying the feeding habits of Cercopithecus sp. and of Cebus sp.,
agreed that they behaved similarly to a young human child. When
presented with seven different acceptable foods in small quantities
simultaneously, the same monkey would eat them nearly always in
the same order. The differences were small between the individuals
or even between the two species. But after eating a very large
quantity of grapes, their favourite food, for instance, this item
might be changed from the first to the last place in the next meal.
In general, whatever was eaten in quantity, was subsequently rated
lower or altogether rejected. 'Their feeding behaviour reminds us of
our own table manners. . . . All the rules controlling the feeding
behaviour of monkeys have their parallels in human beings'.
Most experiments on the insect-feeding habits of monkeys were

16
performed to discover whether aposematic insects with warning
colours or warning behaviour were rejected as food (COTT, 1942,
pp. 256, 271, 290). Extensive experiments in this direction were
made by G. H. D. CARPENTER (1921) in East Africa with Cercopi-
thecus sp. This monkey definitely recognized a difference in palata-
bility between various insects which it encountered when out
hunting with its master or which were offered to it. Insects in
general were readily and spontaneously eaten. Yet some species
were eagerly pounced upon, others neglected, according to its ex-
perience of their suitability as food. Various experiments carried
out by HEIKERTINGER and others have suggested fairly conclusively,
in particular with aposematic behaviour in insects which display
their warning colours conspicuously, that they are instinctively
refused by many monkeys. The possibility cannot, however, be ex-
cluded, though this is by no means certain or even probable, that
experience of unpleasant tastes in the past have influenced this
attitude.
A large and varied assortment of insects offered or spontaneously
found by the monkeys were classified by CARPENTER according to
conspicuousness and edibility, as indicated by the monkeys' reac-
tions. In 615 individual observations the first individual of 244
different insect species gave the following result: of 143 aposematic
species 120 were distasteful, 23 edible; of lOI cryptic species 83
were edible and 18 distasteful. CARPENTER concludes that warning
colours are, in general, a sign of distasteful qualities.
Various cryptic grasshoppers (Cyrtacanthacris spp., Catantops spp.)
formed the staple diet of Cercopithecus. Yet nothing would induce
the monkey to eat aposematic grasshoppers, such as Dictyophorus
productus, a heavy, bloated, sluggish insect which freely e.xposes its
grey and bright red abdomen to view. On one occasion, when one
of the monkeys saw from a distance that a grasshopper was being
brought to him, he became very excited. However, he lost his
excitement when getting a closer view of Dictyophorus. He picked it
up from the ground, smelt it, and put it down again. To encourage
him, CARPENTER pretended to taste it. The monkey then licked it,
only to get a taste of the yellow froth which it exuded. He then
shook his head, as if trying to get rid of a disagreeable taste and
would have nothing more to do with this grasshopper. Shortly after

17
this he seized with great eagerness and ate a huge, ten centimeter
long cryptic Cyrtacanthacris. Four days later, before he had eaten
anything, he was offered another Dictyophorus; this was examined
and licked, then dropped uninjured. When out hunting ten days
later, the monkey caught and ate another large Cyrtacanthacris, ab-
solutely ignoring a Dictyophorus, which was on the ground just in
front of him. Zonocerus elegans, another large, bright yellowish-green
grasshopper with short reddish elytra and with the antennae alten-
ately ringed black and orange, was similarly refused. When offered
one, the monkey just looked at it and took no more notice of it.
He was then shown another large cryptic grasshopper, upon which
he leapt at once, seizing it and eating it with extreme haste. The
large aposematic, green and red Phymeteus viridipes, when attacked,
erects its wings vertically to display their red and black colour. On
two occasions, when the monkey had begun to examine the grass-
hopper, this display prevented any further interference; he never
would eat this species. An unidentified aposematic grasshopper was
avoided by a Slow Lori in Burma (MACKENZIE, 1930).
GUY MARSHALL (1902) confirms this experience for baboons with
the big aposematic caterpillar of the hawk moth Chaerocampa osiris:
'The female baboon ran forward expecting a tit-bit, but when she
saw what I had brought she flicked it out of my hand on to the
ground, at the same time jumping back suspiciously; she then
approched it very cautiously, and after peering carefully at it from
the distance of about a foot, she withdraw in alarm, being clearly
much impressed by the large blue eye-like markings. The male
baboon, which had a much more nervous temperament, had mean-
while remained at a distance surveying the proceedings, so I picked
up the caterpillar and brought it towards him, but he would not
let me approach, and kept running away, until I threw the insect
at him. His fright was ludicrous to see; with loud cries he jumped
aside and clambered up a pole as fast as he could, into his box ....
On concealing the larva I managed to coax him down again, drew
him slowly towards me holding up the larva in the other hand;
he simply screamed in abject terror'.
The observations of W. W. A. PHILIPS, 1931, (also POULTON,
1932), carried out in-Ceylon on a lemur (Loris tardigradus) in capt-
ivity, which was offered a variety of butterflies and moths, are

18
likewise in agreement with the rule that cryptic insects were accept-
ed, aposematic ones rejected. The importance of these and other
experiments for us lies in the unavoidable conclusion that not only
the lemurs, but also the larger monkeys and apes readily eat insects
in greater or lesser quantities, either spontaneously or if offered
them. This removes the last possible doubt that primitive man,
from Pithecanthropus to the recent food-gatherers, was in no way
prevented by instinct from consuming insects. On the contrary, at least
the occasional eating ofinsects belongs to his phylogenetical tradition.
The fact that many primates readily eat the most varied insects
spontaneously can no longer be doubted. Some of the tiny lemurs,
such as the Malayan Spectral Lemur, even exist primarily on
insects. Furthermore, during a study of the influence of aposematic
insects on feeding habits of various monkeys, it has been incidentally
proved that the higher monkeys can make them their staple food.
Therefore, we cannot agree with BRUES (1946, p. 4 I 8 f.), who is
sceptical about the food value of 'these tiny creatures,' which make
a major contribution towards satisfying the Gargantuan require-
ments of such complex social organisms as men only on rare occa-
sions. He agrees, however, that certain insects on occasions form a
real source of food, of which he quotes various instances from the
life of the American Indians. Yet, considering the use of insects as
tit-bits, hors d'oeuvres or medicines, etc., the wealth of information
is so overwhelming, ranging from termite queens to minute stink-
bugs, that it could only be catalogued by a librarian. We will,
nevertheless, try in the following pages to demonstrate that at
certain seasons, under certain sociological and ecological conditions,
insects have formed and remain an important part of the all-the-
year-round food of primitive man; they do not represent a mere
dainty, but a physiological necessity by virtue of their qualitative
as well as quantitative importance.
A. MAUR1ZIO (1932, p. I) in his 'History of Vegetable Nutrition'
defines the gathering of various animal and vegetable foods and
primitive hunting as the lowest stage of human nutrition. Charac-
teristic of this primitive stage is the great and ceaseless effort which
the search for food requires. No part of a utilizable plant is wasted
by the collectors and the same is true for all available animals, from
the highest to the lowest: nothing is refused.
 MAURIZIO mentions the excellent humour into which an African
native is transported when gathering a handful of lice from the
head of one of his neighbours. He also quotes the dramatic descript-
ion of v. WISSMANN (I 890, p. 168) who saw dense swarms of mil-
liards of tiny gnats over Lake Tanganyika forming living clouds
known as kungu by the natives. The natives follow these swarms as
soon as they appear above the ground and collect them while the
gnats are resting after having crossed the lake. A flour is made from
them, which is used for roasted cakes, a highly appreciated food.
The poorest and most primitive peoples, like the Australians of the
deserts or the pygmies of the primeval forests, gather everything
edible in their environment. Unfortunately, we are less well in-
formed on the animal food of prehistoric man, apart from big game,
than on his vegetable food. The meat of big game has been appa-
rently roasted or grilled from the time of the first available records.
Fish was, therefore, the first substantial animal food to be eaten
raw. We have little information about insects and shell-fish, yet it
may be safely assumed that in many cases early man ate them raw.
G. RENARD (1931) also holds that the Australian aborigines well
illustrate primitive conditions. In order to illustrate the primitive
methods and at the same time the ingenuity of their search for food,
he mentions that the Australians sponge honey from flowers for
eating purposes, when no wild honey from bees is available. He
likewise stresses the need to work hard throughout the year for their
daily food supply. Nothing is known about primitive man making
stores of food.

2. SURVEY OF ENTOMOPHAGOUS HABITS ON

THE VARIOUS CONTINENTS.

The most primitive people who have been studied from the point
of view of entomophagy are the Australian aborigines and they will
be discussed in detail later. Here a few general quotations may be
in place, in order to illustrate the role of insects in the diet of these
primitive food-gatherers.
Keith C. McKEOWN (1944, p. 170) says: 'Nothing in the way of
additions to the food supply ever came amiss to the native. Through-

20
out the continent the insects were exploited and played a prominent
part in the aboriginal commissariat, varying often only with the
presence or absence of any of the insects in particular districts'.
Or J. MATTHEW (1910, p. 88): 'In the territory of the Kabi and
Wakka of Queensland, food was plentiful and in great variety. The
animal food embraced almost every living thing from a fly to a man'.
B. SPENCER (1922, pp. 66 ff.) who is, perhaps, the most competent
judge, states in his 'Guide to the Australian Ethnological Collection
at Melbourne' that among the higher vertebrates practically every
mammal, bird, reptile, frog and fish that has enough flesh to 'make
it worth eating, serves as an article of food in some part of the
continent or another. Among invertebrate animals, shellfish of
various forms, mussels, cockles, etc. are eaten in numbers, their
empty shells lying in heaps beside the cooking places, forming on many
parts of the sea coast shell mounds of great size. Various forms of
insects, such as Bugong-moths and the larvae of other moths and
beetles, as well as ants, are much relished and wherever obtainable,
the honey-comb of wild bess is a favourite item of diet. Mupingalu,
pounded up termite hill, is eaten as a cure for colds by the natives
of the Kakadu tribe.
We conclude these few general quotations with two more:
EYLMANN (1908, p. 278 f.) is the only observer to state that the
natives of South Australia eat only few insects. He explains this by
the poverty of the local insect fauna in edible species. We will see,
however, that his own lists of insects are by no means small. The
absence of observations on locusts is nevertheless remarkable.
Recent research seems to indicate that in South Australia locust
plagues are largely a more recent consequence of the changes in
vegetation induced by human colonization. The South Australian
diet is mainly of animal content. Vegetable food is unimportant,
as it is either slight in quantity and rare, or unhealthy. During the
common drought periods vegetable food is especially scarce and fat
game is wanting. The aborigines of the interior then live on a diet
rich in proteins but poor in fats and carbohydrates. During such
periods, they lose in body weight, in consequence of the small
quantity and the unbalanced composition of their food. Generally
their diet lacks non-proteins, as is demonstrated by their perpetual
eagerness to search for fats and carbohydrates. With regard to his

21
 animal food, the native is by no means selective. He eats all verte-
brates and the larger molluscs. Yet many insects are apparently
not to his taste. He even refuses locusts and grasshoppers.
R. SEMON (I903, p. 233), who has made no special notes on insect
food of the Australian natives, makes the following observations on
their diet: 'The great leanness of the native Australians is mainly
based upon their prevalently animal food: Marsupials and Echidna,
birds, snakes and lizards, turtles fish, beetle-grubs, eggs of birds and
reptiles, shrimps and shellfish. Some of the Queensland tribes are not
above eating human flesh. Hunting for game is the task of the men,
whilst the women dig in the scrub for edible roots, collect mush-
rooms and nuts of palms, pods of leguminous plants, seeds of
grasses, sweet gum and manna of eucalyptus. Yet the flora of
Australia is poor in edible fruits and mealy roots, which makes
their diet poor in carbohydrates'.
Africa is perhaps today the continent where insects still play the
most important role in native diet. M. BR1AULT (1943, pp. 82 ff.) has
given us a remarkable review of the diet of the African Negroes:
'Before penetration by the Arabs and white men, primitive Africa
was a continent almost bare of basic foods, especially those of
vegetable origin. The only native vegetable foods were bananas,
citronella, gourd, beans and peas, sorghum, millet, durrha and
perhaps taro. The natives of the primeval forest, including the
Negrillos, have no plantations at all and live exclusively on game
and fishing .... Meat, fish, pastes of caterpillars or big palm worms
are wrapped into a large leaf, which is made supple by passing it
over a fire. Some salt, some spice, some drops of wild lemon, and
a pleasant meal is taken from the hot ashes, yet it never lasts long ....
When game is insufficient or lacking, the worries of supplying the
camp with provisions pass to the women. They go out to explore
the rivulets, the caves, the ponds, the shrub, the bog, and return
with rats, lizards, snakes, occasionally even with a small monkey
or porcupine, or with a collection of caterpillars, palmworms, tad-
poles or young fish. These dishes of lean days are stewed with salt,
spice, and lemon, which are added to every native dish. Neverthe-
less, famines are not rare'.
BEQ,UAERT (I92 I, p. 193) ascribes the wide use of insects as food
in Africa more to necessity than to choice. He maintains that the

22
 climate and the small scale of animal husbandry reduce the amount
of meat eaten, even that of chicken and dogs; hence the perpetual
craving for animal food, which is one of the causes of cannibalism.
Crops often fail. Thus, repeated famines are a contributive factor
towards the inclusion of insects in the regular diet. The reader will
have observed that, while BRIAULT'S description refers to primary
food-gatherers and primitive hunters, the remarks of BEQ.UAERT
refer to agriculturists. But even in this case the conclusion that
entomophagy is primarily induced by famines is inaccurate. The
wide use made in the agricultural societies of tropical Africa of
termites, locusts, and honey contradicts this conclusion. BRYGOO
(1946, p. 27), for example, has pointed out, quite properly, that,
while it would be exaggerated to talk in Africa about tetmite civiliza-
tions, the primary importance of these insects as part of the diet is
very great. The origin of many ceremonies can be traced back to
them and they may even eventually determine a certain rhythm
of life, even among primitive agriculturists.
In tropical Asia all sociological and economic stages of develop-
ment, from primitive food-gatherers to highly specialized agricul-
turists, are still co-existent. Corresponding to this variety of social
conditions one finds a variety of insect feeding habits. Among the
Veddas of the Ceylonese primeval forests strong remjnders and
remnants of an almost pure 'honey-civilization' from a not very
remote past are still found. Another group of Asiatic pygmies in-
habiting the Andaman Islands, who are also primitive food-gathe-
rers, depend so much upon insect food that two of their months are
named: 'the month when caterpillars abound' and 'the month of
honey abundance'. The writer greatly regrets having lost the re-
ference to this most important statement. The level of other food-
gatherers, such as the J akuns and similar groups in the forests of
Malaya, is not much higher (FAVRE, 1865, MARTIN, 1905, etc.).
R. KIPLING tells us in the first chapter of his 'Second Jungle
Book': '110wgli, who had never known what hunger meant, fell
back on stale .poney, three years old, scraped out of deserted rock-
hives. He hunted too for deep-boring grubs under the bark of the
trees, and robbed the wasps of their new brood'. These lines are
certainly taken from tales heard and from observations made by
the wt"iter in India.

23
 In the highly organized agriculture of the monsoon regions, such
asJava, Malaya, Siam, etc., the ill-balanced, monotonous vegetable
diet induces a craving for animal proteins and fats. This is easily
satisfied, both by small game and fishing, and also over wide areas
by gathering insects which are roasted or included in sauces, thus
forming an important supplement to the daily rice. BURR (1939,
p. 209) quite correctly states that the teeming millions of India,
China, Japan and Malaya could hardly imagine life without their
daily meal of rice. The agricultural Negro tribes of Africa feed
almost entirely upon mealies, manioc, bananas or Kaffir corn. Such
people feel an overwhelming hunger for meat and do not even scorn
rats and grubs. The report of VAN DER BURG (1904) for Dutch In-
donesia, ofBR1STOWE (1932) for Siam, ofN GUYEN-CONG-TIEU (1928)
for Indo-China and many others, which will be discussed in detail
later, also support this thesis.
The extremely frugal peasants and urban proletariat of China
and Japan also improve their daily rice meals by the addition of
small quantities of any kind of animal, from toads and mice to
insects. Silkworm pupae which remain ready cooked after the
reeling of the silk from the cocoons are highly appreciated as food.
Grasshoppers are eaten in Japan. In China they form an important
food during the famines, for which they are partly responsible. They
are always a part of the food of the coolies and in Peking they are
offered boiled in salty water in the restaurants. In Burma fried
locusts are a delicacy. Many other insects are used to flavour
the rice.
Locusts and wild honey are still appreciated in the Middle East
as items of food for nomads and mountain peasants, who also
delight in partaking of the various forms of manna.
FROST (1942, p. 63 f.) remarks that in North America the primi-
tive idea of economy suggested the use of insects as food. When
insects emerge in great numbers, such as cicadas from the ground
or mayflies from water, they may be gathered with little effort. He
even suggests that insects may supply salt when this is deficient in
the diet. The scarcity offood might at times lead certain populations
to select insects as food. In the Eastern United States insects were
not frequently eaten, chiefly because the rainfall was generally
sufficient to produce good crops and insect food was not essential.
In the West, where famines were more common, the Indians
resorted to any kind of available food. Again, here also famine is
not the only and not the most potent of the factors inducing ento-
mophagy. The arid plains of the Western areas did not favour
agricultural development and big game was fairly rare. Thus, the
entire level of economic development on the two sides of the Alle-
ghany Mountains was very different, resulting from the general
ecological conditions. It will be shown below that the Indians of
the Eastern states were by no means averse to eating insects.
Reports from the early days of occupation testify to widespread
eating of insects in the West Indies. PETER MARTYR (1612, p. 121 f.,
vide F. OVIEDUS) says that in the houses of the inhabitants they
found great chests and baskets made of twigs and leaves, which
were full ofgrasshoppers,· crickets, crabs, crayfish and snails, together
with locusts which destroy the fields of corn, all dried and salted.
The Indians explained that they kept these insects to sell them to
their inland neighbours. And in the account of J. H. LINSCHOTEN
(vide WANLEY, 1806, II, p. 373) we read that the inhabitants of
Cumana eat 'horse-leeches, bats, grasshoppers, spiders, bees and
lice, roasted or raw. They spare no living creature whatsoever, but
they eat it'.
DE WAVRIN (1937, pp. 58, 122 ff) concludes that the South
American Indians are, in general, far from strict as vegetarians.
Many tribes are exclusively flesh- or fish-eaters, others are omni-
vorous. All like bees'-honey very much and honey-bees abound in
certain parts of the forests. It is a general rule that all tribes eat
almost every mammal, from the largest to the smallest. The Piaroa
eat many fruits and vegetables, every fish and every 'reptile' from
toads to poisonous spiders. Ants, caterpillars, palmworms, etc., are
accepted as food and from the primeval forests of Paraguay VEL-
LARD (1939) has described a real 'honey-civilization'.

3. THE NUTRITIVE VAL UE OF INSECTS AS

HUMAN FOOD.

The most recent work on the food value of insects has been
carried out by French scientists, especially in West Africa. G.
 HARDY and CH. RICHET (1933), in a book devoted to the native
diet in the French colonies, come to the conclusion that most
natives there are undernourished: in wide areas chronically, in
others seasonally. The best measure of the soundness of a people's
diet is its mortality (p. 41). Termites and 'earthworms' are quoted
from the Lower Ivory Coast as sources of proteins (p. 163). All
forest peoples are in need especially of proteins, as game in this
habitat is very difficult to come by. Even if some people are appa-
rently overfed, such as some tribes of Togo according to a survey
of A. CHEYSSIAL, where the average daily diet contains 4000 to 8000
calories, it may be deficient in proteins or in fat (p. 289).
M. SARRE, Professor of Geography at the Sorbonne, in his 'Bio-
logical Basis of Human Geography' (1943), devotes a special chapter
to the geography of dietetic regimes (pp. 247-290). 'The regime
of a group is the combination of foods, either produced locally or
imported, which assure its daily existence, satisfies its taste and
assures its preservation in a given complex of life conditions. This
is a norm. The medical view of the dietetic regime has an individual
character'. The diet is composed of basic foods plus complementary
and luxury foods. Primitive dietetic regimes, such as those of the
aborigines of Australia, show an extraordinary diversity. They pass
their life in a perpetual search for food. Everything edible is accept-
able to them: emu-eggs, crocodiles, tortoises, ant-maggots, cater-
pillars and other insect-larvae, grasshoppers, moths and snails. The
flora yields them water-lily bulbs, a wide variety of grains to make
soups, roots and buds of Xantorrhoea, flowers of Banksia, in all not
less than 300 species of edible plants. Among animals, kangaroo,
opossum, tortoises, frogs, rodents, snakes, many birds including the
emu, freshwater crabs, etc. are eaten when available. Close to the
sea-shores whales, seals, sea fish, molluscs, etc. are added. The
preparation of the food is governed by complicated rites (cf:
WARNER, 1937). VELLARD'S description (1939) of the Guayakis in
Paraguay is quoted, as well as the similar food-habits of the Veddas
in Ceylon. The absence of a permanent domicile and of stores make
necessary a daily search for food, and often result in hunger com-
bined with occasional overfeeding. During times of famine human
flesh is a last resource. Many regimes are one sided, as often in
hunting or fishing peoples, in the animal-breeding nomads or where
one vegetable is the dominant food. In Africa and in the Oriental
monsoon regions mixed diets with a predominant vegetarian ten-
dency are often also one-sided. SARRE goes into the causes and
effects of undernourishment and of famines (pp. 277 ff.) and into
the effects of ill-balanced diets, which may produce diseases such
as beri-beri, pellagra, scurvy, etc.
The best survey on the problems of native nutrition in tropical
regions has been given by P. GOUROU (1947, pp. 76 ff.). Nutrition
is .mainly vegetarian in an environment little favourable for animal
breeding. And even many pastoral people have an essentially vege-
tarian diet. Thus, the Betsileo of Madagascar, despite their herds
of cattle, live on rice, manioc, sweet potatoes, maize, spinach and
fruits. Meat is an exceptional dish and milk is not consumed; cock-
chafer grubs, caterpillars, locusts and small.fish are the main animal food.
Even among the Foulah, the Hindus of India and other people who
eat animal products, we find that the diet is deficient in animal
proteins, fat, calcium and vitamins. In general, animal products
form only 4 to 5 % of the total calories. The ingenuity of the natives
of the warm and humid zones in utilizing the resources of the vege-
table kingdom is great. The large-scale reduction in the number of
nutritive plants so characteristic of modern Western civilization
has not yet begun. GOUROU mentions from the plateau of the Gold
Coast 114 kinds of fruit, 46 kinds of leguminous seeds, 47 kinds of
vegetables. The essential food is always a porridge of carbohydrates
(cereals or roots), seasoned by essentially vegetarian sauces, oil,
fat or spices. Some of the vegetables, such as bamboo (Elaeis-palm),
are prepared in the most manifold manner; in Northern Ceylon
Borassus flabellifer is prepared in eight hundred and one different ways.
And on the coasts of Guinea, when the preparation of Elaeis has
been completed, the palmworms - the grubs of Rhynchophorus ferru-
gineus - are also extracted. The consumption of a daily rate of 1700
calories in tropical countries is nothing exceptional, a rather in-
sufficient quantity! Inadequate nutrition is especially marked at
the end of the agricultural year, when reserves are exhausted.
Then food gathering in the savanna and in the forest commences:
wild gramineous seeds, wild tubers and fruits, mushrooms and
caterpillars are collected. Women searching for these foods are a
common sight in the African scrub. And by a coincidence this

27
season of want usually occurs at the time of the heaviest agricultural
work. Thus, it is not surprising that the undernourished natives of
the tropics concentrate their thoughts on food. The discovery of
a nest of caterpillars is long remembered.
This situation is caused by the poverty of the soils, the irregularity
of the rains and locust invasions. Regions with rains throughout
the year are better off than those with seasonal rains. Malaria and
similar diseases sap the natives' energy and reduce their capacity
for work. In spite of the greater stability of root diets over those of
cereals, the latter are everywhere preferred by habit, tradition and
taste. GOUROU warns against accusing these people too hastily of
improvidence, since the poor soil would be exhausted by more
intensive cultivation and give reduced crops in the future.
Yet, for us the most important thing is that these diets are not
only inadequate in quantity, but deficient as a rule in animal
proteins, fat, vitamins and often in minerals, and also in the so-
called protective foods which help to preserve health. Thus, lack
of calcium and of vitamin E is held to cause the low fertility of the
Haussa women (WORTHINGTON, 1938, p. 572). Taboos often prevent
the use of milk and meat. Tropical diets are usually poor in mine-
rals, such as phosphorus, calcium, iron, iodine and sodium. Lack
of salt is often seriously felt. All these deficiencies are apparently at
the base of the many cases of geophagy (earth-eating). And again
this shortage of protective foods is accentuated towards the end of-
the dry season and the beginning of the rainy season, the period of
the heavy agricultural work.
This brief survey is sufficient to illustrate the great deficiency of
tropical diets in animal components. Here is the main cause and
the main importance of the widespread, large-scale and habitual
use of insects of all kinds by all tropical native populations. Hence,
this diversion clearly shows that insects as food in these regions are
by no means dainties, but offer to primitive man just those elements
in which his basic food is deficient. This will now be easily under-
stood from the few adequate analyses of nutritive insects which we
have at our disposal.
The first report on Nutrition in the British Colonial Empire (193-9:
I, p. 67) comments on the general character of colonial diets: 'The
more normal sources of food supply are supplemented in many
cases by wild animals and insects. We have already mentioned the
consumption of such things as grubs, caterpillars, locusts and flying
ants (read: termites). Many kinds of wild animals are consumed,
though not usually with great frequency. In the more closely popu-
lated countries they are of some significance'. The second part of
the same report (1939, pp. 19, 25, 29) mentions locusts, grasshoppers
and white ants as universally eaten as delicacies. In Northern Rho-
desia grubs, woodlice, caterpillars, flying (white) ants, honey and
dried fish are widely used as local foods. In Nyassaland, although
meat is relatively cheap, its price places it out of reach of the
average villager, who depends on game, small rodents, caterpillars,
flying (white) ants, locusts, etc., augmented by fish, for his intake
of first-class proteins.
Perhaps the most elaborate chemical analysis of termites has been
made by L. TIHON (1946). TIHoN was stimulated by his often
repeated observations of the arrangements made by natives of the
Belgian Congo for catching termites for immediate consumption and
for trade, dried in the sun or slightly fried. The propriety claims of
families or villages on specified termite hills underline the great
importance of this insect crop. In May one usually finds at the
Leopoldsville market baskets full of fried termites, which are sold
for 54 centimes for a small handful. Most natives eagerly devour
the donge-termites, from which they also gain a colorless oil of good
quality, excellent for frying. It was with such slightly fried, brownish-
looking, oily and aromatic-smelling termites from the market at
Leopoldsville that the analyses were made.
Moisture . 6.0% Ashes . . . . . . . . . . 6·4%
Dry matter . . . . . . . 94.0% Fat (petroleum, ether 50%) 44·4%
Nitrogen . . . . . 5. 8 %
Protein (N X 6.25) . . . . 3 6 .0 %
Chitin. . . . . . . . . . 5. 1 %
100 gr. offried termites have a value of 561 calories, which ranges
them among the richest foods; they are superior to other animal
foods and approach the value of groundnuts. These termites are
most important as a source of proteins which are too often deficient
in the usual native diet. Does not this need of protein provide the
main and instinctive reason for the eating of termites, caterpillars,
adult beetles and their larvae? The analysis of the ashes yielded:
Insoluble 60.6% NaO 2.2%
I ron and aluminium oxides 11.8% P 2 O. 7.0 %
Manganese traces
CaO 1.4% Chlorides 8.2%
MgO 0·3% Sulphates. traces
K 20 7.8 %

The insoluble component is mostly sand. The termites are rich in

phosphates and potash, poor in sulphates.
Termite oil extracted by petroleum ether below 50° C. from
lightly fried insects from the same source was brown and of a
typical aroma. It contained:

Density at 28° C 0.906 Contents of brown, solid fat:

Index of saponification 19 1
Index of acidity 18.6 Titer 38c C
Acidity in oleic acid 9·3 Melting point: beginning 40° C
Index of Hehner. 94. 1 : end 41° C
Index of iodine Hubl . " of solidification
Point " 38.5-39° C
55·4
Index of refraction at 40° C 1.461 Index of iodine 57·9
Soluble acids. Planchon in Index of saponification 198
butyric acid. 0.64 Average molecular weight. 283.5
Insaponifiable . 3.7 6 of the corresponding acids

This non-siccative oil has a somewhat high acidity. It yields a

clear, fairly hard soap. Even though this oil may never be of com-
merCial interest, its study has some theoretical value. The termites
themselves form an important part of the diet of the natives in
most districts of the Belgian Congo and their study is of definite
practical interest.
CH. AUFFRET and P. T ANGUY (1947/48), of the French Colonial
Medical Service, have fully confirmed these analyses of TIHON.
They observed in French West Africa that the Soussou soak the
termites in cold water, dry them in the sun and put them into an
iron pot or a jar. The Fullah boil the termites first, before drying
them in the sun, and later repeat the process. Then the insects are
fried. They are eaten as they are or they may be incorporated into a
sauce. Dr. PALES brought back from his journeys samples of fried
termites from Pita in the Fullah country and living termites cap-
tured while swarming near Dakar. Their analysis permitted a
comparison with other common foods of the same region:
 Moisture Fat Proteins Ashes Calories of 100 gr.
Living termites 44·5% 28·3% 23. 2% 347
Fried termites 13. 0 % 36. 2 % 45. 6 % 508
Meat (beef) 75.2% 6.6% 16.9% 1.3% 127
Dried salt-fish 32.4 % 3.1% 43.7% 20.8% 203
Groundnutoil (Senegal) 7.0% 49.4% 27.6% 2·7% 598
No vitamin A was found. The samples were too small to permit
an analysis of the ashes. The calorific value of the termites as com-
pared with that of the other foods is surprisingly high. In addition
the termites offer for certain peoples a most important source of
protein, of fat, and of calories. The writers conclude that their
nutritional value can explain an apparently paradoxical observa-
tion, namely, the healthy condition of some natives compared to
their poor diet, if unknowingly certain dishes, such as the ter-
mites, are omitted from the analysis.
ANTONIE has analysed smoked caterpillars from the Upper Sangha,
which contain 15.7 % moisture, 40. % ashes, 13.7 % fat, total
N 6. I %, carbohydrates 13.9 %, phosphates 0.7%, chlorides 0.4 %,
chitinous remainders 13.5%. A second sample yielded a slightly
lower nutritive value. ANTONINI calculates the calorific value as
268 cal., of which 258 cal. are digestible.
An analysis of silkworm pupae by CH. AUFFRET yielded: 60.7%
moisture, 23. I % protein, 14.2 % fat, minerals 1.5 %, calorific value
207 cal. Vitamin A was fairly ample. Both these analyses have been
made available through the habitual kindness of Prof. TH. MONOD,
Paris.
Y. YONEZAWA and K. YAMAFUJI (1935) give the following ana-
lysis of the eight days old pupa of the silkworm (Bombyx mori L.)
(female), in gr. per 1000 individuals:
Water: 1191 gr; dry matter: 362 gr.
Protein 207.5; fats gO.5; carbohydrates 23.2; ashes IS.S; total
phosphoric acid 7.S; organic phosphoric acid 4.3; total nitrogen
45.0; non-proteinous nitrogen 12.2 gr.
The amino acids of the pupa (E. KATAYAMA 1917) are alanin,
valin, leucin, prolin, phenylalanin, glutamin acid, histidin and
tyrosin in 0.05 to 0.67 % of the live weight.
The properties of the fats of the pupae are (YONEZAWA and
YAMAFUJI 1935): acid indicator 14, ester indicator IS2, Hehner
 indicator 95, saponification indicator 196, iodine indicator 137.
The glyzerides of the oil of the pupae (YAMAFUJI 1933) are in %
of the oil: dioleo-linolenin 0.6, insolin-oleo-dilinolenin 4.3, triolein
44.7, palmito-diolein 8.5, palmito-oleo-linolenin 16.7, tri-lino-
lenin 8.3.
No data are available for the minerals of the pupae but those of the
larvae are in % of the total ashes (LENZ 1867): P 20 5 28.7; K 20
48.1; Si0 2 0.6: CaO 5.9; Fe 20 a 0.7; MgO 8·4; SOa 6.2; Na 2 1.3.
All these data are quoted after K. YAMAFUJI (1937).
Some analyses of a few locust species are contained in the fol-
lowing table. In addition Dr. J. K. GUGGENHEIM informs me that
in the specimens of Schistocerca, which he analysed, 100 grams con-
BODY COMPOSITION OF VARIOUS LOCUSTS
Nomadacris septem- Schistocerca Locusta migratoroides
Species fasciata gregaria
Author Hemsted 1947 Das Gug- Imp. Inst. Anim. Nutr. 1936
1945 gen-
heim
Condition DM. DD. W. SD. DM. W. SD. SD. SD F.
% % % % % % % % % %
Water 8.0 70.6 5.0 10·5 7. 1
Protein 63·5 5 8 .4 18·7 61.4 75. 0 46. 1 61.1 55.0 49·7 47·5
Fat 14. 1 13.0 4. 1 17.0 3·4 9.6 10.1 15.0 18·4 22·9
Carbohydrates 7·5 6·4 9.6 6.8
Fibre 13·5 12·4 4.0 10.0 12·5 4·9 4·9
Total ash 8·7 2.6 5.0 4.6
CaO 0.1 0.1 0.04 0.6 0-4
P 20 0.2 0.2 0.1 1.2 1.6
Cl 1.2
K 20 0.8

DM.: Dry matter; SD.: Sun dried; W.: wet or fresh weight; F.: locust flour.

tained 1.75 mg. riboflavin and 7.5 mg. nicotinic acid. This shows
that even the vitamin content of insects may not be negligible and
the same is true of certain minerals such as phosphates.
B. P. UVAROV (1948) has carefully compiled the available data on
the chemical composition of grasshoppers. The complete mineral
ash content of sexually mature adults of Schistocerca gregaria (E. B.
UVAROV 1931) and of the two sexes of Oxya velox (ICHIKAWA
1936/37) IS:
32
 Schistocerca Oxya
males females
SiO. 11.9% 6.64% 9. 61 %
CuO 0.16
Fe.O. 2.06 1.59 0.69
MnO 0.21 0.26 0.3 1
Na.O 6.2 17.20 20.21
K.O 18.2 21.78 19.23
CaO 6.2 1.05 0·74
MgO 4·9 5·44 3. 8 5
TiO. 0.16
NiO 0.009
P.O. 3 2.4 40. 10 46.26
SO. 2.56 0.67 0.27
Cl 0.40
C 2·4
The desert locust is richer in calcium, sulphur and iron content
but poorer in sodium and phosphorus than Oxya. Other students
found also lithium, barium and strontium in locusts which had been
boiled in salt water and which were being sold on a Moroccan
market. The organic constituents ofOxya spp. (KORIGAWA 1934) were
moisture 11.78, total N 10.85, crude protein 67.79, protein N 7.45,
protein 46.61, amino N 0.78, crude fats 4.52, crude ash 3.78, 'chitin'
9.25, glycogen 0.01. The high protein content of 46% is ·confirmed
by other analyses: in Nomadacris septemfasciata: 46.1 protein and
9.6 fats; in Schistocerca gregaria: protein 56.7 in the males, 42.3 in
the females (LAPP and ROHMER 1937). The protein analysis of
Oxya velox (ICHIKAWA 1938) yielded for the females: water soluble
N 36.49, albumino N 6.68, globulin N 8.51, protamine N 1.51,
gluten N 2 I .84, chitin N 25.00.
TIMON-DAVID (1930) summarized the data on fats and oils of
insects. He found 13% of oil in the larva of Ergatesfaber L., 22.3%
in that of Rhynchophorus palmarum L. For grasshoppers he gives the
following data:
Species % of wet iodine saponifica- reference
body weight index tion index
Dociostaurus maroccanus 3·3 109.9 181.0 Timon David 1930
Anacridium aegyptium 2.6 97. 6 198.9 ditto.
Schistocerca Smale 106·7 224.0 Lapp Rohmer 1937
gregaria ( female 92 .7 224.0 ditto.
Schistocerca paranenis 65. 2 194·5 Tewithick Lewis 1939

33
Oxyajaponica 3. 0 122.6 171.5 Tsujimoto 1929
Taeniopoda, male 2·5 111.5 196.5 F. Giral 1941
auricornis (female 2.8 101.3 185. 0 ditto.
Melanoplus mexicanus 75·3 216·3 J. F. and M.Giral 1943
The role of fats and their bio-chemical changes within the indi-
vidual's development has not yet been studied. The ranges of the
indices given above may equally well refer to changes due to matur-
ation as to differences between species. MATTHE (1945) obtained
a considerably higher fat content in the gregarious than in the soli-
tary phase of Locusta migratoria and Locustana pardalina. For further
information it may be added that chitin, or rather the indigestible
exoskeleton, froms 4 % of the wet, and 10% of the dry weight of
adult locusts. This means that the indigestible part is not large
enough to diminish to any considerable extent the protein value
of the insects as food. Bl and B2 are the vitamins most commonly
found in grasshoppers.
All these chemical analyses may mean very little to the layman,
yet they confirm that precisely those insects which are eaten in large
quantities are rich in animal proteins, in animal fats and in calories.
They include a fair number of mineral salts, which may under
certain circumstances be valuable for human nutrition. The im-
portance of insects as a source of vitamins has apparently been
overestimated by some authors. The richness of insects in animal
proteins and fats and the richness of certain animal products
(honey, honey-dew) in carbohydrates suffices to explain their inter-
national importance in the poor and ill-balanced tropical diets.
And although vegetables in the tropics are especially poor in vita-
mins of the B-group, it may well be that this relatively small vitamin
content has an even greater importance than its small size might
indicate.
Honey is not only one of the few concentrated sweets to which
primitive man had access but, with its almost pure solution of
monosaccharid sugars, represents one of the most stimulating foods.
Physiology has well established that carbohydrates are the most
perfect food for the production of working energy in man and
domestic animals. The nectar in the crop of the honey-bee worker,
as well as in that of most other strongly-flying insects, is the main
source and reserve of the energy expended during flight. The ex-

34
haustion of this store will speedly immobilize the insects or in the
bee-worker bring it to the border of starvation. There is perhaps
no other food, excluding stimulating drugs, comparable to honey
for the prevention of fatigue or for the restoration of strength after
thorough physical exhaustion. The warriors of the Masai and of
the Bushmen relied almost exclusively on honey as food during their
longer expeditions and wars. Many explorers in Africa and Australia
have praised the invigorating qualities of a mouthful of wild honey
This was also well known in Biblical times. In I. Samuel 14 : 27
we read that Jonathan dipped the rod in a honey-comb and put
his hand to his mouth; and his tryes were enlightened. It is almost im-
possible to describe more vividly the immediate restorative effect
of honey, when one is fatigued. Possibly the war with the Philistines
would have ended differently if Saul and his army had all parti-
cipated in the meal of honey instead of fasting. Athletes from the
age of Greek glory down to the mountaineers of our days, pilots and
deep-sea divers, they all by tradition and by belief eat large quantities
of honey during their periods of training, during their actual efforts
and immediately afterwards. Physicians, dieticians and bee-keepets
have always strongly supported this claim of honey as a restorative
of energy and as a means of preventing fatigue (for example, see
BECK and SMEDLEY 1947). The following table gives a condensed
compilation of the main facts known about the composition of
bees'-honey.
Bees'-honey has been thoroughly analysed, especially by the Agri-
cultural Department of the United States of America. Samples
from all over the world were included in these analyses, in order
to establish the standard requirements of commercial honey. The
results are (BECK and SMEDLEY 1947; ECKERT 1936):
I. Main components: Water about 17.7%
Dextrose 34.0 (24-37) Dextrose and levu-
lose are variable,
Levulose 40.5 (39-49) but their combined
total is fairly stable.
Sucrose 1.9 The total of the 3
basic sugars: 76.4%.
Dextrins and gums 1.5
Total:

35
 II. Ash contents: The minerals of the ashes amount to about 0.18%: calcium
about 5 mg., phosphorus 15-17 mg., iron 0.4-1.0 mg.
Traces of the following are also present: silica, copper,
manganese, chlorine, potassium, sodium, sulphur, alumi-
nium and magnesium.
III. Vitamin contents: Vitamin A o
Vitamin C 1-6 mg.
Thiamine (B1) 0.0021-0.0091 mg.
Riboflavine (B.) 0.035-0.145 mg.
Pyridazine (B6) 0.210-0.440 mg.
Pantothenic acid 0.047-0.192 mg.
Nicotinic acid 0.04-0.94 mg.
IV. Miscellaneous: The following substances form a total of 4. I %.
Proteins (mainly from pollen) 0.3-0.5%
Fats 0 to traces.
Acids (formic, acetic, malic, citric, succinic, amino)
Pigments (carotene, xanthophyll)
Chlorophyll (decomposition products)
Bees'-wax
The following occur in even smaller quantities:
Enzymes (invertase, diastase, catalase, inulinase)
Aromatic bodies (terpenes, aldehydes, esters)
Higher alcohols (mannitol, dulcitol, etc.).
Other sugars (maltose, sometimes melezitose, etc.).
The calorific value of 100 grams of honey is 280 to 330 calories, the normal
pH value about 4.5.
An examination of this list shows that there is no rapidly energiz-
ing food-component in honey to explain its quick physical effect
apart from the dextrose and levulose. The great and easy absorp-
tivity of these sugars may be considerably enhanced by the effect
of other components such as vitamins, calcium, phosphorus and
others. None of these latter has the rapid effect of the sugars. It is
the habit of primitive man to eat the honey in the combs, together
with the bee-maggots and bee-pupae and with a large quantity
of pollen. The bees'-wax itself has no digestive value. But the
proteins of the bee-maggots and of the pollen, as well as the high
concentration of vitamins in the latter, add considerably not only
to the calorific value of honey but also result in a greater balance
in its dietetic value. This raised value gains in importance as the
significance of honey increases within the general diet. Thus with
the Guayakis, Veddas and African Pygmies, among whom honey

36
is the principle food, this habit of consuming maggots, pupae and
a certain amount of the stored pollen assumes great significance
and importance.
Among the many medicinal virtues of honey its anti-haemorrhage
and its bactericidal properties are undisputed. Some over-enthu-
siastic statements and claims, however, such as found in BECK and
SMEDLEY (1947, pp. 120 ff.), should be accepted with great reserve.
Honey mead, i.e. a fermented watery solution of honey was the
nectar of the gods, not only in ancient Greece. Another refreshing
drink is hydromel, a fresh mixture of honey and water.
Honey, whenever and wherever available, has always been a
most prized food since Palaeolithic times and was rightly considered
a highly effective restorative and guarantee of physical strength and
a welcome guardian of health. Many primitive people, such as
Pygmies, would not be able to survive in their last refuges in the
primeval tropical forests, if they were not able to live mainly on
honey. Half a kilo of honey provides their daily calorific require-
ment, while the addition of bee-maggots as a rich source of proteins
and also of fats, as well as vitamins and minerals, make honey a
well balanced food, especially as considerable quantities of pollen
are taken simultaneously. Thus honey became one of the most
important items of food throughout the tropics, before man reached a
high degree ofefficiency in hunting or the level of primitive husbandry.
A number of similar sweets playa more restricted role in native
nutrition: certain honeys of social wasps in South America, those
of the 'honeypots', various desert ants (Myrmecocystus in America,
Melophorus in Australia), and the honeydew excretions or mannas
of various cidadas, plant-lice and scale insects in various parts of
the world. Even if the quantity of these sources of honey is limited,
they are easily digestible and of great value as a restorative from
fatigue especially ifbees'-honey is not available.
These analyses and their comparison with the calorific and nutri-
tive value of other basic items of food lead us to the unavoidable
conclusion that insects are most nutritive and only to be compared
with the most valuable types of food. Only where big game or a
thoroughly balanced, rich diet in highly agricultural nations are
available in adequate quantities, can the dietetic importance of
insects be ignored and neglected. Among the primitive food-

37
gatherers, who cannot disregard anything which nature may provide
in the way of edible food, insects form of necessity an important
part of the diet. In this case the only limiting factor is the quantity
available. All insects which are neither poisonous nor aposematic
(these are a small minority) are eagerly gathered for food, whenever
they are present in numbers and of a sufficiently substantial size.
Selection by taste becomes a quite secondary motive. In all higher
stages of sociological development, man is often rather selective.
The sporadic consumption of ants and the slight use made in many
parts of Australia of grasshoppers and termites are indicative of
such selective taste. Yet even for higher primitive men BEQ.UAERT
(1921, p. 197) still reports that caterpillars are often appreciated
as food in direct proportion to their size and abundance. Some of
the descriptions given in the following pages from Australia and
Africa, together with a number of photographs from African mar-
kets, not to mention the many reports of locust gathering for food,
will convince the reader that in the tropics there is no lack of edible
insects. At least in certain seasons, and to some degree the whole
year around, insect life is sufficiently abundant to offer adequate
quantities for human nutrition. In times of drought and famine
these insects may become the decisive food which permits .survival.
The especially important role of honey for all primitive peoples
cannot be over-emphasized. Particularly with locusts quantity has
never been a limiting factor. Today a large amount of locust flour
is sold as manure. 'World Trade' (1936, p. 42) announced that
almost 3000 tons of locust flour containing 9.7% nitrogen and
12.4% fat was available in the Argentine for export. Various
authors have discussed the value of locusts for manure in South
Africa (ADLER 1934) and in India (DAS 1945).
At the outset of the following discussion we can thus be fully
convinced that insects when eaten in quantity are an important
source of food. In cases where the normal diet is poor in animal
proteins or in animal fats, they offer a necessary supplement to the
main food. Honey provides a most important source of immediate
working energy. Vitamins, salts and minerals represent other
possible additions to the value of insect food. We shall now examine
the question of the extent to which primitive man has utilized this
potential food in his normal diet.