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Fig. 1. Side view of honey-bearer x 2 (honey-ant, Garden of the
Fig. 2. Dorsal view of honey-bearer of Camponotus inflatus.
(After Lord Avebury.)
Fig. 3. Dorsal view of Fig. 1.
Figs. 4 and 7. Views of males of Hortideorum.
Figs. 5 and 10. Winged female, or virgin queen, of Hortideorum.
Figs. 6 and 11. Node, or scale, of the petiole of queen of honey-
Fig. 8. Worker-minor of M. Hortideorum x 4. The workers
major and minim, or dwarf, are similar in form.
Fig. 9. To show the striae (sir.), supposed stridulating organs of
Fig. 10. Winged female, or virgin queen, of Myrmica ruginodis.
Fig. 11. Node, or scale, of the petiole of No. 10.
For detailed description, see page 109 and following pages.
AND HOW THEY ARE GOVERNED
A STUDY IN NATURAL CIVICS
HENRY CHRISTOPHER McCooK
"TENANTS OF AN OLD FARM"
ILLUSTRATED FROM NATURE
HARPER 6- BROTHERS PUBLISHERS
NEW YORK AND LONDON
Copyright, 1909, by HARPER & BROTHERS.
Published October, 1909.
GENERAL ANSON GEORGE McCOOK
COMPANION, COMRADE, FRIEND, ALWAYS LOVING AND
WELL- BELOVED. IN EVERY OFFICIAL TRUST
BRAVE, PATRIOTIC, COMPETENT, AND
INCORRUPTIBLE. IN EVERY RELATION
OF LIFE JUST, HONORABLE
LOYAL, AND KIND
A FULL-ORBED MAN 7
PREFACE ............... v
I. FRATERNAL CONFEDERACIES AND COMMUNAL RIGHT-
EOUSNESS AMONG SOCIAL ANTS ....... 1
II. NESTING ARCHITECTURE HOUSING THE COMMUNE . 17
III. ENGINEERING METHODS IN ANT STRUCTURES ... 49
IV. SUPPLYING THE COMMUNAL RATIONS ...... 77
V. FEEDING THE COMMUNE .......... 101
VI. THE LANGUAGE OF ANTS AND OTHER INSECTS . . . 121
VII. How ANTS COMMUNICATE .......... 130
VIII. FEMALE GOVERNMENT IN ANT COMMUNITIES . . . 155
IX. THE PROBLEM OF COMMUNAL DEPENDENTS . . . . 171
X. WARRIOR ANTS, AND THEIR EQUIPMENT FOR WAR . 190
XI. How ANTS CARRY ON WAR ......... 211
XII. ALIEN ASSOCIATES AND AFFINITIES IN ANT COMMUNES 224
XIII. APHIS HERDS AND ANT ASSOCIATES ...... 243
XIV. THE FOUNDING OF SLAVE-MAKING ANT COMMUNES . 261
XV. PROBLEMS OF SANITATION AND PERSONAL BENEVOLENCE 279
XVI. A NOTE IN REVIEW ............ 296
TABLE OF AUTHORS AND REFERENCES ..... 305
HONEY-BEARING ANTS Frontispiece
A GROUP OF MOUND-MAKING ANTS* NESTS 4
WATCHMEN AROUND THE GATE OF A HONEY-ANTS' NEST . . 10
GROUP OF TWO MOUNDS OF F. EXSECTOIDES AT ALGONAC, ST.
CLAIR RIVER, MICHIGAN 12
SINGLE MOUND OF F. EXSECTOIDES, BRUSH MOUNTAINS, PENN-
ORIGINAL CELL OF CARPENTER ANT QUEEN, AFTET, WHEELER 18
EXTERIOR ARCHITECTURE OF HONEY-ANTS OF THE GARDEN OF
THE GODS 19
INTERIOR OF A STORIED COMMUNE OF THE OCCIDENT ANT OVER
EIGHT FEET DEEP
A CONNECTED SERIES OF STORE-ROOMS, OR GRANARIES AND
CHAMBERS, OF THE OCCIDENT ANT 21
PEBBLE-ROOFED COMMUNE OF THE OCCIDENT ANT .... 24
LITTLE HEAPS OF EARTH-PELLETS THROWN UP AROUND THE
GATES OF PHEIDOLE PENNSYLVANICUS, THE SIMPLEST
TYPE OF ANT-MOUND 27
CREMASTOGASTER LINEOLATA 28
CARTON COCCID-TENT OF CREMASTOGASTER LINEOLATA-PILOSA
PASTEBOARD NEST OF DOLICHODERUS BITUBERCULATUS MAYR,
FROM BANKOK 31
NEST OF CAMPONOTUS RUFIPES ON A TREE 33
NEST OF PSEUDOMYRMA BELTI ON MEXICAN ACACIA THORNS 36
PSEUDOMYRMA BELTI (EMERY) MAGNIFIED 36
PRELIMINARY ENGINEERING 39
SECTIONAL VIEW OF ANT-MOUNDS 42
ANT ENGINEERING 43
MOUND-MAKING ANTS COVERING A DOUBLE GALLERY ... 46
ANTS DIGGING OUT GALLERIES 53
AN AGRICULTURAL ANT'S FOOT, ENLARGED 180 DIAMETERS!
THE TOOL USED, WITH THE JAWS, IN DIGGING .... 54
THE CIRCULAR DISK AND ROADS THAT SURROUND THE ONE
CENTRAL GATE OF THE AGRICULTURAL ANT OF TEXAS . 56
AGRICULTURAL ANTS CUTTING GRAIN TO CLEAR ROADS AND
ANT CARRYING A PELLET OF SOIL 57
A SPECIMEN OF GRAIN-STALK ON AN ANT CLEARING .... 57
AGRICULTURAL ANT CARRYING GRAIN-STALKS 57
OCCIDENT ANTS: SINGLE AND DOUBLE GATES OPEN .... 67
OCCIDENT ANTS CLOSING GATES 68
A CLOSED GATE OF CUTTING ANTS OF TEXAS 73
CUTTING ANTS: A GATE WIDE OPEN 74
CUTTING ANTS: A GATE IN PROCESS OF CLOSING 75
GUSTATORY ORGANS OF THE HONEY- ANT . 80
AGRICULTURAL ANT FEEDING FROM A HICKORY-NUT KERNEL 85
MOUND NEST OF TEXAS CUTTING ANT UNDER A LIVE-OAK TREE
PARTLY DEFOLIATED 87
THE HEAD OF A TEXAS CUTTING ANT 88
A PROCESSION OF TEXAS ANTS CARRYING LEAVES TO THEIR
CUTTING ANTS 93
SECTIONAL VIEW OF A CUTTING ANTS* NEST 94
A PIECE OF LEAF-PULP WHICH FORMS THE MUSHROOM GARDEN 96
SPRIG OF DWARF-OAK (QUERCUS UNDULATOR), WITH GALLS
EXUDING DROPS OF SWEET SAP 109
THE DIGESTIVE TRACT OF A HONEY-ANT 110
HONEY-BEARERS OR ROTUNDS Ill
ABDOMENS OF HONEY-ANTS 112
SECTIONAL VIEW OF THE STOREROOMS OF THE OCCIDENT ANT 115
OCCIDENT ANT HARVESTING SEEDS OF WILD SUNFLOWER . 116
POSTURES OF THE FLORIDA HARVESTER IN CUTTING SEEDS FROM
THE STEM 117
SECTION PENNSYLVANIA HARVESTING ANT 118
MIMETIC LANGUAGE IN ANTS' 123
ANTS IN ATTITUDE OF COURAGE, ANGER, AND ALARM . . . 125
PTERATIC LANGUAGE 132
DEATH'S-HEAD MOTH WHICH MIMICS THE PIPING OF A QUEEN BEE 139
STRIDULATING ORGANS OF LOCUST AND CRICKET. RED-LEGGED
LOCUST (MELANOPLUS FEMUR-RUBRUM) 141
JAPANESE CAGES FOR STRIDULATING INSECTS 142
PROBABLE STRIDULATING ORGANS ON ABDOMINAL PLATES OF
THE FACE OF AN ANT, SHOWING THE FLEXIBLE ANTENNA . 148
THE CHALLENGE WITH CROSSED ANTENNAE 151
A QUEEN ANT SURROUNDED BY HER COURTIER GUARD . . 157
A PEEP INTO AN ANT COMMUNE 159
WORKERS TAKING EGGS FROM THE QUEEN MOTHER . . . 161
A TRUANT QUEEN BROUGHT HOME 163
AN EMMET NURSERY FOR THE YOUNG 166
YOUNG WINGED QUEEN OF HONEY-ANT 173
A MARRIAGE-FLIGHT, OR " SWARM," OF WINGED MALE AND
FEMALE ANTS . . 176
ARRIVAL OF THE FOOD-BEARERS AMONG THE WINGED DE-
THE BRAINS OF ANTS 1S3
WINGED FEMALE ANTS AT PLAY ON THE PLAZA 187
WAR-LIKE DWARFS ATTACKING AN OCCIDENT ANT ... 196
A PLUCKY LILIPUTIAN ATTACKING AN OCCIDENT AXT . . . 197
SLAVE-HOLDING ANT 200
THE FLORIDA HARVESTER (POGONOMYRMEX CRUDELIS) . . 204
MANDIBLES WHICH ARE USED AS WEAPONS BY WORKERS OF
AGRICULTURAL ANTS 206
USING THE STING IN FLIGHT 207
THE BEETLE XENODUSA CAVA LECONTE 230
PHEIDOLE INSTABILIS AND ITS PARASITES 233
ORASEMA VIRIDIS 234
PUPA OF ORASEMA 235
SOLENOPSIS FUGAX 254
SECTIONAL VIEW OF HOST-ANTS' NEST, SHOWING THE GAL-
LERIAS WHICH THE THIEF-ANTS ENTER 255
WORKER ANTS DEPORTING CAPTIVES OR THEIR FELLOWS DURING
A THIRD LEG OF ANT, SHOWING HAIRS AND SPINES USED IN
PERSONAL CLEANING 279
VIEW OF OUT-THRUST TONGUE OF AGRICULTURAL ANT . t . 280
MITES THAT ATTACK ARTIFICIAL NEST OF HONEY-ANTS .... 282
ANTS CLEANING STINGING APPARATUS AND BRUSHING BACK-
HAIRS OF THE HEAD 283
HONEY-ANT WORKERS IN ACTION 290
HONEY-ANTS AND ROTUNDS 293
THREE years ago, in his Nature's Craftsmen, the au-
thor presented a series of original studies of the life
history of sundry insects. Half of the book was given
over to one of his specialties the ants. The remainder
embraced accounts of another specialty, the spiders,
and of certain insects that had received particular
attention as a sort of by-product of his special studies.
The author's purpose therein was to give his readers a
veritable natural history of the subjects treated of, in
popular form, and clothed in at least some measure of
the simple graces of good literary style.
The present volume, while aiming to preserve the
above features, differs from Nature's Craftsmen, in that
it is limited to the natural history of ants. Moreover,
it considers mainly those phases of their life that are
developed around their behavior as social animals.
It is here that appear most clearly and fully the habits
which have drawn to these insects from the earliest ages
the attention of man, and have won for them a high
reputation for wisdom.
From this has arisen a secondary feature of the book
-viz., the indication of parallels, more or less distinct,
between the communal actions of ants considered
simply as natural history, and the communal actions of
man, considered, as all human beings are bound to
consider them, in their relations to the highest welfare
of the race.
The association of separate groups of individuals to
accomplish the primary aims of physical life is almost
sure to develop resemblances in methods. What are
those aims? Among ants, as the following pages will
show, they are the establishment of a home; procure-
ment of a livelihood; protection from enemies; preserva-
tion and nurture of the young and other communal
dependents; perpetuation of the species, and the prop-
agation of the commune.
Wherein do these differ from the common necessities
and aims of men in their social aggregations? They
are practically the same. The Great Hand of Sovereign
Nature that has laid upon her children these common
aims has so guided them in the achieving thereof, that,
amid the endless variations which issue from an Infinite
Fountain of Design and Force, one traces resemblances
in methods that suggest their common origin. In our
studies, these likenesses, as well as unlikenesses and
contrasts, will be interesting to note.
In many of the higher and complex duties of hu-
man communities it is impossible that insects should be
models for men, in whom there is an element that sep-
arates from all other creatures by an impassable gulf.
But in the great physical functions of a commune, which
are a bond of sympathy between us, we may have
something to learn from the ants, who manifestly have
kept and still keep to the primitive ways of nature more
closely than we. Sometimes these lessons have been
pointed out, sometimes simply suggested, sometimes
left for the reader to discern. But whether the one or
the other, the author ventures to hope that they may
stimulate thought and discussion, or at least interest,
but in no wise divert from his main purpose: to increase
knowledge of the natural history of creaturelings that
have contributed so much to his own enjoyment of life.
It has been more than thirty-two years almost a
full generation of men since the author published in
the Transactions of the American Entomological Society
(Philadelphia) his first observations of American ants.
Since then he has given to the world in books, in publica-
tions of scientific societies, and in magazine articles, his
studies of various species, chiefly devoting himself to
their habits rather than to their systematic classification.
In this volume, Ant Communities, taken together with
the first part of Nature's Craftsmen, he presents a sub-
stantial summary of these prolonged observations.
And he has brought them down to date by associating
therewith the latest observations of some of the leading
naturalists of the scientific world.
" Brookcamp," Devon, Pennsylvania.
FRATERNAL CONFEDERACIES AND COMMUNAL RIGHT-
EOUSNESS AMONG SOCIAL ANTS
ORGANIZED society, whether among insects or
men, implies some form of government; and that
implies citizenship. And fidelity to the just and natural
service of citizens is communal righteousness. May we
apply such a term to insects? And if so, what is the
character of such a quality; or, if one may venture so to
put it, what is the quality of such a character? And
is it in any measure comparable with communal right-
eousness as the phrase goes among men? The inquiry
will here be limited to ants; but the study requires the
statement of some preliminary facts, so that readers
may have a true conception of the field which our
thoughts are to explore.
Some insects are "solitary"; they live alone. Others
are "social"; thev live in communities. There is such
a striking contrast between the manners of the two
groups that one wonders how the distinction arose.
True, at the beginning of life most insects are massed,
since their mothers lay their eggs in compact clusters.
But if one start with the theory that this may have left
in the germ of being a tendency which, under favorable
conditions, might be transferred to the adult, he is met
by certain facts that may confound his reasoning.
For example, the eggs of ants and bees are dropped
separately, yet they produce insects of the strongest
social habits. The moth of the tent caterpillar oviposits
in clusters, and her progeny keep together in the larval
state. The eggs of the garden orbweaver, like those of
most spiders, are laid in carefully sheltered masses, and
the young are partly reared together in the silken tent
which the mother overspins. Moreover, they start in-
dependent life in a self- woven silken compound. The
lycosid, a ground - spider, drags her round cocoon be-
hind her until the eggs are hatched, and then bears the
younglings about clustered upon her back. Yet soon
the centrifugal factor in vital force drives the young
of moth, orbweaver, and lycosid asunder, and thereafter
their life is solitary.
With social insects the tendency is reversed. Be-
ginning life solitary, as in the case of the maternal
founder of an ant's nest, the individual becomes a family,
and the family a community, and this may develop into
a vast commonwealth containing many thousands or
even millions of individuals. When the circle of life is
complete, the vital centripetal force which binds these
communities together is relaxed, in a movement of im-
passioned communal fervor, to allow the outgoing of the
winged males and females, as with ants ; or the swarming
of a new community, as with bees. This is the " com-
mencement" time in the insect calendar, when a matured
sliver of the community is struck off and pushed into
Among ants these communities vary in population
FRATERNAL CONFEDERACIES AMONG ANTS
from a few score to many thousands. There are villages,
towns, cities each, for the most part, independent of
all others, and each complete within itself, a separate
tribe, a sovereign state. That the orderly and successful
conduct of such communities must spring, consciously
or unconsciously, from some system, is self evident.
What is that system? What are its laws, its customs,
its methods of administration? Is an ant-hill a mon-
archy, a republic, a democracy, a socialistic commune?
How does its government compare if in any wise com-
parable with the civil governments of men? And
what lessons in civics can we learn therefrom?
Surely, an interesting inquiry here opens up; for,
whatever the result, it must give us a glimpse of nature
pure and simple. To this the author's purpose is mainly
directed; but, as a by-product of his studies, he con-
fesses a keen interest in those reflections that traverse
the field of human civics, and which inevitably arise
as one pursues the history of life in ant communes.
In many parts of the Alleghany Mountains, and in
middle and eastern Pennsylvania, in New Jersey, in
the White Mountains and elsewhere, are distributed
the large conical nests of the mound-making ants of the
Alleghanies, Formica exsectoides (Fig. 12). These vary in
size from newly begun colonies a few inches high to
mature hills, measuring thirty-seven feet in circumfer-
ence at the base, though rarely more than three feet
high. They occur in groups; and in one site near Holli-
daysburg, Pennsylvania, within a space of fifty acres,
the writer counted seventeen hundred well-developed
mounds. At two other localities in these mountains sim-
ilar groups were observed even more thickly placed. At
"Pine Hill" about thirty acres were occupied, of which
five were found to contain two hundred and ninety-three
mounds, an average of fifty-nine to the acre, or eighteen
hundred for the whole section. At u Warrior's Mark"
^^^S^S^::?B:V::. . i>' ^ ; SSS ' tj.
. -. p : ^^t^'/^iM^^^
\ ^ v ; >^;ffX; i i MmHfe
Fig. 12 A GROUP OF MOUND-MAKING ANTS* NESTS
another large settlement of nearly two hundred hills
was visited. Experiments made in the Hollidaysburg
group proved that all therein formed substantially one
community, in complete fellowship, although the in-
FRATERNAL CONFEDERACIES AMONG ANTS
dividual mounds appeared to be conducted indepen-
dently. The following will illustrate these experiments.
[McC. 2, p. 282.] 1
A small oak-branch covered with aphides and their
attendant ants was broken from a tree and placed
erect upon a mound twenty rods distant. It was
thought that if anything would incite to hostility, it
would be the intrusion of members of a separate com-
munity upon a congener's feeding -grounds. On the
contrary, the ants being called from the hill, came out
and mounted the branch with the usual excited bearing,
and then mingled with its original occupants on friendly
terms, and began to feed quietly from the galls and
aphides. A larger branch having many more ants
upon it was cut, and planted upon a mound a consider-
able distance beyond the first one. The insects were
called out by tapping upon the surface. The usual
whirl of angry sentinels and other workers followed,
and then all blended with the intruded ants without a
sign of hostility.
A spadeful of earth was swiftly cut from the mound,
and with ants, cocoons, and broken cells, thrown into a
pail, carried to a cone fifty rods distant, and cast upon
the surface and around the lower gates. One could not
distinguish between the citizens of the two mounds as
masses of excited ants poured out and began their
usual movements, but no marks of hostility appeared.
After the first sharp challenges with crossed antennae
the imported ants melted away into the general com-
munity as though at home.
The only other test of this nature which need be
mentioned was made with three hills (D, E, F), to
1 See " Table of Authors," Appendix.
which reference will be made hereafter as the " hysterical
hills/' on account of the abnormal state of excitement
which marked their inmates, and for which no reason
was apparent. Large pieces of the mounds D and E,
which were twelve feet apart, were interchanged, tossed
violently from one to another. Although swarming
with insects intensely agitated, there was no appear-
ance of hostility at either mound.
I then proceeded to F, one hundred and fourteen feet
distant, and called out the ants until the cone was
fairly black with them. From the densest centre of
life was cut out a section about six inches square and
borne hurriedly to D, catching en route the dropping
ants in a hat. The contents of shovel and hat were
thrown upon the cone in the midst of its hosts of in-
habitants. Even this violent invasion which, with an
alien species would have been a signal for war and
slaughter, was not resented. There were sharp antennal
challenging and quick response, and then the new-
comers melted away into the mass of their enforced
hosts, as fellow-citizens "to the manner born." There
was complete fraternization, which was not afterward
disturbed by any breach of the peace.
The final test w y as an artificial nest prepared in a large
glass jar within which earth, sticks, and surface litter
were placed. Ants taken from a number of mounds
situated in parts of the field and wood most remote from
one another were put in. Cocoons from yet other
cones were added. Aphides, water, and honey were
then given them. This miscellaneous assemblage united
with the utmost harmony in building galleries, caring
for the cocoons, and defending the nest from intruded
ants of separate species and from spiders. From time
FRATERNAL CONFEDERACIES AMONG ANTS
to time ants and cocoons collected from widely separated
hills were put in, and these were always and at once
adopted. This amity and co-operation in the duties
and responsibilities of good citizenship continued until
the composite republic of drafted citizens was broken up.
The natural explanation of these rare conditions is
this: the antennal interchanges between the various
parties at once showed that all were fellow-citizens of
one commonwealth, equally entitled to communal wel-
come and place, which were accorded at once upon the
recognition of the one common nest-odor which is the
badge of citizenship.
Thus it appeared (and to the writer it was then an
astounding revelation) that among the myriads of
creatures occupying these more than seventeen hundred
mounds there was complete fraternity if, indeed, they
were not one mighty confederacy! Here was a republic
which in the number of its separate states for every
mound was an independent community of ants and in
the multitude of its total population exceeded the most
sanguine prophecy of the future American republic.
It would be hard to conceive of anything like local or
communal loyalty, an inflated devotion to "state rights,"
or that jealousy and conflict of interests which are apt
to develop among neighboring communities, as leading
on to war among the insect commonwealths which were
the subjects of the above experiments.
If a city be (as it has been defined) "a place inhabited
by a large, permanent, organized community," the
name "Ant City," by which it is popularly known, is
fitly given to this vast concourse of united emmets, or,
indeed, to any one mature colony thereof. Naturally
the question often occurred, How many ants are here
assembled? An exact census in such a case is im-
practicable, but at least a reasonable approximate is
possible. Dr. August Ford, the eminent Swiss myr-
mecologist, has described a community of two hundred
mounds of a closely related species (Formica exsecta)
among the mountains of Switzerland, as having each a
population ranging from five thousand to five hundred
thousand. If one were to apply the lowest estimate
(five thousand) to our American community, it would
give a total population of eight and a half million living
creatures! That is quite enough to justify their claim
to the title of "city," but, in truth, a conservative esti-
mate would make them many times as numerous.
In his Die N ester der Ameisen (Ants' Nests) [F. -1 and
5], in commenting on my observations upon this sodality
of the Alleghany Mountain ants, Doctor Forel says:
"These ant kingdoms have, in all probability, a popula-
tion of two hundred to four hundred million inhabitants,
all forming a single community, and living together in
active and friendly intercourse." Think of it! A
population equal to that of the whole empire of China!
And this is -not a wild guess of an enthusiastic vision-
ary, but the sober calculation of the veteran chief of Eu-
rope's myrmecologists, and one of its foremost medical
specialists. I have spoken of this mighty concourse of
organized insects as a city; but doubtless kingdom, or
empire, might be a better title, for there was through-
out the settlement a marked tendency to groups of
mounds of different sizes, which might represent the
cities or large centres of population distributed through-
out a commonwealth.
No North American ant exceeds these mound-builders
in the size of the structures reared by them. But in
FRATERNAL CONFEDERACIES AMONG ANTS
some tropical lands even larger mounds are found.
Livingstone speaks of ant-hills in South Africa that
dotted the face of the country like haycocks in a harvest-
field. In the woods they were twenty feet high and
forty to fifty feet in diameter! [Li. 1, p. 590.] These
rival the great gothic erections of the termites. Whether
the African ants show the communal unity that exists
among our Alleghany mound-builders has not been
determined. But such unity must obtain among the
vast hordes that occupy each hill.
One who studies the economy of these communes
soon notes a well-defined division of labor among the
three worker castes viz. t workers-major, workers-minor,
and minims or dwarfs. There are sentinels or police-
men, masons or builders, foragers, nurses, and courtiers
or queen's body-guard. These are not so differenced
as to form fixed classes which embrace always the same
individuals with duties limited to one sort of service,
as is the case in some other species. Apparently, all
branches of service have recruits from all the castes, and
these pass from one duty to another at will. On the
surface (as far as human intelligence discerns) it is a
"go-as-you-please' 1 arrangement, which nevertheless is
dominated by some occult principle that brings orderly
results out of seeming chaos.
There appears to be no specialized warrior caste among
these lormicans, but there are sentinels, or policemen,
whose duty it is to guard the community from hostile
approach. Their internal affairs call for no domestic
police. Among these millions of citizens there is not
one criminal, not one degenerate! I do not recall, in
all my long and varied observations, a single example of
an ant whose actual offending called for the administra-
tion of civil punishment. Nor do I remember to have
read of such a case in natural history. Emmet out-
lawry is unknown. These vast communities are self-
policed. Their citizens are so perfectly self-controlled,
so absolutely free from even the desire to violate law,
that as against them a domestic police would be a sine-
cure. Do you look for the perfect social commune whose
citizens are all perfect in that " righteousness which is of
the law"? One may find it here.
But public enemies abound. Eternal vigilance is the
price of peaceful industry and security. Here are
millions for defence, though not an ant be needed to
support home government. And to this end every
citizen, if need be, is a soldier-policeman. Watchmen
continually guard the various gates, or entrances, to the
cone, most of which are ranged along its base just above
the ground, but some are placed between that and the
summit. These sentries lurk inside the gates, whence
they issue, with every mark of intense excitement and
watchfulness, if one approaches a finger or drops some
object near them. Frequently they patrol the vicinity
of the gates, and
IrLTrXt-fcW*?*;- >^/*.~- <*'A xr <T\J.*I tSVkVSJ '^,\ t
This is a p rev-
rig. 13 WATCHMEN AROUND THE GATE OF
A HONEY-ANTS' NEST alent manner oi
ants. There is
but one large tubular entrance to the pound-cake-like
mound of the honey-ants of the Garden of the Gods
(Myrmecocystus hortideorunt) in Colorado. Around its
upper rim, with their yellow heads and quivering antennas
FRATERNAL CONFEDERACIES AMONG ANTS
just in view, one sees a ring of sentinels. At the door
into the pebble-sheathed cone of the Occident ant (Po-
yonomyrmex occidentalis) of the American plains, which
opens into the breast of the cone, the watchmen wait,
intent and vigilant. So it is elsewhere, and almost
everywhere, that ants are found in large communities.
It is the law of emmet as it is of human society, that
"Some must watch, while some must sleep;
So runs the world away!"
These watchmen do not always belong to a soldier
caste. Every emmet citizen who has passed the brief
callow stage of first emergence from pupahood is a
policeman or soldier on occasion, and may, as far as the
facts now appear, go on sentry as on any other duty.
It would not be strange if, in the gradual development
of such a social system, certain individuals should have
shown special aptitudes for police service that kept them
more or less continuously therein, and so have arisen
something like a soldier class. In some species such
has been the case, as with those of the genus Pheidole,
and the leaf-cutting or parasol ants of Texas.
But it is not so with our mountain mound-builders.
They remind one of the militia organization of our earlier
frontier States Ohio, for example, which made every
adult male, not disqualified by age or otherwise, subject
to military duty. Indeed, such is, in theory, the relation
of all citizens of the American republic to the general
government. Among our ants that duty is never dodged.
There are no desertions. Lazy, cowardly, and skulking
ants one does not see. With heartiest good-will the
call to service is met, and a " clear call," apparently, is
simply a perception of the commune's danger and need.
Then, at a touch, every citizen becomes a warrior, and
the outer walls swarm with defenders.
Here one may note a remarkable trait of these ant
citizens their devoted patriotism. At the approach
Fig. 14 GROUP OF TWO MOUNDS OF F. EXSECTOIDES AT AL-
GONAC. ST. CLAIR RIVER, MICHIGAN
(Photo by William S. Cooper, Detroit)
of an enemy they attack it, absolutely regardless of con-
sequences. The personal factor has no place nor even
consideration in the act. Whether the supposed hostile
be great or small, beast, bird, creeping thing, or man
himself, the brave little creatures fling themselves upon
it with the utmost abandon.
For example, here comes to the edge of the mound a
large black carpenter ant (Camponotus pennsylvanicus),
a ferocious and formidable insect, almost twice as large
as the formican and a hereditary enemy. Forth from
a gate leaps a sentinel, and launches its quivering body
straight against the sable giant. One snap of the
Camponotid's jaws, and the assailant's brown head is
FRATERNAL CONFEDERACIES AMONG ANTS
severed, and its beautiful life extinct. Another sentinel
follows, and another, only to meet the same fate. But
others crowd to the combat, eagerly facing and meeting
wounds and death. Overwhelmed by numbers, the black
warrior is at last conquered and dragged into the
formicary, where its dismembered body is sucked dry
and its shelly parts dumped upon the refuse-heap, or
mayhap built into the growing walls, along with vegetable
debris of various sorts.
This courageous and unselfish disregard of person
and absolute devotion to the communal safety at the
cost of life or limbs is characteristic of ant citizens ; not
of a few, but of all; not rarely and occasionally, but
always; not under compulsion, but freely and without
Fig. 15 SINGLE MOUND OF F. EXSECTOIDES, BRUSH
reward of any sort. Not even the high stimulus of
applause of comrades and of honors from their fellows
urge and sustain them. Such conduct is so much a
matter of course that no one notes it as extraordinary,
and a war-scarred veteran may be seen dragging its
maimed limbs into action, or in some obscure corner
licking its hurts and waiting for the end without nurse
or comforter. It has done its duty, and accepts the
result with imperturbable unconcern, as do its fellows.
Apropos of these studies of police administration of
our ant commonwealth is an observation incidentally
made while conducting experiments to determine the
mode of recognition among ants. Starting upon the
theory that it was a specific odor or emanation analogous
thereto by which our mound -makers recognize one
another, the matter was tested by subjecting individuals
to baths of clear water, and infusions of wintergreen,
cold coffee, and tea, and then returning them to their
mounds. The individuals thus treated were immediate-
ly attacked by roundsmen, a dozen or more sometimes,
and dragged away like culprits. These assailants were
then taken with their victims, submerged, and festered
to the hill with the same result. So with a third
series; the assailants of the assaulted ants were in turn
attacked, and invariably the same measure meted to
them that they had measured to others. They had lost,
for the time at least, the "mark" 1 of their citizenship.
[McC. 2, p. 281.]
In some cases the parties assailed were soon released,
as though the mistake had been perceived. But for
the most part there was every indication of a mortal
purpose and a fatal issue. It was here that a curious
trait was developed. The demeanor and conduct of the
immersed and "tainted' 1 ' ants were in marked contrast
with their character for valor in battle and pluck
generally. They were quite passive under the fierce
FRATERNAL CONFEDERACIES AMONG ANTS
assault of their fellows, and succumbed with little or no
effort to resist. They seemed to have the carriage of
persons detected in some meanness or crime a " hang-
dog'' sort of air.
Could it be that these unfortunates tacitly recognized
the fact that they had become obnoxious to the com-
munal police? And, although this had come about by
no fault of their own, was their instinctive sense of
obligation to submit to the " legal authority' which
dominated the commune so imperative that they
yielded themselves to their fate, temporary captivity
or death as the case might be, without the least show
of resistance? One's judgment is so apt to be biassed
by his interest in and sympathy with these wise little
creatures that he is inclined to distrust even his most
careful observations, and fear that unconsciously he may
have interpreted their behavior by the operations of
his own mind. But in this case so many tests were
made, all yielding like results, that the above conclusion
seemed to be justified.
And why should it not be so ? The higher animals are
not insensible to the public sentiment of their kind,
as one may see from the actions of domestic flocks and
herds and of gregarious wild beasts. It is what might
be looked for in social insects, though therein less notice-
able by human senses; for ages of hereditary com-
munal life must have wrought upon their sensibilities,
so keen in certain quarters though defective in others,
a marked response to an environment of active dis-
One does not speak of this as a conscience, perhaps not
even as a remote analogue thereof. But it seems to take
the place of that sentiment, or experience, or inward
impulse and restraint in man, without which no com-
munal government is long possible. What is it that
imparts to our genus elements of chaos, crime, misrule,
and misery so far beyond the qualities of social insects
(if also so vastly above them), and which starts up in
the path of history records of communal disorder that
one seeks in vain among ants, hornets, and wasps?
Why should a creature with a conscience ever be less
steadfast and exemplary in communal righteousness
than a citizen of a commune of mound-making ants?
NESTING ARCHITECTURE HOUSING THE COMMUNE
THE housing of the commune is a duty that springs
up side by side with the existence of the commune.
In the typical beginning of an arit community by the
single fertilized queen, the first act of the incipient
foundress is to scoop out and heap around her, in earth
or wood, a cell whose diameter is somewhat greater
than the length of her own body. This is the rudimen-
tary house of the commune the primitive cave which
bounds the architectural aim of most animals, and
which is the starting-point with man himself (Fig. 16).
With great numbers of species, this cave will be found
under a stone. A flat stone, not too large and not
deeply imbedded in the ground, if lifted up in the early
spring, or at any time during summer, will be found to
serve as a rocky roof which overspreads the vestibule
and protects certain galleries, halls, and passages into
an underground formicary. The mere fact of choosing
such a location for a nest is significant; for, besides the
protection and defence afforded, the stone absorbs the
sun's ravs and serves thus as a natural furnace, con-
tributing to the warmth of the ants and of their imma-
Like the ancient catacombs of Rome, which served
primitive Christians equally for home, for sanctuary,
and for cemetery, these subsurface chambers and
galleries are arranged in irregular stories, one above
another. They are simply the primitive cave in multiple,
with intercommunicating passages. And they increase on
the principle of any other social settlement to meet the
Manv of them reach
i m m e n s e propor-
tions ; most of them
With the great
army of woodwork-
ers the same simple
type of architect-
ure prevails, modi-
fied simply and not
largely by the ma-
terial from which
the public build-
ings are wrought.
The storied subdi-
are crowded within
a narrower space
and ai'C leSS
courtesy of the American Museum of Natural History)
Fig. 16 - ORIGINAL CELL OF CARPENTER
ANT QUEEN, AFTER WHEELER tmctty HI a 1' k C d .
One who carefully
studies the architecture of a long-established nest of
carpenter ants will find himself unconsciously tracing
out in miniature pillars, arches, aisles, vaults, and
domes of different orders of architecture. It takes but
a slight stretch of fancy to imagine that one is gazing
upon the ruins of an ancient seat of a diminutive type
of his own race, who had carved out their toy-like homes
and temples in the solid wood. 1
One of the most interesting examples of the storied
type of underground architecture is that of the honey-
ants of the Garden of the Gods (Myrmecocystus hor-
tideorum), Colorado, the farthest north they had been
observed. The approach to their nest was a small,
low, pebble-covered mound with a large central gate
which penetrated it vertically for a few inches, and then
was diverted into various passages that followed the
slope of the ridge on which the colony was planted
In one nest, chosen for complete exploration, excava-
tion was carried forward during three days and several
^=%y 5p3$^fflp' ->
Fig. 17 EXTERIOR ARCHITECTURE OF HONKY- NTS OF THE
GARDEN OF THE GODS (Myrmecocystus hortideorum)
parts of days, two men working with mallet and chisel
and with knife in the soft, red sandstone, or " pudding-
stone," of which the ridge is composed. The entire
length of the formicary was seven feet eight inches.
1 Nature's Craftsmen, p. 120.
The point at which it ended was forty and a half inches
below the level of the main gate and twenty-nine and a
half inches beneath the level of the hillside. In all, the
ants had excavated thirty-six cubic feet of rock, and
this space was honeycombed with galleries and rooms.
The latter varied from five to six inches long, three to
four wide, and about three-fourths of an inch high.
The walls and floors of these rooms were smooth, but
the roofs were left in their natural roughness, thus
forming a better foothold for the rotunds, or honey-
bearers, who were perched upon them, clinging thereto
with their claws, and closely clustered together. [McC.
4, pp. 36, 37.]
The Occident ant (Pogonomyrmex occidentalis) is closely
related to the agricultural ant in structure and habit
(Fig. 18). But the typical forms vary decidedly in their
exterior architecture, the Occident having its commune
overbuilt with a prominent cone coated with pebbles,
while the typical agricultural keeps the space around its
gate free from all growth. Both species, like the mound-
making ants of the. Alleghanies, are among those that
found and maintain vast communities, and therefore
have a special interest to us in our present studies.
Their homes are often wrought in a tough clay that is
almost as hard to excavate as the red sandstone of the
Garden of the Gods, and equally taxes the resources of
the workers. The arrangement of rooms into stories
is here also carried out, and to a surprising extent. In
one nest of the Occident ant a story was found at a depth
of over eight feet beneath the surface (Fig. 19).
Those who are curious in such comparisons might
find grounds here for a striking parallel between the
achievement of an ant three-eighths of an inch high
(long), and of a man one hundred and seventy-six times
as high (five and one-half feet). [McC. 4.] \Yere we
to reckon a proportionate rate of progress between
the two on the basis of height, our man would have to
'-^ i- ";'' -^^fe^'"^ ' %,.
Fig. 18 A CONNECTED SERIES OF STORE-ROOMS (A, B, C, D) , OR
GRANARIES AND CHAMBERS OF THE OCCIDENT ANT
be credited with a storied structure one thousand four
hundred and eight feet deep. Apart from such fanciful
comparisons, it is certainly well calculated to excite our
wonder that such insignificant creatures can, by their
united exertions, bring about results relatively so vast .
unaided by mechanical contrivances.
The numerous chambers which in honey-ant structures
.;-.- ' ^ ' . -. .:.'/:
: -. . '' .-...-.
;.;.;..;: v . . v - .:
Fig. 19 INTERIOR OF A STORIED COMMCNE OF THE OCCIDENT
ANT OVER EIGHT FEET DEEP
are occupied by those living honey-pots, the rotunds, in
the Occident nests are used as store-rooms. Herein
one finds various sorts of seeds put away for food. In
a few cases rooms were found rilled with husks and
apparently sealed up, as if empty spaces had been
utilized in trie rush of business for "dumping-grounds,"
to save transporting the waste matter of the seeds to
the outer gates and the kitchen -middens. Perhaps
these "relief chambers 75 were merely a temporary
makeshift, and would have been cleared out in due
course had not the commune suffered a destruction as
dire as that of ancient Troy or Carthage.
Such great structures as have been described here
imply the work of years, and it is probable that some
of them were several years old. They showed every
mark of such age; in fact, the continuous life of an
ant community, in such sharp contrast with that of our
hornets and yellow - jackets, which do not survive
October, would naturally demand permanent or con-
tinuous residences, the permanency of the community
and the permanency of their dwelling going naturally
hand in hand. By calculations made from the levelled
floors of the mountain charcoal-burners, which had been
occupied by large mounds since their abandonment, I
concluded that some communities of Formica exsectoides
were at least thirty years old, and I believe that they
remain active for a longer period if unmolested.
Livingstone (South Africa) speaks of ant-hills which
dotted the face of the country like haycocks in a harvest-
field. In the woods they were seen twenty feet high
and forty to fifty feet in diameter. He also notes the
fact that these spots are more fertile than the rest of
the land, and are the chief garden ground for maize,
pumpkins, and tobacco. This statement has a signifi-
cant bearing upon the part assigned in nature to ants
and other insects in making the earth habitable by
agricultural man. [Li. 1, p. 590.]
The pebble roofing of the cone of the Occident ant is a
permanent feature (Fig. 20) at least, of the immense
number seen by me, all were covered with pebbles of the
gravelly soil in which they stood. In the vicinage of the
Garden of the Gods the pebbles were red sandstone.
The mounds in Wyoming observed by Prof. Joseph Leidy
were covered with a white stone. Mr. R. Hill saw them
Fig. 20 PEBBLE-ROOFED COMMUNE OF THE OCCIDENT ANT
on the Sapa Creek, in northwestern Kansas, roofed with
pellets of the limestone rock in which the great fossils
are found, and in one or two cases even of portions of
the fossils. Thus the conditions of the famous riddle
of the Judipan Hercules are repeated in this far Occident,
and the hymenopterous allies of the bees who nested in
the skeleton of Samson's lion burrow and build a home
among the bones of extinct creatures of the geologic
These roofing pebbles are not i.or but sparsely) inter-
mingled with the soil of which the interior bulk of the
cone is composed, but form a stone covering, or roof, about
a half inch thick, more or le . Mr. H. L. Viereck in-
formed me that he had seen bits of cinder and coal,
evidently gathered from the railroad track, for roofing.
This is confirmed by the statement of Mr. G. A. Dean
[D. 1, p. 1(59]. who further says that on the old town
site of Wallace. Kansas, they used bits of glass, mortar,
and small fragments of rusted iron from the debris of
ruined house-. Thus the roofing habit, though it may
have originated from the accidental deposit of ex-
cavated pebblets. seems to have grown into a fixed
purposeful instinct that prompts to gather supple-
mentary material from any available quarter.
The pebbles are handled with ease by the worker-
ants, who nip them with their outstretched mandibles
and then move off. rarely stopping en route to adjust
the burden or to rest. The body is lifted up. the head
well elevated to prevent bumping against the surface,
and the load held well to the front or somewhat beneath
the body. The portage was amply observed during
ordinary excavations, in opening and closing gates, and
in repairing breaks caused by rains or purposely made
for experiment. In the last-named work the ants
would descend to the clearing at the base of the cone,
and carry the stones up the slope with as little apparent
effort as when moving downward.
This, however, must be an easier task than transport-
ing them from distant sites or from their interior beds
up the galleries to the surface. The space traversed in
this underground portagf- is sometimes equal to a per-
pendicular distance of nine feet, which has little me-
chanical relief from the inclination or roughness of the
gangways, sfome of the pebbles have from six to ten
times the weight of their carriers. I never saw anv
copartnerships in these port;._ s. No ant came to aid
a struggling worker, and none seemed to need assistance.
I have often admired the vigor and skill shown by
baggage-porters in shouldering and bearing up several
flights of stairs the immense trunks which American
ladies take with them on their travels. But here, if
we may be indulged in the comparison, is an insect
three-eighths of an inch long and the worker-minors are
shorter . who can cany up sharp inclines and perpen-
dicular surfaces, over a distance three-hundred times
its length, a burden six to ten times its weight. If. as
heretofore, we estimate the average man at five and a
half feet in length and one hundred and fifty pounds
in weight, our baggage-porter would needs carry a half-
ton trunk up one-tenth of a mile of stairway, to meet on
equal footing the emmet athletes of the Occident ant-
The simplest type of ant architecture, as we have
seen, is a single cave excavated in the earth, or in wood,
or formed bv detritus cemented bv salivarv secretions.
This grows into (second) an enlarged chamber or cham-
bers, with vestibule and connecting galleries. Thenre
third) developing downward, the simple cave or con-
nected chambers have grown into vast and deep-storied
rooms and avenues, like those of the agricultural. Oc-
cident, honey, and cutting ant-
Expanding in the opposite direction a development
upward instead of downward fourth) the little heaps
of earth-pell . _ 1 thrown out around the gate of
the cavern home of Pheidole. or the garden Lasius, be-
come (fifth) the great conical structures of the mound-
iking ante of the Afleghanies. which are in themsel
ii ::.:. : . : :> This 15 an important difference. 7
3 are thoroughly honeycombed with avenues and
.nd galleries, which are the actual Kving-
ters of the commune, and form, each mound in it-
-. . --
_ _ -
**= -: &
Flff. 21 f -TTFl.lt H " - 7 _ ABTS-PEIXETS THBOWN'
-_:- rr-iz - -. T j. - . -
: i z ~ r :
self, into a densely populated city although in full affi-
ance of citizenship with all like mounds in the vicina^
We come nc vpe c: -:: jcture which
char^ rues a number of genera in Europe and America,
but which is particularly developed in varic - spe
and varieties of Cremastogs-Trr. 7 - eeiea of t:
genus are small, and are widely distributed throughout
:errl: They have a hean-shaped abdomen or
gaster. flat above and rounded below, and this th
have the odd habit of turning up and directing forward
(Fig. 22), so that it is almost parallel with the line of the
thorax. [McC. 9, p. 188.]
These ants, besides nesting in the earth near the
surface and under stones, are apt to choose a site in a
heap of stones, on an
old stump, or in the
debris of fallen and de-
caying logs. A colo-
ny settled among the
crannies of a bowlder
wall at our country
built a covered ap-
proach to their main
entrance, using there-
for particles of dust,
earth, etc., that had
accumulated upon the
rocks. The nest itself
was within the inter-
stices formed by the
rounded exteriors of
the big bowlders, and
was quite out of sight.
This covered vestibule
was a mild suggestion
of the vast mud-cov-
Fig. 22 CREMASTOGASTER LiNEOLATA 6i*ed ways made by
a, b, c Worker, d, e Queen. the Eciton, or " driv-
/ Worker with turned-up gnster.
their route when out upon one of their devastating
On a vine twined about the cloistered porch connect-
(By courtesy of the American Museum of Natural History)
Fig. 23 CARTON COCCID-TENT OF CREMASTOGASTER LINEOLATA-
PILOSA PERGANDE. (After Wheeler)
ing my city manse with the church, I have noted similar
enclosures built around the stock at points where a
branch diverged therefrom. They were apparently
wrought out of ''carton' -that is, a combination of
wood-dust, loose earth, minute particles of straw, hay,
and horse-feed (the droppings of passing animals), such
as drift from the street into corners and crevices of walls
and upon the foliage of city plants (Fig. 23). This
material had been mixed into a sort of mortar and fixed
by a natural cement secreted by the ants, until it formed
a woody composite that easily crumbled between the
fingers, but held together well enough to answer its
purpose as a temporary tent. Structures of a similar
character have been observed around the bases of the
needle-like leaves of pine-trees in New Jersey and else-
To what purpose are these rude shelters made?
Chiefly to obtain exclusive and undisturbed possession
of aphides and other insects that excrete the honey-
dew of which they are so fond. [\V. 9, pp. 1-18, Plate
ii. copied.] This enclosure not only serves to restrain
the insect herds from wandering to inconvenient sites,
but shuts out alien ants that, when strolling about, as
is their wont, in search of food, might happen upon
the pre-empted aphidian flock, and encroach thereon.
In short, it is the Cremastogaster's way of " staking her
It is interesting to notice that this tendency to build
carton tents exists in this species not in full vigor as a
thoroughly developed and fixed habit, but as a sub-
sidiary tendency, a survival, it may be, of an ancestral
habit once strong and persistent, but which, in course
of time, has been gradually weakened and well-nigh
K - ' ' - / .:
Fig. 24 PASTEBOARD NEST OF DOLICHODERUS BTTUBERCULATTJS
MAYR, FROM BANKOK
atrophied. Still, under the stimulus of special favoring
conditions, the latent ancestral tendency springs into
active force, and shows itself in the occasional and
temporary structures above described.
I have alluded to this phase of ant industry and vent-
ured on the suggestion of its origin, not simply because
of the intrinsic interest of the facts, but because they
lead up to what is (seventh) a distinct and well-defined
form of nesting architecture. Not only Cremastogaster,
but other genera have acquired the habit of making
carton nests in and upon trees. They are at times quite
large (a foot or more in diameter), and they form true
habitations, as are the nests of hornets, in which larvae
are reared, dependents housed, and all the functions
of an ant commune carried on.
These tree-dwellers are for the most part habitants of
tropical and subtropical countries, and the accounts of
travellers give one a vivid conception of their power,
when excited by intended or accidental aggression, to
swarm forth in legions from their domiciles and punish
invaders with stings that seem pointed with fire.
Of the ants of this form of arboreal nest, Fig. 24 is
taken from Dr. Forel's Ants' Nests, and shows a paste-
board nest of Dolichoderus bituberculatus Mayr, taken
from the bough of a tree from Bankok, Siam. Fig. 25
is a tree-nest after a figure published by Dr. von Ihring
[Yon 1. 1, p. 334] of Camponotus rufipes, of South America.
We come now to note (eighth) the existence of nests
for which the framers have called in the use of silk.
That this should turn up at even the most unexpected
points in the insect world will not seem strange to one
who knows how largely the spinning habit enters there-
into. Many ants in their larval forms follow the role
Fig. 25 NEST OF CAMPONOTUS RUFIPES ON A TREE
(After Von Hiring)
of other orders, and spin themselves within their tough
pupa cases or cocoons. Thus we are quite prepared to
learn of Professor Forel's Ceylon species (Polyrachis
jerdonii), which builds upon leaves a small nest com-
posed of pebbles and minute fragments of plants,
cemented together by a fine web, or woven together to
form a web-like wall of bright grayish brown. In the
East Indian ant Polyrachis dives the nest wall is a pure
silken web of a brownish yellow, by which the enclosing
leaves are lined and bound together.
Polyrachis spinigera, of Poonah, India, makes for its
nest a fine brown silk web pliable as the finest gauze, but
thicker. This is fixed on the ground, where it forms the
lining of a funnel-shaped cave that widens out into a
But among the woven ant -nests thus far made
known the one that seems to show the highest type
of nidification is that of Mcophylla smaragdina, a com-
mon ant of tropical Asia and Africa. The workers are
large, long, reddish to greenish in color, of a war-like
and fiery temper. Their females are grass-green, their
males black (a rather striking color combination), and
they maintain populous communes among the branches
of trees. The common habitation is formed by joining
together the borders of leaves with white spinning-work
and binding them into a large mass, something after the
fashion of certain spiders and tent-caterpillars.
According to Mr. Aitken [Ai. 1, p. 422], the method of
construction is as follows: A worker stands at the point
where two adjoining leaves diverge, and holding to one
with its claws, seizes the other with its jaws and draws
the two together. As the leaves gradually approach each
other they are held in place by the outspun threads,
until, as the leaves touch and overlap, they are overspun
and firmly bound down. And so the work passes from
leaf to leaf until a sufficient housing is provided. This
process in essential method resembles that which I have
studied among spiders, particularly those large native
orb-weavers, like Epdra insularis and E. domiciliorum,
that build as a domicile large leaf-tents above their
great geometric orbwebs.
It seems almost past belief as a bit of natural archi-
tectural ingenuity, but the observer, whom Professor
Forel thinks entitled to credit, states that if in clamping
the leaves the space separating the edges be overwide
for one ant to manage, other workers, from two to five
in number, will "join hands" to form a chain, each
grasping the body of its neighbor until the last link on
one side holds a leaf in its mandibles, while the last link
on the other side grasps a leaf with its claws. Thus, all
drawing together, the chain is gradually shortened until
the breach is closed and spun over. When enough
leaves have been fastened together, the whole is over-
spun with a compact silk web, and made water-proof.
It is then divided into connected living-rooms as
Many ants are opportunists in the choice of a habitat.
Instead of working out a nest in earth or wood, they
exercise a sort of "squatter sovereignty/' and preempt
for use some available locality. For example, I found
a large commune of Aphenogaster tennesseensis in an old
pine-tree stump at Bellwood, Pennsylvania. Moss
and lichens grew freely upon the stump and its great
bare roots. In the scant soil that had formed upon
the top sundry wild plants were growing, as in a roof-
garden. At various places over the surface large, dry
toadstools were attached, looking like ancient sea-
shells. Within these the ants had settled. The in-
teriors swarmed with workers and larvae; and, as evi-
dence that the task of enlargement was going on inside
Fig. 26 NEST OF PSEUDOMYRMA BELTI ON MEXICAN ACACIA
of both toadstools and stump, the outer bark of one
large root was covered with fresh wood-dust.
In tropical and subtropical countries examples of this
habit are common on certain thorn-bearing trees and
shrubs, as the Acacia. These thorns
are often quite formidable in size.
I have had specimens of them which
had been inhabited bv ants of the
widely spread genus Pseudomyrma.
Near the points circular gates were
cut out for entrance and exit, and
the commune was established in the
hollow interior (Figs. 26 and 27).
It is believed that some ant-inhabited plants, as
Cecropia adenopus, furnish in themselves a supply of
Fig. 27 PSEUDOMYR-
MA BELTI (EMERY)
food in small pyriform or ovate albuminous growths
(Midler's corpuscles), which are eaten by the insects, ap-
parently without disadvantage to the tree. They present
another example of that interesting form of symbiosis-
a comradeship of support and defence which not infre-
quently exists between plants and insects. 1 For in
return for the nurture yielded by the plant, the ants
protect them from the incursions of such enemies as
the leaf-cutting Attidie, that defoliate them in their leaf-
Our studies of what, for lack of a fitter name, has been
called the "architectural' and " engineering'' skill of
ants will now return for a while to the great confederacy
of the communes of mound-making ants. Let us note
the citizens of this large colony of Formica exsedoides,
who are adding a story to their communal cone. It is a
lively and interesting scene. The utmost activity pre-
vails, greatly quickened by a smart shower which has
made the building material more available for use; for
hot, dry weather had left the soil powdery and un-
We soon get an insight of their mode of operating.
On the outer surface, especially around the gates, small
warts or pillars are thrown up, as though to gauge the
height of the addition; and between these the infilling
is made with pellets of earth, most of which are brought
from the interior, where new galleries and rooms are
being excavated. The vehicles for this portage are the
insects' jaws, that serve alike for carriage, as basket or
barrow, and for digging, as pick and shovel. The irreg-
ular faces of these pellets fit into and fasten upon one
1 See Prof. A. F. W. Schimper's The Varying Relations Between
Plants and Ants, Jena, 1888.
another, uniting the whole in a way that, perhaps, may
not be characterized as " do vetailing " or " ball-and-
socket" jointing, but which raises the suggestion of
It is said that formic acid, which is extruded freely
by ants, forms with silica a natural cement. Can it be
that these pellets, which are composed largely of sand,
are thus cemented together? Probably not; but at least
the moisture of the late rain has aided their adhesion.
Grass straws, cut from tufts growing along the base and
strewn over the summit of the cone, are dragged into place
and skilfully wrought in with the pellets. Besides these,
bits of decayed wood, the needle-like leaves of pine-trees,
which are abundant here, and leaves of low shrubs are
intermixed with the soil. These insect masons are not
forced to the hard service laid by the Pharaoh upon
ancient Israelites of making bricks without straw.
At another hill the builders had undertaken a special
work of construction, or, rather, of repair. An errant
cow grazing on the mountain-slope we can hear the
tinkle of her bell from a distant corner of the wood-
had strolled by that way and set a hoof upon the edge
of the mound, leaving a deep and wide impression. Just
one; for a swarm of irate insects must have instantly
attacked Brindle's legs and caused her to beat a rapid
retreat. But she has left a footprint on the cone that
must needs be repaired. As a full day's work is before
us, let us bring camp-stool, note-book, and drawing-pad,
and sit down before it. We shall see something that
looks like a well-planned system of engineering in filling
up the hole.
The cow's foot had made a nearly circular pit between
eight and nine inches in diameter and depth. At the
unbroken edge of the pit the task of filling up had begun.
From the lower points (A) there extended a series of
elevations (a, b, c, d), which marked the upper outline
of an arc. Beyond this, toward the base of the hill,
and parallel with it, was a like series (d, e, /), bending
around the depression next to and parallel with its
Fig. 28 PRELIMINARY ENGINEERING
Mound-making ants filling up a break made in a mound
by a cow's foot
lower rim. These little pillars were not alike in shape,
and it was quite noticeable that their height gradually
increased from the unbroken margin of the mound
toward the centre, where it was highest. Yet, through-
out, their tops conformed to the general surface of the
cone, the diminishing depth from centre to edge being
met by a corresponding lowering of the columns. From
these, and from similar elevations around the rim of the
track, as centres of operation, the work of covering-in
proceeded with great advantage (Fig. 28). A number
of straws were worked into the columns apparently as
braces ; and in one of the little hollows were piled many
shells of cocoons from which antlings had just been
delivered, and which previously had been carried from
the hill and dumped among the stones outside. Within
three days, so vigorously did the work go on, two-thirds
of the track had been filled up. The new work did not
connect with the broken parts on the side toward the
summit, but a deep trench, or open gallery, was there
preserved all the way across. This seemed strange
until it was considered that it was needful, or at least
convenient, to keep such an opening into the network
interior of galleries in order to allow easy entrance to
and exit from the works within the track.
Could this systematic order of work, with such plain
marks of an intelligent plan, and carried forward much
after the manner of men, in their like though larger
undertakings, have come about by mere chance? It
does not seem reasonable to think so. Yet there was
no trace in any quarter or in any act of chief-engineer,
or local foreman, or gang-boss, or of. any visible or-
ganized directing body, or official supervisor, or regulator,
or prompter. Every individual had a mind to work,
and every one wrought, but unprompted and un-
governed save by its own impulse.
Here, also, the observer was impressed by the presence
of that invisible, secret, mysterious Something which
he has called the Spirit of the Commune bevond the
veil of science and philosophy, which kept all these
active sovereign integers in such harmonious co-opera-
tion in the execution of a fixed plan, sustained without
intermission for half a week. Is analogous action
possible among men? And if not, why not? And
wherein lies the superiority if it be superiority in
this respect of ants over men?
Everywhere we note examples of this co-operating
instinct of construction. In order to get a view of the
interior of an ant city a mound was sawed through the
centre with a large cross-cut saw and one-half thrown
aside with shovels. This required vigorous and rapid
movements to avoid the attack of the myriad of angry
insects thus assaulted in their home. This, however,
was less formidable, because the calamity was so unique
and terrible like the earthquake shocks which lately
wrecked Charleston and San Francisco, Messina and
Reggio that the ants at first seemed stunned, and
moved about as though distraught. Such an over-
throw was beyond their limited powers to grasp. But
they soon rallied, and promptly set themselves to restore
their ruined commonwealth. Yet the blaze of passion
was hot enough on the part of those who swarmed from
the quarters untouched by the shovellers. Fortunately,
the heat and fury thereof were soon expended.
Let us examine the interior of the mound thus laid
bare. The view of the perpendicular face of the half-
cone exposed was truly remarkable (Fig. 29). Tubular
galleries three-eighths to half an inch wide rose in regular
series one above another, from the base to the domed
summit. The cone within was a rough reproduction in
soil, and on a gigantic scale, of the celled structure of the
combs of bees, wasps, and hornets.
Throughout this network of galleries were scattered
cavernous rooms, the common lodging-places for the
young and other dependents, although the galleries
also served this end, as well as being the roadways be-
tween all parts of the community. It is an amazing
structure for so small a creatureling, and must have re-
quired immense labor and pains to rear it. That this
huge hill of sandy earth, mixed with and fixed by vege-
table 1 matter, and perforated from top to bottom and
from side to side with numberless tubes, could be made
to stand the stress of mountain weather rains and
floods, frosts and snows was marvellous. The inference
' - Jl
s> - "=:- ' " - *V^4s^ r .,^trv,>x ^^O'-sfl.M
Fig. 29 SECTIONAL VIEW OF ANT-MOUNDS
seems inevitable that some sort of hardening material
must have been used secreted from the jaws as saliva,
or extruded from the stinging organs at least upon
the inner surface of the galleries. But, however done,
the ants had successfully solved their problem of build-
ing a stable structure with friable materials, and had
wrought it into a true communal home.
To further learn their method, let us fix attention
upon one point. A small cavity with a bit of projecting
clod on the foundation of the removed part, and close
to the remaining one, was the centre of active operations.
Just above towered the perpendicular face of the half-
cone, along which everywhere squads of workers were
Fig. 30 AXT ENGINEERING
Covering-in a broken horizontal gallery and repairing a broken
continually thrusting their brown heads out of gallery
openings and dropping pellets of soil. They were clear-
ing up the inner debris from the broken passageways,
and doubtless some of them were calmly carrying
forward the improvements begun before the Dearth-
quake ' (Fig. 30) . These pellets were taken up by the
workers beneath, who for some reason best known to
themselves seemed to prefer them to those that lay
everywhere around them, the crumbled particles of their
.shattered home. Two galleries running side by side,
the upper parts of which had been destroyed, were being
covered over. They were directed along the bottom
of the cavity for three inches, and then slanted up-
ward to connect with the standing half-cone.
The work progressed by continuously adding earth-
pellets to the outer edges and pressing them into place.
As the sides rose they were gradually arched, and the
springing of the arch was plainly seen. The curved
edges approached in irregular lines, and at various spots
the two projecting points drew near and nearer until
they almost touched. It was quite exciting now to
watch the delicate manipulation of the masons. Here
came a worker with a pellet of larger size than usual.
She climbed the arch, moving more daintily as the top
was reached. Holding on the while with her hind
feet, she stretched across the wee chasm and dropped
the ball of soil into the breach. The bridge was
And now, with surprising rapidity, it widened as the
roof of the arch was covered. Until this was done,
openings were left through which the ants moved back
and forth, and which were closed over as sections of the
arch were completed. They were temporary arrange-
ments " manholes," so to speak for the convenience
of the builders. Through these one could see the ants
at work upon the inner surface, smoothing it with their
jaws, as a mason would work with his trowel and
mortar. The outside of galleries and rooms was left
rough, as laid, but the interior was smoothed. Salivary
secretions probably gave the additional moisture needed
for this. At one point the gallery was widened from
half an inch, the usual diameter, to one and a half
inch, as though a store-room or living-room were being
formed. Close by, a vertical gallery, one side of which
had been torn away, was being repaired by the infilling
of the broken side, and this work was done precisely as
in the case of the horizontal arches.
One was reminded, in all these actions, of the methods
of bricklayers at work upon an arched sewer or culvert,
or of masons putting up a rubble-stone wall. There
were some marked differences the profound silence of
the worker ants and the absence of overseers. The
ant is no "spendthrift of her tongue." She "talks to us
in silence." No one is prompted or driven to work;
no one needs to be, for here there are no shirks. No
regular hours of service are kept, and there are no fixed
intervals of rest. Labor goes on all the time; and, view-
ed in the mass, there is no cessation, at least at this
juncture, by day or night. Each individual determines
for herself the period of work and the time for rest, and
so strong is the sense of duty, or the instinct of fidelity,
in every ant, that such individual liberty and respon-
sibility are not abused, and the public works of the
commonwealth are not damaged or delayed.
Building operations were not limited, as in the above
cases, to the original site of the cone. A fragment half
the size of one's head, which had been shovelled to one
side, was a centre of special activity. It had already
been made the nucleus of a new mound. Columns,
corridors, and halls, corresponding closely with those out-
lined upon the under side of the fragment and united
therewith, had been erected. In one of these halls was
a small collection of dead ants, a token of a custom
sometimes observed among these insects to show a sort
of funereal respect to the dead of their own household
This was one of the most interesting and puzzling of
the activities developed by the cutting-down of the
mound. Access to the uninjured part was easy, and
Fig. 31 MOUND-MAKING ANTS COVERING A DOUBLE GALLERY
a, a, a Double gallery, c, c, c Chambers.
knowing the habits of these Fonnicans and their wide
range of daily venture, it seemed strange that they did
not at once, as did so many of their fellows, rally to the
reconstruction work on the old foundation. Had they
been found huddled impassively underneath the scat-
tered fragments, keeping refuge for the nonce and wait-
ing results, it would have seemed natural. But this
immediate launching upon an apparently fresh enter-
prise turning to housemaking instead of home-repair-
ing, so near the gates of the old republic beginning the
upbuilding of a new this mystified a mere human brain.
Did the familiar savor and associations of a bare lump
of their former home deceive them ? Were they so little
impressed by the commune's partial wrecking as to
think that nothing serious had happened? Was this
an instinctive act of self-protection, for the present
exigency alone, a sort of bivouac and makeshift, like
the temporary camps around a despoiled city? Would
these adventurers persist, and build up a permanent seat,
or soon return to the old quarters? Was this a proof of
superior wisdom, the act of keen opportunists quickly
adapting themselves to strange conditions, or of faculties
far more limited than we have been wont to credit to
ants? Was it simply the result of a physical necessity
to be doing, an uncontrollable impulse finding vent in
But here we stand, vainly speculating and philoso-
phizing, while the field of observation just before us is
alive with busy insects who have much to show us. Let
us go back to the practical and objective. We have
seen something of the way in which the citizens of an
ant community labor in mass on their public buildings
and roads. The manner of the individual now requires
to be noted. And here comes a worker-minor who will
answer our quest. She has strolled along over the
irregular surface of the old foundation of her home,
feeling with her antenrue here and there without any
visible purpose. At last she pauses. She seems to be
reflecting upon the ruin around her and without apparent
emotion. Ah ! if we could but command the mediation
of some fairy interpreter of her thoughts !
But see! Suddenly she leaps upon a pile of earth-
crumbs, and, seizing one in her jaws, lays it down at a
little distance with a sharp pat. Another and an-
other and many others follow, all gathered and placed
with amazing activity. The little body, from the tips
of her ever-moving antemue to the apex of the abdomen,
quivers with the intensity of her energy. She reminds
one of a small harbor tug, forging ahead, trembling from
stem to stern under her great engine as she draws in
her wake a huge ship. Only, there is no puffing! All
goes on "in solemn silence, " like the shining orbs in "the
spangled heavens/' as sung in Addison's paraphrase of
the nineteenth Psalm.
Will the day ever come when even a remote approach
to this noiseless toil shall characterize human communes ?
To be sure, since men's work is so largely wrought by
the aid of machines, the racket thereof and the audible
strain of their motors cannot be wholly subdued. But
it is certain that some of our mightiest and most effective
machinery does its work with the minimum of friction,
and so of noise. Really a mechanical contrivance might
be fairly counted valuable in proportion to the silence
of its operation. Always force is wasted in noise-
physical and mental as well. Silence is a mighty
economist of man's wealthiest powers. Racket is a
Beyond doubt, at least, abatement in large degree of
our city noises is not only desirable, but wholly prac-
ticable. They are needless. They are wasteful. They
are often cruel to the invalid and nerve-worn. They
are remainders of a crude stage of development, and
an enlightened people should not tolerate them. They
are chiefly the products of thoughtlessness, stupidity,
penuriousness, unthrift, and a selfish rudeness. Here
also our city magnates and lords of industry might go to
the ant and consider her ways with profit.
ENGINEERING METHODS IN ANT STRUCTURES
A TJ"E return to our lone pioneer laborer of the mound-
T T making ants, left, at the close of the last chapter,
in the act of beginning a work of repair upon her desolat-
ed commune. Her movements will give us an insight
of some emmet methods in this field of engineering
construction. One soon begins to see some purpose in
her work, for slowly the suggested outline of a gallery
takes shape. Meanwhile a second ant has wandered
that way. She halts and, with \vhat appears a. careless
mien, surveys the scene. Then, struck by an impulse
that probably is as mysterious in its origin to her as to
her observer, she joins the first adventurer in her attack
upon the pile of earth-pellets and in their transfer to
the growing gallery. By a like process the squad of
workers increases from two to four, from four to ten,
from ten to fifty or more, until a busy company swarms
over the works, which are rapidly taking distinct form
as an arched gallery.
The pioneer of this enterprise has long ago been lost to
sight among her comrades, and one regrets the lack of
brush and white paint wherewith he might have marked
the black abdomen, and thus have kept track of her.
It is certain, however, that the fact of her having been
the first citizen and founder of that settlement had
given her no claim to authority or superiority of any
sort. Her fellow-citizens seemed ignorant of the fact,
and it is doubtful if she remembered it herself. Such
sentimental considerations have no weight in this wholly
utilitarian government and society.
Perhaps this is she who slips out of the throng, and,
ascending a bit of a clod hard by, squats upon her hind
legs and begins to preen her downy coating. What to
her now is all the busv scene beneath her? Let the
emmet world wag on as it will; she must be clean.
And so, in peaceful unconsciousness of all and sundry in
her sphere of being, she proceeds with the one present
purpose of life, and thoroughly cleanses herself in the
approved mode, as hereafter will be described. And now,
her purifications being finished, she yawns, stretches
her limbs, gives her antennre a final brush, and
leisurely descends from her perch. A moment she
stands as though undecided, then plunges again into the
whirl of activity on and around the new works, and
soon is indistinguishable from her fellows.
In the review of these building operations several
reflections arise. Here was a test of the ability of ants
to meet a new experience, such as was the destruction
of one-half of their republic. Accidents like the breach
made by a cow's foot they had known and remedied,
but no such misfortune as then faced them had ever
befallen. Yet they met it with admirable spirit and
method, and with success. They set themselves at once
to the work of reconstruction, not only with vigor but
with practical wisdom, and with ready adaptation of
means to the new conditions. Their instinct was
sufficiently elastic to cover a strange and colossal ad-
versity; or, may we infer that the appeal in the
emergency was to something other than routine instinct
-something, in fact, that nearly approached and cer-
tainly suggested a process of adapting means to ends,
that bore the earmarks of reasoning?
Again, their swift and perfect reaction from the first
shock and excitement of a disaster that well might have
overwhelmed ambition and endeavor was noteworthy.
They went straight on with the ordinary duties of life
in the uninjured part of their city, and took up the ex-
traordinary ones without a sign, understandable by
human intelligence, of grief, or passion, or discourage-
ment, or deep emotion. No time was wasted in useless
moping, no vigor in aimless schemes. At once they
aroused themselves to action, and attacked the emer-
gency with admirable energy and poise. Among men
such self-control has been called the fruit of philosophy.
If such it be, will we be able to deny our mound-making
ants the title of insect philosopher? Certainly they are
apt are they automatic? imitators of the philosopher's
role. And not to a few rare spirits, the sages of the
commune, but to all and equally is the honor due.
Nature maintains in the ant city Voltaire's ideal con-
dition that a philosopher should live only among
A curious observation as to how active work may
affect the physical condition of ants was made upon
three mounds listed for daily special study. For
several days they were found in such an unusually
excited condition that they were down in my note-book
as the "hysterical hills." The cause of this agitation
evaded all inquiry, but the cure was most interesting.
Rains that succeeded the first dry days of our coming
were observed to have imparted activity in building
operations to a number of mounds. The prediction
was thereupon ventured to one of our company that we
should find our "hysterical hills'' busily building up
their cones like their fellows of other mounds, and as a
result settled into their normal composure. And so we
found it. They were working at the top of their bent,
and were subdued in temper and manner. Honest,
hearty physical toil had quieted them, as it often does
over-nervous human beings; or, perhaps it had filled
their natures with a present and pressing duty, thus
diverting them from that useless expenditure of force
that often comes from purposeless inaction.
One must also note the immensity of the labors
wrought by the insects. These may seem trivial as one
watches them lifting up and placing here a pellet and
there a pellet of soil, and building them into the walls
of the common structure. But if the results be con-
sidered, they will seem astonishing for such small creat-
ures to accomplish. Perhaps a comparison with a noted
building achievement of our race, the great pyramid of
Egypt, may here be allowed. It is true that such com-
parisons are apt to be superficial and misleading, but
from a purely popular standpoint the}' are allowable
and may be instructive. The cubic contents of one of
the largest mounds was calculated to be in round num-
bers two million cubic inches. We may estimate the
bulk of an ant to be equal to that of a cylinder three-
eighths of an inch high and one-sixteenth of an inch in
diameter. Taking thirty-five one-hundred-thousandths
of a cubic inch as the bulk of a single worker ant, the
size of the builder would be to the size of the edifice as
one to fifty-eight hundred million. Let us compare
this with a corresponding estimate of the work of man,
taking his bulk as six cubic feet, and accepting the solid
contents of the great pyramid as reckoned at seventy-
six million cubic feet. We shall have the following
formula of comparison:
Man's bulk to his building is as 1 to 121 millions;
The ant's bulk to her building is as 1 to 5800 millions.
A simple calculation will show how greatly this ex-
hibits the comparative superiority of the insect in the
mere quantitative re-
sults. It should also
be considered that in
these and all other
such works the ants
have no mechanical ap-
pliances such as mul-
tiply the effect and
ease the fatigue of hu-
man labors. Her sole
tools are her jaws and
feet (Figs. 32, 33),
the bodily appendages
with which nature has
provided her, although
it must be admitted
that these are highly Fig. 32 ANIS DIGGING OUT GALLERIES
effective. (Drawn from an artificial formicary)
There is another c Claws, d Particles of dirt,
comparison --in sooth p Point being dug out.
it is a contrast -
which needs here to be drawn. Such knowledge as
has come to us of the building methods obtaining in
ancient Egypt shows that the laborers were driven to
their hard tasks by overseers who urged on gangs of
workmen with the lash. Theirs was unwilling service,
Fig. 33 AX AGRICULTURAL ANT*S FOOT, ENLARGED 180 DIAM-
ETERS! THE TOOL USED, WITH THE JAWS, IN DIGGING
a c I Attached claw, a t Its articulation with the joint.
i c I Inserted claw, i n The inserted parts, s r Ser-
rations on same. / m Foot muscle, p v Pulvillus or
cruelly exacted. We have already seen that the ants
rendered free and willing service, and that their toil was
without overseers, and wholly of individual selection.
From the beginning to the end there was no discord
among them; no protests; no strikes, sympathetic or
otherwise; no walking delegates or their insect analogues;
no oppressing (or oppressed) contractors or owners.
Indeed, there was no occasion for any of these frequent
appendages of great modern structures whereon human
workingmen artisans, mechanics, and common laborers
And yet the work was done, and on undertakings
relatively many times greater, in the most perfect har-
mony, good temper and content of all. Is it possible
for man to draw some lessons from this example of
natural civics? Is it beyond hope that some goodly
measure of such results may lie within the sphere of the
practicable for our current organized society? Does
our "civilization' 1 hopelessly encumber us from ever
attaining the ideal commune? Must it lie in the bright
cloud-realm of the optimist's dreams, until alas! can
it ever be?> the whole race, reborn and disenthralled,
shall return to the unsullied simplicity of nature?
It may be drawing too fine a distinction in the build-
ing work of ants to discriminate between architecture
and engineering. Yet we seem to note such a distinc-
tion. The commune of the agricultural ant, already
described, is differenced from its fellows by the circular
disk (Fig. 34) that surrounds its central gate or en-
trance. The construction of this disk, and its main-
tenance as a free and open plaza in the midst of the
surrounding subtropical foliage, are works for which no
little skill and energy are required (Fig. 35).
Fig. 34 THE CIRCULAR DISK AND ROADS THAT SURROUND
THE ONE CENTRAL GATE OF THE AGRICULTURAL
ANT OF TEXAS
But this is only the beginning of the enterprise, the
pivot upon which more important undertakings centre.
At various points around the circumference of the disk
enter a series of cleared trails, widest at their point of
contact, that radiate into the surrounding herbage,
whose denseness at once suggests the reason for their
construction. In short, they are roads laid out to
penetrate the harvest fields of these granivorous ants,
and are used and admirably adapted for that purpose.
The method of transportation in use by harvesting
and other ants is primitive enough, consisting simply in
personal carriage by a host of individuals (Figs. 30, 37,
38). It is the method of the African explorer, the
method of primitive man when unable to utilize the
beast of burden. But it is effective. It is here that
the engineering quality of the roadways comes in; they
converge upon the entrepot of the colony. They facili-
tate transportation by making communication easier and
quicker. However they may have arisen in the history
of the harvesting habit's development, or with what pur-
pose (if any) originally constructed, the facts are as stated.
And they do not stand alone. In the summer of
Fig. 35 AGRICULTURAL ANTS CUTTING GRAIN TO CLEAR
ROADS AND DISKS
Fig. 36 ANT CARRYING A PELLET OF SOIL
Fig. 37 A SPECIMEN OF GRAIN-STALK ON AN ANT CLEARING
Fig. 38 AGRICULTURAL ANT CARRYING GRAIN-STALKS
1887, while visiting Scotland, I spent two days at the
Trossachs Hotel, which is located in the glen known as
the "Pass of Achray,' 1 through which flows the little
Achray river (or creek) which Sir Walter Scott describes
as "the stream that joins Loch Katrine to Achray."
Here I found a number of nests of Formica rufa, the well-
known "horse ant," or "wood ant," of Great Britain.
They were built on either side of the footwalk that leads
from the Trossachs glen to "the sluices," as they are
popularly called, which regulate the stage of water in
Loch Katrine, the source of supply for the city of Glas-
gow. The demands of humanity are imperative; but
the lovers of romance cannot but mourn that the spell
which Scott's genius has thrown over such beautiful
spots as "Ellen's Isle" and the "Silver Strand" is
being dissolved before the engineer's need for greater
The mounds raised by the rufous ants were found to
be cones of earth intermingled with chippage of various
sorts. They were about three feet high, and some of
them from six to seven feet in diameter across the base.
They resembled those of our mound-making ants of the
Alleghanies in general appearance, but their builders
seemed to make a freer use of leafage and chippage to
work up and cover their nests. The surface was quite
thickly thatched with bits of straw and leaves, stalks of
grass, pieces of fern, and various like materials. Num-
bers of openings appeared upon the surface at irregular
intervals from the summit to the base, and at 4 P.M.
many workers were dragging the chippage back and forth,
as though arranging to close the doors for the night.
[McC. 23, p. 336.]
These huge cones stand in the midst of the tall
bracken a large, coarse fern that overhangs them, and
at times almost hides them from passers-by, as a forest
might hide a castle standing in its midst.
My attention was especially attracted to the character
of the roads leading from the ant-hills to various points
in the surrounding wood. These were distinctly marked
upon the surface of the ground, having in places a width
of from two to four inches stained dark brown or black
by the formic acid exuded from the insects as they
passed along. The fallen leaves and crushed grass
upon which the trails were made were pressed down
and smoothed by the constant action of innumerable
legs upon the surface. So well marked were these
trails that they were easily traced even without the
presence of the columns of ants that marched back and
forth upon them.
While following up one of these roads, I was struck
by the fact that it showed scarcely any deviation from
a straight line. In order to test this matter more care-
fully, I selected a large mound from which three roads
radiated. These were all traced to their termination at
three several oak-trees, up which the columns of ants
ascended to obtain the honey-dew supplied by numerous
aphides that infested the branches. The roads were
carefully marked out by stakes set at short intervals,
a plan made necessary by the high bracken, whose stalks
stood so closely together that they had to be pushed aside
to trace the roads.
The following facts resulted; Road No. 1 was sixty-
five feet in an almost perfectly straight line from the
nest to the tree. Road No. 2 was seventv feet long and
varied less than three inches from a direct line drawn
from the nest to a point within two feet of the terminal
tree. There the column made a detour of about six
inches. But an abandoned path continuous with the
main road, which apparently had been used at a recent
date, was traced for a considerable distance farther.
Road No. 3 was the longest, being more than a hun-
dred feet long. It extended for nearly twenty feet in a
straight line, at which point it touched an old stump
that deflected it at a slight angle. Thence it was
continued in a nearly direct course as far as a beaten
footpath through the wood. Here the ant trail was oblit-
erated by passing human feet, although the ants still
thronged the pathway, there much broadened by the con-
tinual interference and loss caused by foot-passengers.
The trail, however, was resumed at a point nearly oppo-
site that at which it touched the path, and was con-
tinued again in a straight line about twenty feet farther
to the tree where it ended.
When the entire trail (No. 3) was staked off, it was
found that its terminus deviated less than three feet from
the straight line drawn from the point of departure at the
ant-hill. The greater deviation in this case was doubt-
less caused by the peculiar difficulties in the track.
The three roads so radiated from the parent nest that
they were included within about one quadrant of a
circle, of which the two shorter trails might represent
radial boundaries of the quadrant, while the longer
trail was midway between the two.
Looking simply at the results of these observations, it
is manifest that these rufous ants showed an accurate
sense of direction in marking out and following their
approaches to the trees. It would not be reasonable to
attribute such mathematical accuracy as above shown
to mere accident. The roads in point of directness were
as well laid out as are works of a corresponding nature
done by the engineering skill of men. And these are
not isolated cases, but mark the general rule. The
mound-making ants (Formica exsectoides) of America,
which so closely resemble Formica rufa in their archi-
tecture and general habits, show like characteristics in
their road-making, though I do not remember to have
seen it displayed under such difficulties.
The question inevitably arises, How did the ants
manage to lay out these roads with such precision?
When a corps of engineers or road-builders produce like
results, we easily call up certain steps that have been
taken. We think of two or more persons, provided with
surveying instruments, all dominated by the sense of
sight, and controlled by reasoning upon the facts and
figures entering into the problem, making out certain
lines, and, as far as conditions allow, laying out the
course in a straight line between the points to be
In the case of the ants we start under the embarrass-
ment of not having the facts to reason from. Were
the roads marked out at once, or are they a gradual
growth? We do not know. Has the direct course been
determined by a great number of experiences, of which
the errors in direction were gradually eliminated, and
the final result viz., the shortest path to the desired
point retained? We do not know.
It is easy enough to understand how, after a course
has once been fixed, the ants follow it unhesitatingly.
Their antennae, which are continually waved before them
and to every side, at once detect the strong odor of
formic acid on the trail. This is a perfectly accurate
guide, and beyond doubt it is thus that the workers pass
so swiftly and surely between points on the established
The sense of sight, it will be observed by the reader,
is not here considered. It is probably a negligible
factor, or at least does not appear to play a considerable
part in determining results. The visual organs of ants
are good as far as appearances show, and as compared
with those of the winged hymenoptera, as bees and
wasps, ought to contribute something effective toward
a visual memory of the localities over which their owners
operate. They may do so here. But myrmecologists
seem at one in the opinion that the vision of ants is
extremely limited; and if so, it could hardly have ef-
fectively directed them in laying out trails of such
length as here considered.
Moreover, the great ferns referred to as closely en-
closing and overhanging the ant-hills and the whole
surrounding region would probably have hindered the
effective exercise of vision, unless we suppose that
the course of the trails was fixed in the early spring
before vegetation began and the vicinage was quite
Suppose we indulge in a bit of speculation? Let us
imagine that when the winter has so far broken as to
allow the insects full liberty of out-door life, a group of
workers start out from the parent commune to explore
the neighborhood for food. After the fashion of their
kind, we see them passing to and fro in zigzag lines, in
arcs and parabola. They are thus storing their mem-
ories with impressions by which to localize their route
and insure their return. Farther and farther, and by
gradual recessions, they reach an oak-tree. To ascend
it is a part of the natural inquisitiveness (and acquisitive-
ness) which so strongly marks these insects, and forms
an important factor in the feeding of the commune.
A tree is a veritable bonanza for insects of various
sorts at sundry seasons. The sappy exudations of the
plant; the opening buds; the bleeding galls; the hosts of
insects, as coccids and aphids, that infest various parts,
and prey so freely upon its generous bulk that their
superabundance yields a rich harvest for many other
insects one or more of these may attract our foraging
And now, full laden, they are ready for the return
journey. Their path down the tree is easily traced by
their ascending trail. The real difficulty must arise as
they reach the ground and face the home commune.
Yonder it lies; the general direction is not hard (for us)
to determine. But amid all these involved crossings
and recrossings, of their first outgoing trail, how shall
they find a straight path home ?
The author is loath to resort to the supposition of
some occult power, although he believes in a divine Over-
force as the one intelligent source of all creatures and
all their actions. His own explanation (held without
positiveness) is that, by means of the odors left during
their approach and recognized by the sensitive antennae,
assisted, perhaps, in some degree, by visual impressions,
the ants discern the general course of their trail. It
may be also that a spirit of venture which possesses most
living things, and a profiting by happy chances which
befall, may aid in giving them the first bent homeward.
Moreover (and I am inclined to emphasize this point),
the home commune, as the central abode and scene of
activity of a vast multitude of ants, must be a huge
reservoir of formic fumes that strongly impregnate the
surrounding locality, and serve as a sense-signal that
affects, even at a distance, the sensitive antennoe of
the workers, and so points the direct way home. Thus
it falls out that the return is apt to be far more direct
than the outgoing.
The next outgoing naturally would be over a some-
what more direct trail than even their first return, and
so, in the course of a few trips, the first indirections
would be eliminated, and the trail established in its
lines as when I saw it. Something like this, perhaps,
may be a natural history of the method by which the
ants perform what seem to us notable engineering feats
in laying out their roads.
But there are cases which cannot be explained so
satisfactorily. While studying the cutting ants of
Texas, near Austin, I took occasion to follow up the
underground routes of some of this species. A planter,
in order to get rid of the depredations of an immense
commune near his residence, had set his men to dig it
up and utterly root it out. In order to reach the central
nest he had traced the ants from a tree inside his home
premises, which they had stripped of leaves, to a point
six hundred and sixty-nine feet distant. The nest
occupied a space as large as a small cellar, the lowest
and main cave being as large as a flour-barrel. In this
central cavern were great numbers of winged males and
females, and innumerable lame and workers. From
this point radiated the various avenues over which the
leaf-cutters marched on their raids.
With the aid of a young civil engineer, I proceeded to
survey the main course of the insects. For part of the
way we had but to follow the diggings of the planter's
laborers. For the rest, it was only necessary to sink
holes here and there along the estimated course to the
main nest, and, when the tunnel was struck, take an-
other bearing. These bearings were afterward handed
to a friend, 1 who had them translated into a chart. In
some places the tunnel was as deep as six feet beneath
the surface, the average depth being about eighteen
inches. At the "exit hole," four hundred and eighty-
four feet from the nest, the tunnel was two feet deep.
Besides this main way there were two branch tunnels,
which deflected from the trunk-line near the country
road, in order to gain entrance to a peach orchard one
hundred and twenty feet distant.
This chart shows better than any verbal description
the problem in underground road-making which the
cutting ants faced and effectually solved. [McC. 6, p.
224.] It quite confirms their ability, at least, to achieve
such an undertaking as described by Dr. Gideon Lin-
cecum, who observed a raid made by a colony of cutting
ants upon a garden situated on the bank of a creek that
flowed between their nest and the garden. In order to
reach the desired plants they drove a tunnel beneath
the bed of the stream, and, ascending on the opposite
shore, successfully raided the garden. [Li., p. 327.]
Gen. S. W. Fountain, of Devon, Pennsylvania, a re-
tired officer of the United States army, recently (1909)
related to me an incident that quite confirms Lincecum's
statement. AVhile stationed at Fort Clark, Texas, during
the summer of 1879, with Troop "E," U.S. Cavalry, the
troop garden, whose conduct was assigned to Captain
(now Colonel) A. B. KaufTman, was so persistently raided
by cutting ants, who stripped the vegetables of their
1 The late Mr. Strickland Kneass, C.E., Assistant to the Presi-
dent of the Pennsylvania Railroad.
leaves, that the site had to be abandoned. It was
transferred to a spot near by that was surrounded by
an irrigating ditch, and, thus insulated, was supposed
to be quite safe from emmet assault. But in course of
time the cutting ants appeared within the island gar-
den. The officers, supposing them to be the former
troublesome community, concluded that they had en-
tered by tunnelling beneath the stream. What other
way? But Captain Kauffman, sceptical of their ability
to accomplish such an engineering feat, resolved to in-
vestigate. He drained the irrigating stream (some four
or five feet wide), dug up the bed, and traced the in-
sects' trail from their point of entering the garden to
the old nest on the opposite side. The ants had in-
deed tunnelled beneath the water, guided unerringly by
their remarkable instinct in engineering.
I have no explanation to offer of the method by which
the Attidie accomplished these feats in subway en-
gineering. The facts are given, and they are without
question. But by what peculiar topographical sense
or gift, or by what faculty or process they carried out
schemes which compare favorably, relative powers
and proportions considered, with underground roads
and tunnels of our own species, this author con-
fesses his inability even to suggest an explanation.
One may see how upon surface trails the antennae aid
in fixing the direction of the ants' course. But it is not
possible to see how they could aid in laying out tunnels
We turn to another and widely different use of the
engineering faculty. In earlier days the gates of great
buildings and of walled towns were kept with a high
degree of diligence and ceremony. This was warranted
by their relative importance in the military methods of
the period. It has now wellnigh ceased, except as a
quaint survival of mediaeval ways in castle and town,
or as a relic of military days and discipline. It is in-
teresting to find among the communal habits of some
species of ants a like marked attention to opening and
closing the public gates. And this is not a mere survival
Fig. 39 OCCIDENT ANTS : SINGLE AND DOUBLE GATES OPEN
of a habit out of which the soul has gone, but is kept up
seriously as an important part of the communal de-
fences. Though the writer confesses that more than
once he has found himself questioning its utility; which
probably means his inability to perceive the same.
Among the communities most persistent in keeping up
this habit are those of the Occident ants, and examples
of the custom as it obtains among them, and in one
other species, will sufficiently illustrate the author's
observations. In form the gates of Occidentalis are
funnel-shaped openings through the gravel roof of the
central mound into the interior, at an inclination of
about forty-five degrees. There is usually but one gate,
located about one-third of the way from the base. It
is single or double, according as it opens into one gallery
or two, the former being about three-fourths of an inch
wide, the latter from one and a half to three inches (Fig.
39) . Within, the terminating galleries are quite smooth;
without, they present the appearance of a rude stone
wall in miniature. Around these gates gathers the daily
out-door life of the ants. Back and forth through them
citizens of the commonwealth are continually moving
during the working hours of the day.
My observations throughout the latter part of July
showed that the gates are ordinarily opened near or
shortly after 8 A.M.,
but full activity of
the colony did not
begin until toward 9
o'clock. The general
statement is warrant-
ed that the gates are
opened between 8 and
9 o'clock A.M. This
is not an early hour
to go on duty for in-
-v / sects that have such
a high reputation for
one must remember
that the interior work
of a formicary, which
Fig. 40 OCCIDENT ANTS CLOSING GATES 1S VG1 Y g 1 6a ^' Hiay
still going on behind
closed doors. Moreover, I found that in a special exig-
ency as injury to a mound by floods the ants modify
their habit, and are found hard at out-door work at
The manner of opening a gate cannot be fully de-
scribed, because the work is done chiefly within and
behind the outer door of gravel. Doubtless the mode
would be shown correctly by reversing the process of
closing gates (Fig. 40), presently to be described. What
one first sees is a pair of quivering antennae above one of
the pebbles, followed quickly by a brown head, and feet
thrust through the interstices of the contingent gravel-
Then forth issues a single worker, who peeps to this
side and that, and after compassing a little circuit round
about the gate, or perhaps without further ceremony
seizes a pebble, bears it off, deposits it a few inches from
the gate, and returns to repeat the task. She is followed,
sometimes continuously, sometimes at intervals of ten,
twenty, even thirty minutes, by a few other ants, who
aid in clearing away the barricade. After that, the
general exit occurs.
On other occasions the method is not so deliberate, or
at least it does not appear so. There is a rush of workers
almost immediately after the first break, who usually
spread over the cone, bustle around the gate, gradually
widening the circles, and finally push out into the sur-
rounding herbage. At first the exit hole is the size of
a pea, and plainly shows that sand and soil have been
used under the gravel to seal up the gate.
The process of closing gates is even more interesting
to the observer than the opening, as the various steps
are more under his notice. It will best appear by trans-
ferring from my notes a few records: About 6 this
evening (July 19th) the closing of doors began. At
nest A the work was chiefly from within. The workers
pushed the sand from the inside outward with their
heads. A grass straw about an inch long was brought
from the interior and pushed out until it lay across the
gate as a stay for the infilling material. Soil was here
used principally for closing, a few pebbles being added.
The gate was not filled up quite flush with the surface
of the mound. At nest B, which had a double gate,
two workers-minor were the last and chief operators.
They brought gravel from near-by parts of the cone and
filled in the two openings flush with the surface.
At nest C, with a single gate, a worker-major was
operating as at B. A number of ants had been en-
gaged at first filling and gradually closing the inside,
but all had retired within except one major.
When the gate was nearly sealed a straggling minor
came out of the grassy commons and essayed entrance.
Several trials and failures followed, whereupon she
commenced dragging dirt from the opening.
While thus occupied the major came up with a huge
bit of gravel, which she dropped upon her comrade with
as much nonchalance as though she were one of the
adjoining pebbles. At last the minor dug out a tiny
hole through which she squeezed into the nest, and the
major, who was again deliberately approaching close
behind, carrying another pebble, straightway sealed
up the opening. During this amusing episode the
straggler did not try to aid the closer, being wholly
bent on entering, and the gate-closer paid her no atten-
tion beyond the first satisfactory antennal challenge.
Each moved forward to her own duty with the undis-
turbed placidity of a machine.
At nest E by 6.30 P.M. most of the commune had
entered the cone; at 6.45 the gate was being closed;
at 6.55 only two ants were outside, slowly working at
the gate, then half shut. An ant came out with a bit
of straw, carried it to the refuse-heap, and returned. At
6.58 two ants came with chopped leaves, and at 7 P.M.
yet another. None of these attempted to help the gate-
closers, who slowly and steadily filled up the entrance.
Now occurred the usual side-play with late-comers.
At 7.07 a straggler came along and tried to get in. As
the gate was nearly closed, she deliberately proceeded
to break it open. A pebble was taken from the gate-
covering and carried three inches up the mound. An-
other was tugged still farther up, and yet another.
Then in steps the gate-closer, quite undisturbed by the
counter-working of her fellow, and quietly plugs up the
little break made with a big pebble, and slips within at
a by-cranny that had escaped the straggler's notice.
A second straggler appears while the closers are ad-
justing the material from within, as may be seen by the
agitation of the surface. Meanwhile the first straggler
has grappled with the last big pellet dumped, which she
succeeds in dragging aside of which straggler number
two takes advantage, and steps into the nest. There-
upon one of the closers reappears from the inside, and,
without the least token of vexation at thwarted plans,
restores the piece to its place and returns.
Straightway the first straggler renews her opposition
effort, and has just set the pebble aside when a small
black beetle comes up. This the straggler seizes, puts
down, turns, reseizes, and tries to push into the gate
therewith. The beetle, however, escapes; and the for-
aging instinct which led the straggler to forego for the
nonce her house-breaking is not strong enough now
to divert her from the home trail. So beetle goes her
way unpursued and the contrarious straggler disappears
At 7.20 a gate-closer comes out and adjusts several
pebbles. My imperfect perceptions cannot discern the
advantage thereof, but doubtless the ant quite under-
stands. The other closer is seen reaching up and ad-
justing pellets from within. At 7.21 the outside closer
goes in at the small opening between the top of the gate
and the inlying gravel. At 7.27 I still see, by the mo-
tion of the pellets of soil and agitation of the pebbles
and occasional glimpse of the tips of antenna? and
mandibles, that the final sealing-up of the communal
walls is being accomplished within. In a moment all
is quiet, and the gate of the emmet city is shut for the
night. Externally it now seems to be a simple semi-
circular or triangular depression in the gravel armor of
The other illustration of the gate-closing habit among
ants is drawn from the cutting ants (A tta fervens) of
Texas, observed in 1877. [McC. 10, pp. 33-40.] One
immense commune was assembled around the trunk of
a live-oak tree (Quercus rirens) on a road-side. The ex-
cavations from the interior had gathered into a mound
twenty-one feet long and about four feet high. This
accumulation, called by the natives a "bed," was evi-
dently one of many years' standing, and when sub-
sequently opened was found to be inhabited by legions
of ants from the size of a bumblebee to that of a small
garden ant. It seemed incredible that such hosts of
living creatures could dwell within such a narrow com-
pass and all find nourishment. And to think that
they live on mushroom gardening! as we shall pres-
My first view of the mound was a disappointment.
It was in broad daylight, and not a sign of life appeared.
Could it be an abandoned nest? Having satisfied my-
self that the mound was inhabited, I arranged for an
evening visit. Here and there were scattered over
the surface small, irregular heaps of dry leaves, bits of
leaves, chips, and broken twigs, which seemed mere
accidental drifts and piles (Fig. 41).
Returning about nightfall, I found the scene wholly
changed. Hosts of ants of various sizes were already
Fig. 41 A CLOSED GATE OF CUTTING ANTS OF TEXAS
hurrying out of open gates into the neighboring jungle,
and two long double columns were stretched from the
bottom to the top of the overhanging tree. The ants
in the descending columns carried above their heads
portions of green leaves that waved to and fro, and
glanced in our lanterns' light, giving them a weird
seeming as they moved along. It is this habit that has
given the insect the popular name of "parasol ant."
I first directed special attention to the opening and
closing of the gates, which occurs before and after every
exit. The opening began about dusk. First appeared
from beneath the heaps of dry leaves and chippage
scattered irregularly over the surface a number of
minims, very tiny fellows indeed. They carried from
within small grains of soil. Perhaps an interior cave
may have been used as a dumping receptacle for the
earth-pellets and smaller rubbish. This was a tedious
process, and little seemed to be accomplished until this
squad was joined by larger forms, who began to carry
away bits of chippage. This was also a slow process,
but it seemed to avail in gradually loosening up the
massy material crowded into the gate; for now came
a grand rush from within, the workers, major and minor,
and the big-headed soldiers in the lead. They broke
forth, bearing before them the larger bits of gate-closing
rubbish, which was scattered here and there, and in a
few moments was cleared away from the gallery and
strewed around the margin of the gate (Fig. 42).
This chippage appears to be a part of the communal
treasure, for it was easy to identify a number of the
pieces as having been used several days in succession.
The ants having found out just what shapes and sizes
Fig. 42 CUTTING ANTS: A GATE WIDE OPEN
were best adapted for effective infilling, were wise
enough to keep them close by and use them again and
The closing of gates began early in the morning and
dragged along until ten o'clock. The galleries, of which
the gates were the terminals, sloped from the surface,
at as great an angle as forty-five degrees, a conformation
that favored the process of closing, as it gave purchase to
In shutting the doors of the commune, the minors
appear to begin work by dragging the dispersed chippage
toward the gate. One after another they were taken in,
and lodged and adjusted. It was certainly not a mere
Fig. 43 CUTTING ANTS: A GATE IN PROCESS OF CLOSING
anthropomorphic fancy that in this process the work-
ers showed admirable ingenuity and a rude but effect-
ive sort of mechanical skill (Fig. 43). For example,
the longest stalks and leaves were stretched across and
wedged into the opening and vestibule so as to form a
rough scaffolding upon which the shorter pieces could
As the gate gradually filled up, smaller castes of
workers appeared upon the field, and took up the work
to which their slighter frames are adapted. The last
touches were carefully and delicately made by the
minims, who in small squads fill in the interstices with
small pellets of soil. Finally, the last laborer steals in
behind some bit of leaf, and the gate is closed. The
infilling material occupies the opening to a distance of
(sometimes) an inch and a half within the gallery. The
exterior of the gate now presents the appearance already
described of a small heap of dry chippage accidentally
accumulated upon the surface.
SUPPLYING THE COMMUNAL RATIONS
FOR all living creatures food is a first demand of
nature. The struggle to obtain it in the vegetable
and animal kingdoms keeps the wheels of life in active
motion ; and day and night, secretly and openly, silently
and with sound and stir of mighty conflict, it goes on
among and around all beings. Its influence in shaping
life and habit is constant and incalculably great. In-
deed, in many, perhaps in most cases, it is decisive at
least in certain epochs of the individual and communal
career. Thus, a study of the food supply of ants is of
highest importance in determining their natural history.
As a general rule, covering most of our common ants,
the founder of the future commune is a single fertilized
female. After the marriage flight she seeks in the vici-
nage of her alightment a suitable site in the ground
or in wood, according to her instinct. Therein she pre-
pares a brooding-cell, which is commonly forced into an
oval shape by her rotary movements in forming the
wall that shuts her in. This cell becomes the tomb of
the great majority of females, but a few survive to be
the founders of communes.
The eggs laid by the queen are tended and the young
are fed by her during her isolation, which may last
three-quarters of a year. As she never leaves her
hermitage, whence comes her food supply? Nature has
provided a store in the voluminous body, generously
nurtured during her virginity by the laborious and self-
denying workers of her home nest, who thus uncon-
sciously had wrought out a further part in preserving
their species. This stored-up substance, together with
the degenerating wing muscles, is transmuted into food,
which passes as a salivary secretion from the mother's
mouth to the mouths of her progeny. In many cases
this supply is supplemented by her own eggs, a con-
siderable percentage of which she eats.
The first individuals matured are naturally scantily
nourished; and for this reason, perhaps, appear as
minims, or ants of the smallest caste. At all events,
the firstlings are minims, and their smaller demand for
food well accords with an empty larder. The known
ability of worker ants to endure a long fast is shared to
some extent by these callows. But as their philopro-
genitive instinct at once awakes, and prompts them to
feed and care for the larval dependents in the cell,
the supply of rations is a pressing problem. Before the
double demand of hunger and devotion to the commune,
their primitive cell walls melt away, and the young ants
break forth into a new world. What a great, strange
world it must seem, even to their imperfect perceptions!
Doubtless the first circles of adventure which these
pioneers permit themselves have a short radius. That
will be measured by their initial success in foraging;
and that, in turn, will depend upon the site whereon it
has been their hap to fall. In any case, their foraging
journeys will sweep over an enlarging space, as the
demands of their growing commune increase and their
To a limited human vision the supply of available
SUPPLYING THE COMMUNAL RATIONS
edibles seems small. But to the omnivorous appetite
of ants it is, under favorable conditions, practically in-
exhaustible. Ants have proverbially u a sweet tooth."
In the vegetable kingdom, nature's vast fecundity of
flowers and blossoms and sappy sw r eets, so far beyond
what seems required for continuing the species, may seem
a great waste. But to the insect world it presents a
bountiful harvest. One sees them, of all the orders,
winged and unwinged, as larvae, as nymphs, as imagines,
in countless hordes drawing upon this exhaustless store.
Among these insects, ants are everywhere most
prominent ; on the low-growing plants, close to mother
earth, high up on the lofty trees, and in all grades be-
tween they peek and mouse. In companies, in bands,
in ones and twos, their busy inquisition is pushed, and
their restless antennae wave and tremble. They dip
into the flower-cups, and drink of the nectar there.
They scout over leaves. They exploit the trunks and
boughs. They are everywhere in Flora's beautiful
domain, lapping her sweets, filling their crops with her
treasures, growing rich from her redundancy, not for
themselves alone, but for the helpless dependents of
their communes. They jostle their winged kindred, the
bees, the wasps, the hornets, the yellow-jackets, who
come by the shorter aerial ways, but are fewer than the
persistent and ubiquitous ants, who plod and climb by
the roundabout routes which apterous beings must take.
There is enough for all ; and although I have seen thou-
sands of these various forms feeding cheek by jowl upon
some rare harvest feast of bountiful Flora's spreading, I'
recall no scenes of violence arising from the casual con-
tact. Let the reader give no credit for this to the peace-
ful temper of the insects. Simply, it is hard to quarrel
when the crop is full, and when all may have who will
all that they may will to have (Fig. 44) .
It is pleasant to contemplate this phase of the latent
helpfulness that lies in creation, and to see the inanimate
world, the fields and forests, extending to the hosts of
the Insecta so magnificent a hospitality. It is ill repaid,
Fig. 44 GUSTATORY ORGANS OF THE HONEY-ANT
(Face viewed from beneath)
Neck, fm Foramen. Ib Lahium. mar Maxilla, mx. p Max-
illary palp, mb Mandible, to Tongue. Ib. p Labial palp
SUPPLYING THE COMMUNAL RATIONS
no doubt, at times especially by the hordes that prey
upon our gardens, orchards, and fields, and compel a
ceaseless vigilance to save our crops. But, on the other
hand, it is gratifying to reflect that Flora's bounty to
ants and their insect allies is repaid by a service which
preserves her domain by perpetuating the life of plants ;
for the insects that pass from flower to flower and plunge
into the cups, distribute the fertilizing pollen upon which
fruitfulness and life depend. This symbiosis between
plants and insects is thus a necessary condition for
both ; and that it has existed from the beginning, a study
of fossil insects shows.
The above seems, as indeed it is, a vast field where-
from to cull a living. But one, perhaps as wide and even
more lasting, is open in the waste products of nature.
Ants are universal scavengers. They are fond of animal
oils and juices. Countless millions of insects perish
every season. What becomes of them? They drop by
the waysides of their lives, and drift into all manner of
crannies and corners. Hereto the ants follow them.
The searching power of the antennae is something mar-
vellous. It has been compared to that of men's hands
were the sense of smell to be located in the tips of all
their fingers also, where such a delicate sense of touch
abides. What human hands could do, in such a sup-
posed case, to follow up and search out odors, the
movable organs of smell, the antennae, do actually
accomplish for ants.
Thus are revealed to them the carcasses of the innumer-
able hosts of fallen insects; and often they may be seen
headed for their homes, dragging with them whole
bodies or parts thereof, and making painful headway
therewith through the jungle of grasses and weeds.
Commonly, however, they are successful, when not way-
laid and robbed by stronger individuals, or by roving
bands of alien species, or congeners of other communes.
The fondness of ants for animal fats and juices may
be tested by placing a fresh bone on the lawn or in a
field. It will be covered soon with a crowd of emmets
greedily lapping the oily particles upon the surface and
exuding from the pores. Some housekeepers avail
themselves of this appetite to collect groups of the little
red ants (Monomorium pharaonis) that infest houses.
These being destroyed, the "trap" is set again and again.
Another source of food supply is the various fruits in
season, wild and cultivated. The windfalls lying be-
neath the trees and bushes are usually bruised, or stung
by insects. Around and into these broken parts the
ants gather and feast. They climb to the laden boughs.
They scout among the ripening fruits. They have a
quick touch for a spot of decay, which has opened a way
for their gustatory attack. Or a bird's bill has been
before them, cutting a little trough from which to sip
the sweets they love, and herein one will see a bunch of
ants scooping out the pulps and drinking their fill of
the fermenting sap.
When September's sun has mellowed the grapes, you
may see legions of ants, joint pilferers with birds and
bees, hornets, wasps, yellow- jackets, and flies, many
with heads buried deep within the berries. They are
lovers of the new wine of the grape, and many empty or
partly emptied skins, hanging among the broken
clusters like cups drained of their contents, show how
often and deeply they and their winged comrades have
drunk. Doubtless ants are apt to be a bit injurious to
our orchards and vineyards. But their share in the
SUPPLYING THE COMMUNAL RATIONS
general scavenger work of nature, as well as their part
in fertilizing the blossoming plants, may be set to the
credit side of their account. Besides, if folk would
follow the author's rule to put in one root-stock for the
birds and insects for every two set out for himself, there
would be enough and to spare for all.
Preying upon insect remains and animal oils, lapping
the nectar of flowers and the sweets of fruit, by no means
exhaust the sources from which foraging ants may draw
their rations. They are free-lances, and they do not
scruple to ply their freebooting against all and sundry
whom they are able to better in a quarrel over booty.
After the manner of human cannibals, they feed upon
their vanquished foes; indeed, the formal raids of
slave-making ants are chiefly for food. Many thousands
of their victims are carried home and eaten. The
tender larvae and pupse are kept in store for the slaughter
as human butchers keep live-stock and fowl, though one
cannot aver that the ants deliberately fatten them for
that purpose. Some captives, and at times a number,
escape the shambles and become auxiliaries or slaves.
But large communes of these kidnappers have been
known to end an active season of slave-catching with
but few if any increase in the number of slaves. All
their captives had been eaten!
The same methods are quite commonly carried out
on a smaller scale among various species. I have turned
up a flat stone, beneath which was a large nest of small
ants. Their larva*, still smaller than themselves, lay
in heaps against the under surface. Scarcely had the
stone been lifted ere several larger ants, representing
two other separate species, rushed in and began plunder-
ing the colony. They evidently had been prowling
around the confines of the nest, waiting for an oppor-
tunity to break through the barriers, or snook into some
chance opening by which they could reach their desired
prey. And this is a typical incident in ant world.
Other insects, both in the larval and imago state, are
victims of this passion for hunting live game as fierce
and high as ever fired human devotees of the chase.
Who has not seen an unfortunate caterpillar writhing in
mortal agony beneath the assaults of a large squad of
small ants? In vain the victim struggles to throw 7 off
its assailants. Its fate is sealed. Spurred into violent
contortions by the smarting thrusts of stings and cuts
of mandibles, it flings its tormenters to this side and
that. They hold on grimly amid all the thrashings
until the quiet of death gradually falls. Then the great
carcass is dragged and pushed home by a gang of workers
or carried thither piecemeal, a vast addition to the
While studying the agricultural ants in Texas, I
observed, after a summer shower, a great commotion
upon a large circular disk of one of the colonies. The
rain had beaten down a great number of the winged
forms of swarming termites, and upon these the red
agricultural were charging from all parts of the plaza.
They seized them in their jaws and ran toward the
central gate, out of which a file of their fellows was
eagerly streaming, intent upon sharing the rare find
of booty. The outgoing and incoming columns met.
The gate was soon choked up. The tiny rivulets caused
by the rain were setting in the same direction, and
presently a mass of excited insects was balled and matted
around the gate., pushing and tumbling over one another,
and splashing in the water. But the possessors of
SUPPLYING THE COMMUNAL RATIONS
captives from the fallen swarm held to their prizes amid
all the tumult, until the jam was relieved, and they
could get entrance within their nest. Their under-
ground granaries were at the time full of stored ant-rice
and other seeds, their provision for winter food ; but the
eagerness '.with which they welcomed this chance supply
of soft, juicy insect flesh showed that their granivorous
habit had not weakened their insectivorous taste.
I have seen the mound-making ants of the Alleghanies
make raids upon the workers of our northern species of
termites (Termes ftavipes), capturing and carrying them
away with an eagerness that showed what precious morsels
their soft, white bodies must be to the Formican appetite.
Here we may consider the vast food supply that lies
in the product of certain seed - bearing grasses, the
garnering and storing of which has led to the popular
names of " agricultural," "farming," and "harvesting"
ants. 1 The habit is especially developed in the genera
Pogonomyrmex and Pheidole, which collect from the
ground and from plants certain grain-like- and nut-like
Fig. 45 AGRICULTURAL ANT FEEDING FROM A
seeds (Fig. 45). These they carry to their nests, and,
after removing the husks, and deporting the latter
1 See the author's Nature's Craftsmen, chap. vi.
from the nests, they store the kernels in large granaries
excavated in the ground, where they are kept for food.
It has been noticed that these seed-eating ants are
marked by the presence of large-headed workers, whose
unusual development of the muscles of head and jaws
particularly fits them to crack and crush the seeds, so
that from the meaty kernels may be rasped or squeezed
the edible starchy or oily parts.
A remarkable example of the unexpected way in
which Nature varies her methods of feeding her children
is seen in habits of the Attidse, the "parasol" or "leaf-
cutting" ants. This popular name is due to their
manner of sending out expeditions of workers, who cut
from the leaves of certain trees and bushes small pieces
which they bring into their nest to convert into food.
These leaf-cutting excursions are striking sights to the
novice, and are not apt to lose their interest even to
familiars. The author's chief studies of Atta fervens 1
were made in the vicinity of Austin, Texas. [McC. 10, p.
33 sq.] Most of their nests were "beds" (as the natives
called them), or spots of denuded surface in the flat
open prairie, eight or nine feet long and of almost
equal width. Over this barren space were thrown up
twenty or thirty circular, semicircular, and s-shaped
elevations of fresh earth pellets. The circular mound-
lets were about the size and form of a "pound-cake "-
pan, or spittoon, the resemblance being emphasized by
a round open entrance in the centre. All these had been
naturally formed by the gradual accumulation of the
1 I have used throughout this, the old and well-known name of
this species, although Professor Wheeler has restored the yet
older specific name of Buckley, by which it will doubtless be known
SUPPLYING THE COMMUNAL RATIONS
, : ; - -:
Fig. 46 MOUND NEST OF TEXAS CUTTING ANT
UNDER A LIVE-OAK TREE PARTLY
sandy soil as it was brought out and dumped upon the
heap, being massed at the base and gradually sloping
to the top. Another of these beds, the one which was
especially studied and finally opened, was a large mound
at the foot of a live-oak tree (Fig. 46) . This had prob-
ably been formed by a road or trail that passed with-
in several yards of the tree, thus restricting the limits
of the gates and throwing the separate moundlets back
upon one another. [McC. 6, p. 231.]
At my first visit to this great commune it seemed like
an abandoned nest. On the spreading branches of the
overshadowing tree and on the defoliated vines at its
base were marks of recent raids of the leaf-cutters.
But no life was anywhere visible. The surface was
covered with earthen knobs or warts of various sizes,
and here and there were scattered small irregular heaps
of dry leaves and bits of leaves, and twigs. As evening
began to fall the scene changed. Hosts of ants of various
sizes, in countless numbers, suddenly burst from gates
that mysteriously opened for them, and began a hurried
march into a near-by jungle. Two large double columns
began to ascend the trunk of the live oak. Along their
flanks, both going and coming, moved the soldiers,
Fig. 47 THE HEAD OF A TEXAS CUTTING ANT
Enlarged eight times, to show furrow and spines and cutting jaws
marked by their immense heads (Fig. 47). They rare-
ly handled the leaves, but seemed to act as scouts or
pioneers or attendant guards.
SUPPLYING THE COMMUNAL RATIONS
In a few moments the two-ranked army had reached
the top of the tree, and the work of defoliation began.
In order to view the mode closely, small branches broken
from the tree were set in the mound
near the gates. They were soon cov-
ered with ants, and in the lantern
light their method could be seen read-
ily. It was like that of Megachile, _\
the leaf-cutter bee. [McC. 7, p. 145.]
The cutter, seated on the leaf, grasped
it with outspread feet, and made an
incision at the edge by a scissors-like mo-
tion of her sickle-shaped and toothed mandibles. She
gradually revolved, cutting as she moved, her mandibles
thus describing a circle or a portion thereof. The feet
turned with the head. The cut was a
clean one, quite through the leaf. The "
Fig. 48 A PROCESSION OF TEXAS ANTS CARRYING LEAVES TO
A. XT COMMUNITIES
cutter would .sometimes drop with the excision to the
ground; sometimes it let the section fall; sometimes
carried it down. At the foot of the tree lay a pile of cut
leaves, to which clippings were being added continually
by droppings from above. Squads of carriers from
the nest took these up and bore them away (Fig. 48).
This is the manner of loading the cuttings: They are
seized by the curved mandibles; the head is elevated;
the piece is thrown back by a quick motion, and lodged
on its edge within a deep furrow that runs along the entire
median line of the face, except the clypeus, and is sup-
ported between prominent spines on the border of this
furrow and on the prothorax. These peculiar features
of the Attidse thus serve a useful end. As far as noted,
the cutting and carrying were not done by the minims or
smallest castes, but by the worker-minors; the soldiers
rarely engaged therein. As the ants moved along down
the branches and trunk of the tree, and over the ground
to their gates, holding above their heads the bits of green
leaves, which waved to and fro and glanced in the lantern
light, the column had a weird seeming.
The citizens of this commune, and of some others ob-
served, made their leaf-cutting sallies in the night.
But this is not the universal habit. I afterward saw
carriers marching with their loads during the day. I
also observed them frequently in day marches in the
vicinity of Santiago de Cuba, during the Spanish-Amer-
ican war, carrying on their quaint industry among the
graves of fallen American soldiers and in the tropical
trees that sheltered them.
They were abundant on the great terraced height of
the Morro, or castle, at the mouth of Santiago Bay,
which I visited just after the surrender, and before it
SUPPLYING THE COMMUNAL RATIONS
was occupied by the American troops. When ascending
the path that zigzags around the cliff to the summit, a
column of these insects was seen marching with their
bits of cut-off leaves. On the summit, workers were
found close by the dismantled eastern battery. Their
fresh-made tumuli were cast up almost beneath the butt
of the great guns. The workers were thronging into the
central gates, bearing aloft their leafy banners. One
could not but wonder: Were these industrious creatures
plying their task while Spanish cannons were firing
and shells from American ships were bursting around
them ? No doubt they did so a type of the army of in-
dustry in the insect world prosecuting the humble arts
of peace amid the roar of human battle and the clash
of arms. If their wee brains could be deemed capable
of thinking on such matters, we may fancy their thoughts
taking shape in the familiar words: "What fools these
mortals be!" -maiming and killing one another when
they might be comfortably cutting juicy leaves and
chewing them into pulp!
At an afternoon visit to the grounds of a nurseryman
and gardener near Austin, Texas, the leaf-cutters were
seen at work. They had come up through the garden
from their colony, three hundred feet distant. From
this gentleman it was learned that these ants prefer
trees with a smooth leaf; are severe upon grapes, peaches,
and the china-tree. They take radishes, celery, beets,
young corn, and wheat, plum, pomegranate, honey-
suckle, cape jessamine, crape myrtle, and althea. They
do not like lettuce, nor the paper mulberry, nor figs, nor
cedar, except the bud ends in the scant days of winter.
They love sugar, grain, and tobacco. This proprieter
assured me that the ants made foraging excursions into
his house, entered his desk drawers, and carried away a
portion of his chewing-tobacco before he discovered the
robbery. He had to be careful thereafter where he put
the delectable weed. At a plantation not far from this
nursery I saw an immense column of Attas plundering
a granary of wheat, which was being carried away in
quantities, grain by grain. This pilfering was also
carried on in the daytime. I have no explanation to
give of this remarkable difference in habit in the same
species, in the same locality, and apparently under the
same conditions. Can A tta fervens have entered upon
a transition period in its history?
How do the cutting ants dispose of all this material so
laboriously imported into their underground city? Is
it used, as with the cutting bee, simply to line the
chamber or cells in which the young are reared? Let
us see. It was no light undertaking to open and ex-
plore a mound occupied and defended by hundreds of
thousands of irate ants. But it seemed necessary.
Two trenches were made, one ten feet long and five
feet deep, and a second at right angles to it wide enough
to allow free entrance for study. The number of in-
sects that swarmed to defend their home was incalculable.
It amazed us to see such hordes of creatures domiciled
in one commune. They were, however, not so difficult
to manage as when disturbed at their night work, as
the swift use of the spade by the assailants and the
general convulsion of their emmet world seemed to daze
them. But when the author entered the trench to work
with trowel, knife, foot-rule, and drawing materials, the
ants rallied and attacked so freely that all the help-
ers were required to brush them off. The wound in-
flicted by them was sharp, but nothing to compare
SUPPLYING THE COMMUNAL RATIONS
in painf ulness with the sting of the agricultural ant
The interior of the great formicary (fifteen feet long,
eight or nine feet wide, and four feet high; I do not
Fig. 49 CUTTING ANTS
1. Winged female. 2. Male. ?. Soldier. 4 Worker-major
of the cutting or mushroom gardening ants.
know how deep, but certainly five and probably ten
feet) may be briefly described as an irregular arrange-
ment of caverns communicating with one another and
with the surface by tubular galleries. These caverns, or
pockets, were of various sizes, three feet long and less
by one foot deep and eight inches high and less. Within
these chambers were masses of a light, delicate leaf-
paper wrought into what may be called "combs." Some
of the masses were in a single hemisphere, filling the
central part of the cave. Others were arranged along
the floor in columnar masses two and a half inches high,
in contact with one another. Some of these columns
hung like a rude honeycomb, or wasp's nest, from roots
that interlaced the cave. No leaves were intact; none
used, like the leaf-cutter bee, for lining wall or floor
In color this material was either of a gray tint or a
leaf-brown. It was all evidently composed in great
part of the fibre of leaves which had been reduced to this
Fig. 50 SECTIONAL VIEW OF A CUTTING ANTS* NEST
Showing mushroom garden caves in sight. (From nature)
form within the nest, doubtless by the joint action of the
mandibles and salivary glands. On examination the
mass proved to be composed of cells of various sizes, an
irregular hexagon in shape, narrowing into a funnel-
SUPPLYING THE COMMUNAL RATIONS
like cylinder. Ants in great number, chiefly of the
small castes, were found in these cells. In the first large
cave opened there were also great numbers of larvae.
Large circular openings ran into the heart of the mass.
The material was so fragile that it crumbled under even
dainty handling, but some specimens were preserved
and exhibited in the Philadelphia Academy of Natural
Sciences. Thus the query was answered in part, at
least : What do the ants do with the leaf-cuttings carried
into their nests?
But the most remarkable part of this history remains
to be told.
While pursuing these investigations, the author knew
that the suggestion had been made by Belt that the
leaf-paper masses of Atta's nest were used as a sort of
mushroom garden for cultivating a minute fungus which
the ants used as food. Examination with a lens showed
him the presence of these growths. But as this was
only what might be expected in such underground con-
ditions, and notwithstanding all that he knew of the
ingenuity of ants in providing for the natural wants of
their communes, he put aside the theory as improbable,
and failed to push experiments which he might have
made. His incredulity thus lost him the opportunity to
anticipate in part, at least some of the brilliant dis-
coveries of such later investigators as M oiler, Von
Ihering, Doctor Goeldi, and Jakob Huber. Briefly
summarized, these discoveries are as follows:
In the case of Atta sexdens (the Brazilian cutting ant),
after her marriage flight the fertilized female begins to
dig in some open space a burrow about three-fourths of
an inch in diameter. It is at first so small relatively
that she cannot turn around in it, but has to back out
in order to get to the surface. But gradually the burrow
grows as the queen cuts off and squeezes together little
balls of earth, which are deported beyond the entrance.
When the terminal chamber is finished, the tubular
entrance is sealed up. Later a little packet of eggs is
laid. Beside it appears a small heap of loose white
substance which gradually enlarges until it reaches
the form of the spherical or elongated masses of gray
comb-like matter heretofore described. And now the
transparent pyriform globules of fungus - hyphse begin
to bud out, which Moller has called "kohlrabi." On
Fig. 51 A
PIECE OF LEAF-PULP WHICH FORMS THE MUSHROOM
GARDEN (After Moller)
these the ant feeds frequently. In truth, this is her
fungus garden. It becomes in time the source of her
food, just as an artificial mushroom cave or cellar pro-
duces nourishment for men (Fig. 51).
In time the first workers are hatched, and they too are
fed upon the kohlrabi. As they increase in number
SUPPLYING THE COMMUNAL RATIONS
they break the cavern seals and go forth to gather
leaves, which they chew and knead into pulp, and add to
the fungus garden. And this is the purpose of those
leaf-cutting expeditions which have enlisted the curiosity
of casual observers, and excited the ardent and attentive
study of many naturalists. They are to supply sub-
stance for the fungus gardens from which Attid ants
glean food for their young and themselves.
Before this strange form of plant culture has reached
this advanced stage, however, Huber introduces us to a
remarkable observation, in answer to his query: How
does the Atta female keep the fungus alive ? For, plain-
ly, the scant substance in the mother pellet must soon
be drained of its original nutriment. How, then, are the
growing fungi fed? The queen tears with her mandibles
from her little bed a minute bit of the fungus, and applies
it to the tip of her gaster, which is bent under for the
purpose. She then emits a clear yellowish or brownish
droplet, which is at once taken up by the spongy tuft of
hyphse. Thereupon it is returned to the garden patch,
and patted into place with the forefeet. This per-
formance may be repeated once or twice an hour or
oftener, and several of the enriching droplets may be
seen at once scattered over the bed. If a piece of the
fungus growth from a maturer nest be supplied to an
Atta female at this stage, she is prompt to appreciate and
utilize the gift. She divides it, drenches it with her fecal
droplet, and builds it into her garden. Meanwhile the
mother feeds largely upon her own eggs, and when the
first larvaB appear feeds them also with eggs, pressing
them directly into the little creatures' mouths.
Soon the first adults begin to appear, which are always
minims, or workers of the smallest caste. This intro
cluces a new order into the commune. They take charge
of the garden; they feed the larvae; they feed upon the
kohlrabi; they begin to enlarge the central chamber,
and in seven weeks after the founding of the colony they
are out in the open dumping their earth-pellets upon
their circular moundlets, and ere long the colony is send-
ing out its leaf-cutting excursions.
Thus we see ant communes, under the exigency of the
need of food, developing the habit of what has not
inaptly been called mushroom gardening. They have
mastered the method of liquid manuring, and of inoculat-
ing exhausted "soil" with an infected culture. They
have learned the value of triturated vegetable matter as
furnishing substance and enrichment for their gardens,
and apparently they have found out that for this pur-
pose certain plants are more valuable than others.
If such a principle or practice of plant culture were
to be as generally and as faithfully applied to gardening
and farming among men, it would need no prophet to
predict that a new era would dawn upon the agricultural
world, and such abundance would follow as our race has
Now very naturally arose the inquiry : Whence did the
Atta queen obtain the fungus germs with which to stock
the original garden? This was solved by Von Ihering
in the brilliant discovery that on leaving the parental
nest the young queen carries with her in the posterior
portion of her oral cavity a very minute pellet of hyphae
of Rozites gongylophora, and small fragments of bleached
or chlorophylless leaves. This, it is believed, is held in
the mouth until she has prepared her foundation cham-
ber, when she ejects it, and infects therewith the be-
ginning of her fungus garden.
SUPPLYING THE COMMUNAL RATIONS
Thus the sober and well -attested facts of scientific
truth prove stranger than the widest stretch of fancy
would have dared to invent. Even in the face of un-
impeachable testimony, one finds one's self startled and
wellnigh staggering before such a remarkable instinct
in an insignificant insect.
In the summer of 1880 my attention was called by the
Rev. George K. Morris to a small ant \vhich he had
discovered at Island Heights, a seaside settlement on
Toms River, New Jersey. I recognized it as a species of
Atta which I believed to be new, and gave it the specific
name of Septentrionalis. 1
I was so strongly impressed by the appearance of this
tropical species so far north that I at once visited Island
Heights to study the insect in site. I found that the
ants harvested the needle-like leaves of the pine, which
were borne into their nests and treated in a way quite
like that observed by the Texas Atta, but on a great-
ly reduced scale. The nests examined were without
an exterior mound. A single gate communicated with a
short tubular gallery with a small spherical vestibule,
which again opened into a similar but larger cave three
inches in diameter. Hanging to the roots that thread-
ed this cave were several masses of gray leaf-pulp, the
analogues of those in the fungus gardens of Atta fervens.
As one contrasted the extensive excavations and the
formidable and vigorous communes of the Texas species
with the small numbers, slight excavations, and sluggish
movements of these Northern allies, he could not for-
bear the thought that the New Jersey colonies of
Trachymyrmex septentrionalis are the feeble remnant of
1 Professor Wheeler has placed it in the genus Trachymyrmex
(T. septentrionalis McCook).
a once mighty people left or thrust by some untoward
change upon unfavorable sites which must work toward
their extinction. [McC. 15, p. 360.] Or, on the other
hand, may we conjecture that here are the first stages
in the origin of a new species already on the march, and
against unfavorable environment, toward the consum-
mation of such a splendid confederacy as has occupied
the hills of Austin?
FEEDING THE COMMUNE
CONTINUING our studies of the emmet modes of
\J feeding the commune, our thoughts once more re-
turn to the great confederacy of mound-making ants
among the Alleghany Mountains. We fix our attention
upon a column of workers pressing along a well-worn
path straight from a large mound to an oak-tree that
stands by a boundary stone wall eight rods distant.
There the column leaves the ground, mounts the trunk,
and is lost among the branches.
But here several interesting things are noted. There
is a descending as well as an ascending column. More-
over, there is something like the sentry service establish-
ed at the gates. There is a tree-trail one to three inches
wide, to which the ants steadily keep, and which is
blackened by the continuous fumes of formic acid issuing
from them. On either side of this are watchmen, who
persistently challenge passers-by. There follow swift
crossings of antennae and mutual recognitions how one
longs to know the countersign! prompt withdrawals,
and the pilgrims pass on and are soon distributed among
the principal limbs. A goodly number lead off upon
one of the lower boughs which overhangs the stone
Mounting this, one has the key to the movements of
the marchers on the avenue beneath. At various points
along the branches are vast numbers of aphides, small
black insects with brownish thorax and head. Note
this one, whose abdomen is raised at an angle of forty-
five degrees. Upon the apex is shining a tiny globule of
transparent liquid. It is lapped up by the attendant
ant, who all the while with alternate strokes of antenna?
gently embraces or pats the insect. Again and again in
rapid succession the sweet excretion, pumped by the
insect from the sap of the tree, and converted by it into
the honey-dew of popular speech, gathers in droplets,
and is removed by the ants, several of whom have en-
joyed the refection in turn.
At last the aphis, one of mature size, leaves its position
and moves along the branch toward the trunk. Its ab-
domen is now flattened. Many of its fellows have that
organ full and rounded out, and must be uncomfortable.
The ants, however, are fast relieving them as the sweet
excretion flows, and in the mean time their own abdomens
are undergoing a noticeable change. They swell and
elongate until the folded membranous bands that unite
the segments are pushed out into narrow white ribbons.
This is caused by the rapidly expanding crop into which
the collected sweets are stored. At last the honey-dew
gatherer, whom we may now call a "replete," is satisfied,
and turns toward home. It is such as she that compose
the descending column of ants upon the tree-trail; and
their full, elongated abdomens and white bands form
quite a contrast with the round black abdomens of their
fellows of the ascending column.
We are now on the verge of one of the most interest-
ing facts in the history of this remarkable community.
These repletes belong to a section of the communal
foragers, of whom thousands are elsewhere abroad, not
FEEDING THE COAiAlUXK
simply feeding, but collecting food supplies which they
are taking home in their mandibles or stored in their
capacious crops for the natural dependents and oth-
ers of the formicary entitled thereto. Following with
closer attention the trail of the repletes, you observe
some of them suddenly disappear at the roots of the tree.
Turn back the sod, clear away the leaves ; what do you see ?
Masses of insects are huddled together in the angles
of roots at the foot of the tree and in sundry depressions
in the soil. Some are repletes, some are ordinary work-
ers; and the latter are stopping or trying to stop the
former, who seek to avoid them and to push into certain
openings that lead into galleries beneath the surface,
which evidently communicate with the central mound.
A few succeeJ in this, but many yield to the friendly
force and halt.
And now what? See this replete. She has raised
herself upon her two pairs of hind legs until her body
slants in a wide angle toward the horizon. And one,
two yes,three workers, assuming a like rampant position,
have placed their mouths against the replete's mouth.
Look closely now, and you will see a droplet of amber
or whitish, syrup-like liquid gather upon the delicate,
thread-like maxillae beneath the replete's jaw. It is
the honey-dew obtained from the aphides upon the oak.
It has been forced up from the crop by pressure of the
contracting muscular sac that encloses it in other words,
by regurgitation. It is greedily lapped by the three
" pensioners," and the replete breaks away and disap-
pears within one of the gallery doors. All around the
foot of the tree are like scenes wrought visiting ants
taking toll of the foragers. 1
1 In connection with these facts, see Nature's Craftsmen, chap. iii.
Who are these visiting ants? Are they highway
robbers? They are certainly not aliens, for the rela-
tions of all concerned are most friendly. There is, in-
deed, here and there a slight show of force in the deten-
tion of a replete who has more than usual reluctance to
part with its stored sweets, but there is no element of
real hostility therein. Plainly repletes and pensioners
are citizens of one community, and their behavior must
torm a part of a natural social arrangement. What is it ?
Not all at once, but gradually, the facts dawned that
repletes, acting as communal foragers, were carrying
supplies to the formicary ; that numbers of their fellows,
engaged as builders, sentinels, and nurses, had left their
several duties for a little while to feed, and instead of
spending time and energy due to the commonwealth in
gathering food afield, had come out to tap the garnered
stores of their comrades, and, having relieved their hun-
ger, would return to their labors. In short, they had
been drawing rations from a sort of field commissary
department. They are no devotees, these adventurers,
of the theory that,
"To feed were best at home;
But thence, the sauce is ceremony;
Meeting were bare without it."
There certainly seemed to be scant ceremony in this
method of banqueting abroad. In truth, it had the
outward look of levying mail or highway robbery, al-
though there was no real violence on the part of those
who bade the repletes "stand and deliver." Indeed,
upon due reflection, the affair resolved itself into a benefi-
cent social function, of which the following appears to
be the spirit and intent: The ants at work in or about
FEEDING THE COMMUNE
the home premises leave the collecting of food to others
of their fellow-citizens, not only for the public dependents
but for themselves. Content with satisfying the simple
wants of nature that they may have strength to toil,
they leave their work and visit the feeding-grounds to
get food from the repletes. The stations for this pur-
pose are wisely chosen; for, as many of the foragers are
overladen, their progress homeward is eased by yielding
somewhat from their stores.
Besides, it seems probable that the instinct which
urges repletes to gather supplies for home dependents
might, after the formicary had been reached, prevent
parting with them to others. Moreover, since ant nature
in some degree is partaker of the weakness of human
nature, it is supposable that the surplus honey-dew, after
feeding dependents, would be kept for individual de-
lectation, and the home working-force be compelled to
leave their work and forage for themselves. The general
movement, therefore, to arrest repletes at stations near
the feeding-grounds is evidently for the public good.
It would be an odd speculation to consider the effect
upon society were such a rule to prevail among men.
Suppose the citizens of cities like New York, Phila-
delphia, and Chicago, or of such states as Georgia, Ohio,
and Massachusetts, were to agree that one moiety of
their number should take the duty of earning or collect-
ing food supplies of every kind for the entire community,
leaving all other duties to the rest? Further, suppose
that these gatherings must be divided with equal hand
among all sorts and conditions of people young and
old, active and dependent, high and lowly, rich and poor,
with sole regard to their real natural needs ?
Stop! Our phrasing is faulty; for in this ideal state
of society, if fairly conformed to the type of an ant
city, there would be no rank or grade, no rich or
poor, no personal distinctions, no individual property.
All things would be in common. There would be one
and only one property-holder the State; nor would
even the faintest desire for separate possessions ever
cross the thought of the most fanciful. There would
be no lust for riches or superior place or an easier lot
in life. One purpose would dominate all with absolute
sway : to serve the All the whole community with all
one's powers, in any line of required duty, without
hesitation, without stint, without reserve, and without
This is truly a wild speculation! This is to conceive
of the inconceivable that human beings could attain
the social standards of an ant-hill! One must first
suppose a moral revolution which even the dreams of a
Golden Age or a Millennium such as idealists in every
century have had would dimly depict; a revolution
more radical than that implied by a literal conformity
to the precepts of the Sermon on the Mount and the
Saviour's summary of the moral law. It would be a
revolution not only in socjal characteristics, but in in-
dividual character, a veritable palengenesis of every
member of the commonwealth. Would it be for the
better or the worse? Would our race gain or lose by
achieving the communistic - individualistic type of the
government of ant cities?
It is evident that there must be a good deal of varia-
tion in the food supply even under favorable conditions.
This would be felt at times when other labors of the
commune, as extension and repair of the living quarters,
interfere with the regular foraging. Moreover, there
FEEDING THE COMMUNE
are periods when the reproduction of eggs by the queen
is especially active, and the vast increase of larvae, all
needing food and care, greatly multiplies the demand
both for food and labor. The pinch of such conditions
must fall inevitably upon the helpless young. Should
they chance to come contemporaneously with a few
days of scarcity, which may arise from various causes,
the communal dependents must certainly fare ill, and
the death-rate be enlarged among them.
And it befalls communes of ants, as it comes to nations
of men, that great deviations from the ordinary course
of nature bring about disasters, at times so great that
the very life of the community is at stake. Famine
follows in the wake of war and floods, and untimely
frosts and droughts consume the sources of food in the
world of ants even as of men. The plough turns up the
fallow field, and multitudes of ant-nests are destroyed
by agriculture. Thus some of our noblest species of
native ants are vanishing before the advance of man,
as are higher types of animals.
However, the vitality of some species under the
strain of famine is remarkable. Miss Fielde has shown
(Tenacity of Life in Ants} that the workers of Cam-
ponotus americanus may live nine months without food.
They thus rival, in their ability to endure a prolonged
fast, the queens that go solitary and draw upon their
reserve tissue enough for self-sustenance and also to
nourish the firstlings of their flock. The common
mode of feeding the young, as heretofore described, is
by transfer from the nurses' crops by regurgitation.
But a wholly different manner has been observed that
is more like our way of giving food to domestic fowl and
animals. This grows out of the fact that the larvaa of
Pachycondyla and some other genera are able to feed
themselves; perhaps have been educated thereto,
though the natural aptitude must have underlain the
habit. While lying upon their backs the larvaB suck the
juices of particles of food given them. The nurses of
Leptogenys dismember termite nymphs and scatter the
pieces among their larvse, who thrust their beaks into
the soft parts and feed thereon. So also workers of
Odontomachus will tear off the heads and legs of house
flies, cut the thorax and abdomen into pieces and feed
them to their Iarva3. In the above cases the food was
not first masticated, as is done by social wasps, but simply
cut into pieces to expose the soft parts to the larval
mandibles. Adlerz has made like observations of the
larvae of Leptothorax, Stenamma, and Pheidole, who are
fed with solid as well as liquid food. [Quoted W. 11,
p. 709.] Such increase in the variety of food and feed-
ing the young must add to the chances of their whole-
some survival by lessening the danger of a failure of
food, since it greatly widens the field from which avail-
able supplies may be gathered.
An example of the strange exigencies that befall the
inhabitants of an ant commune appears in the case of
certain workers of Pheidole commutata that become in-
fested with large internal parasites, and are therefore
known as Mermithergates. This condition is accom-
panied with an enormous appetite, and they con-
tinually beset the nurses for food, which they get often
at the expense of the hungry larvae.
The voracious creatures not only ply the nurses with
mimetic entreaties, including the out-thrust tongue,
but keep up a stridulating chant of solicitation. At
times they resort to more vigorous measures, and
FEEDING THE COMMUNE
seizing a replete, hold down its head with their large
forefeet, and compel it to give up the contents of its
crop. This greediness has its penalty in times when
food is scarce; for in order to rid the commune of
such voracious and non-productive mendicants, they
are killed outright or starved to death by the
The honey-ants as studied by the author in Colorado
made their night expeditions into a scrub-oak copse, and
the sweet liquid with which their crops were filled on
their return was collected from oak-galls formed upon the
twigs and branches 1 (Fig. 52). But, doubtless, like other
Fig. 52 SPRIG OF DWARF-OAK (QUERCUS UNDULATOR), WITH
GALLS EXUDING DROPS OF SWEET SAP
ants, they know the value of aphides, and, as the seasons
change, gather from them and from other sources the
1 Nature's Craftsmen, chap. x.
supplies for current sustenance, as well as for their
peculiar mode of providing for future wants. [McC.
v, p. 17 sq.]
Among the honey-ants the workers, though varying in
size, are structurally alike (Fig. r>3). Yet certain indi-
viduals, quite independent of caste, and following an
impulse unknown, but apparently fixed in the germ and
Fig. 53 THE DIGESTIVE TRACT OF A HONEY-ANT
Showing asophagus, a, as a nearly straight tube from the mouth
to the abdomen, c Crop, gz Gizzard, s Stomach
early manifest in the callows, begin to store up food in
their crops, and thus develop into rotunds or honey-
bearers (Fig. 54). It must be allowed to be a curious
manifestation of communal philomyrmicry which causes
one of the most active of creatures to become little more
than an animated honey-pot, that the food supply of its
fellow-formicans may not lapse. But so we find it; and,
after all, it is little more than a development to its cli-
max of an instinct that urges ants of other species to
charge their crops with an excess of food in order to
impart it to the commune dependents.
The insectivorous habit of ants has been utilized as a
check upon the increase of certain destructive cater-
pillars. The author's attention was called to an article
on the "Utilization of Ants as Grub-Destroyers in China,"
by Doctor Magowan, of Wenchow, and this led him to con-
FEEDING THE COMMUNE
Fig. 54 Honey-bearers or rotunds (a) in situ in a natural nest.
6 Same with workers in an artificial formicary
sider at some length whether and how far those insects
could be used in the United States. 1 [McC. 8, p. 263.]
1 The information was received from a copy of the North China
Herald of April 4, 1882, sent me by the Rev. Dr. Hunter Corbett,
a Presbyterian missionary at Cht>f<><>, China.
Fig. 55 ABDOMENS OF HONEY-ANTS
Showing the progressive development of the crop in various
worker forms from the worker-minor to the honey-bearer
See footnote page 113
FEEDING THE COMMUNE
According to Dr. Hunter Corbrtt, in many parts of the
province of Canton, where cereals cannot be cultivated
profitably, the land is given up to orange-trees. These
are subject to attack by a species of "worms" -the
specific name is not given which work serious injury
in the orange orchards. A peculiar mode of protection
from these enemies is adopted by the proprietors, ap-
parently suggested by the fact that the injurious larvae
are preyed upon by certain ants native to the orangeries.
But these are not bred in sufficient numbers to be of
much practical advantage. Resort was therefore had
to the "hill-people" of the countries, who find the ant
nests suspended from the branches of the bamboo and
A. Synthetic figure exhibiting the entire course of the alimentary
canal in ants.
B to J compose a series illustrating the progressive distention
of the crop from its normal condition to that of the honey-bearer.
In C and F the crop is normal.
In B and E (workers-major or semi-rotunds) the distention is well
advanced. In F the crop has shrunk after distention. G shows
the same process in the abdomen of a worker-minor.
H. Abdomen of a honey-bearer, opened at a slit (s), to punc-
ture the crop, and show by its shrinking that it had filled the abdomi-
nal cavity. / shows the full crop of a honey-bearer with the lower
part of the alimentary canal shown through the abdominal wall
against which it is pressed.
J is the abdomen of a honey-bearer, wherein the full crop has
pressed the gizzard, stomach, etc., into the cloacal cavity. / and
J were apparently in normal health.
K is an abdomen of Camponotus inflatus, the Australian car-
penter ant, which exhibits the characteristic distention of At.
hortideorum. Drawn from alcoholic specimen. Other specimens
are quite spherical.
Key to lettering, uniform in all figures: ab, abdomen; ab pi d,
abdominal plate dorsal; ab pi v, abdominal plate ventral; an, anus;
be, buccal or mouth sac; col, colon; gz, gizzard; il, ileum; in, intestine;
m th, mesothorax; met th, metothorax; mpg, malpighian tubes; nd,
node; ce, oesophagus; pr th, prothorax; px, pharynx; re, rectum;
other trees. There are two varieties, a red and a yellow,
whose nests resemble small cotton bags. These are
captured by the Chinese mountaineers by means of pig
or goat bladders baited inside with lard. The mouths
of the bladders are stretched across the gates of the
ant-nests, and as the insects are fond of oils and greasy
food, they enter in, are trapped in great numbers, and
are sold at the orangeries. They are colonized upon the
trees by turning them loose upon the branches. Once
established, they begin their work as insecticides by
capturing and killing the destructive larvse. To enable
them to pass freely from tree to tree, all the trees of an
orchard are connected by bamboo rods.
Whether such a method is practicable in the United
States, at least to an extent to justify extensive use,
may be doubted. If successful at all, it would prob-
ably need the painstaking patience of Chinese men with
the Chinese ants. However, a somewhat similar experi-
ment has been tried upon a Ponerine ant (Ectatomma
tubercMlatum) , popularly known as the "Kelep." This
ant was imported into Texas by the United States De-
partment of Agriculture as an insecticide, with the
special purpose of directing its insect-destroying energies
against the cotton boll- weevil. It had shown marked
tendencies in that direction in its native Guatemala.
Apart from the more or less complete success of such
experiments, the fact remains, which is here relevant,
that it is one of a great army of ants that feed upon
That this habit is widely distributed among the native
ants of our Southern States was shown in a report made
by the author [McC. 9, p. 182] a number of years ago
to the then entomologist of the Department of Agri-
FEEDING THE COMMUNE
culture, Prof. J. H. Comstock. Several species therein
described were found to prey upon the eggs, the larva?,
and the pupa? of the cotton caterpillar over a wide
section of our southern territory.
It is not difficult to suggest a theory as to how the
taste for cereal foods may
have arisen among ants.
Whether it be the true one
or not is another matter.
Following their habit of gen-
eral scouting for supplies,
and of putting all promis-
ing objects to the test of
antennal or gustatory ap-
proval, they would be sure
to fall upon seeds in the milk
Stage. Being then soft and Fi g- 56 SECTIONAL VIEW OF THE
STOREROOMS OF THE OCCI-
easily crushed, and to the DENT ANT
ants a toothsome relish, all r Seed-rooms, rd Dumping-
grain-like seeds would soon rooms ^Gliierfes PebbleS '
commend themselves, and
easily pass into the accepted and fixed menu (Figs. 57
As the outer shell gradually hardened, the growing
taste for such food would prompt to break it open, and
so would come, little by little, the habit of removing the
husk. Although the flavor of the seed would change
with its ripening, one readily conceives that the taste
for it might have a corresponding gradual change; and
also the power of utilizing it for food by rasping off or
breaking up the starchy substance instead of crushing
and lapping it, as in the milk stage.
In quite the same way the use of nutty or oily seeds
would gradually form, aided by the natural emmet appe-
tite for animal and vegetable fats and oils of all sorts.
The tendency to carry all these seeds to the common
centre, the nest, would in clue course be followed bv,
first, the taking off and deporting the useless husk or
shell, and, next, the retaining for temporary, and so at
last storing (Fig. 56) for more permanent use, the grain
or nutty meat of the seed. The harvesting habit in ants
since it was first scientifically confirmed by Moggridge [Mg.
1] has excited an exceptional degree of interest and sur-
Fig. 57 OCCIDENT ANT HARVESTING SEEDS OF WILD SUNFLOWER
prise. But in truth, when one considers all the condi-
tions, the wonder is that it is not more widely distributed.
Here we may notice a peculiarity that appears in the
communes of a Texas species, PJieidole instabilis. A
study of its feeding habits presents a striking example
of a sharp distinction between the functions of the
soldier and the worker caste. The workers are much
FEEDING THE COMMUNE
the smaller but far more numerous, and the bulk of the
commune's work is done by them. They collect and
store seeds and dead insects. They dig rooms and gal-
leries, care for the huge royal larva? and pupa?, feed the
brood, and aid the callows break out from the pupal sac.
Fig. 58 POSTURES OF THE FLORIDA HARVESTER
IN CUTTING SEEDS FROM THE STEM
The soldiers do none of this work. They are the
communal trenchers. They crush and carve the tough
insects and hard seeds stored by workers, a service for
which their large muscular heads and jaws are well
fitted. The same organs are efficient weapons for de-
fence of the commune, a service which attaches to them
as soldiers. As such they may be seen, as if on sentinel
duty, surrounding the communal dependents. They
are stolid in temperament. They decline, in Indian
fashion, to take part in nursing the communal young.
Their big heads, though of such value to the commune,
may sometimes be a serious incumbrance to themselves;
for when dropped upon their backs on a polished sur-
face, they are not able to recover themselves, and may
die literally standing on their heads.
It seems to be an odd characteristic that these mega-
lacephalous creatures appear never to feed upon the
oily seeds and insect juices which they make available
by cracking the material brought in by the workers,
but live on liquid food regurgitated by the workers.
One must note in this a beneficent arrangement; for the
soldiers, nut being exposed to the temptation of feeding
directly upon the food which they carve for the workers,
and which must thus all pass through their "hands,"
make sure that the dependents are not stinted or
starved and the community thereby imperilled. This
characteristic seems all the more important in view of
the philoprogenitive defects of these soldiers. A species
of Pheidole is found in and around Philadelphia, and it
- - ^SffW.^W^.-^,^ :
.- ;;->:>,.<,. &&'
Fig. 59 SECTION PENNSYLVANIA HARVESTING ANT
Sectional cutting, showing storerooms, or granaries, in site, of
too is characterized by a big-headed worker caste. I
have made observations of its seed-storing habits (Fig.
59), but it remains to be learned whether the soldiers
have acquired so remarkable a role as that of communal
trenchermen. [McC. 4, p. 148.]
Some of the feeding habits of the Indian Leptogenys
are interesting. Writing of Lobopolata distinguenda, Mr.
Wroughton says [Wr. 1, pp. 50-58] that it is occasion-
FEEDING THE COMMUNE
ally seen going about solitary, probably when acting as
scout, but ordinarily is only met in the early morning
or late in the afternoon travelling in an unbroken column
four to six or eight abreast, by the straight or the easiest
road to the scene of operations. This is usually a colony
of termites, or white ants, whose galleries have been
broken open by the hoof of a passing beast or some like
accident. Apparently they do not have the initiative
faculty of breaking into the termites' nest, but wait for
an accidental opening. Arrived at their destination,
every ant seizes her termite prey, swings it under her
thorax in the usual way of these porters, and the attack-
ing column then moves homeward. But the return
formation is much less regular than the advance; it is,
in fact, a " march at ease."
The same writer gives a note on the allied species
Lobopelta chinensis. A populous community of this ant
had settled in a cavity of the house foundations of Mr.
Aitken, who reports the incident. From this nest there
ran a well-marked ant road which crossed a broad gravel
path and then branched out over the tennis-ground.
After sunset the workers would come out and march
along one of these branches, or break up into parties and
take different routes. Their point of approach was a
termite's nest; and when they reached a place where
these insects had thrown up new earthworks, and were
busy eating dead grass underneath, they collected in
dense masses, awaiting an opportunity to break in.
This came when the termites sought to extend their
works on any side. Then the waiting columns of ants
were precipitated in mass upon the unprotected creat-
ures, and the slaughter began. Sometimes the ter-
mites were killed faster than they could be carried off.
After one raid, as late as 7 A.M., the ground was still
heaped with the slain, and an unbroken stream of ants
fifty-six yards long was taking them away, every porter
having two or three of the dead in its jaws.
Sometimes the tables would be sharply turned upon
the plunderers. If they chanced to cross the territory
of a commune of harvesting ants after they had opened
their gates and were abroad on morning duty, the
Lobopelta hordes had to flee before their betters, often
abandoning their booty. Yet, per contra, the observer
once saw a Lobopelta, who had come to the aid of a com-
rade assaulted by a harvester, after vainly trying to tear
off the aggressor, deliberately pick up both comrade and
assailant, and carry them off together! Apparently
both were so intent upon the personal combat that they
gave no heed to the deportation.
Leptogenys elongata feeds largely upon the common
wood slaters (Omiscus and Armadillidium), which abound
under stones and logs in shady sites where the formi-
caries are placed. [W. 8, p. -253.] The workers have re-
peatedly been seen carrying dead slaters in their mandi-
bles, and the space surrounding the gates is white with
bleaching limbs and segments of the crustaceans, a proof
that great numbers of these animals must be destroyed
by the ants. Their long, toothless mandibles resemble
scissors, and are well adapted for piercing the inter-
segmental membranes of their prey and exposing edible
parts. This ant appears to be the only one known to
feed on crustaceans as a regular diet. Other species
are insectivorous, granivorous, mycetophagous (fungus
eating) feeders on the sweet, liquid excretions and secre-
tions of insects, or the juices and sugary exudations of
fruits, plants, and galls, and on animal fats and oils.
THE LANGUAGE OF AXTS AND OTHER INSECTS
LANGUAGE is essential to effective government
among social creatures. Without means of com-
munication of some sort, it would be impossible for
societies to hold together and to act together in those
communal movements which are alike the evidence and
the end of social organizations. Thus we infer that
some way of making known the common will and aim
must exist among such insects as ants, bees, wasps, and
termites that maintain permanent sodalities. And so
we find it in ant communes. Thus is preserved unity
and efficiency, by holding the citizens together; by dis-
seminating purposes and influences important to civic
success; and by securing at once mobility of action and
the concentrated force of the republic, for peaceful ser-
vice, for common defence, and for aggressive enterprise.
Men commonly think of language as a vocal medium
for conveying thought and emotion from one individual
to others. As thus defined, insects are dumb, for they
have no true voice nor organs of speech such as belong
to " articulate speaking men." They also lack the means
of uttering such cries as characterize birds and brutes.
But if we take language as simply an understandable
medium for expressing emotions, insects are thus en-
dowed. By certain movements of the body and of
parts of the body, especially the wings, antenna), and
jaws, and by sounds made by various organs in sundry
ways, they convey to one another the primitive and
simple emotions of their kind and of all animate beings.
In taking up the subject as it bears upon ants, we shall
best reach such conclusions as seem at present attain-
able by considering it in relation to insects in general.
The language of insects may be regarded as mimetic,
when emotions are expressed by gestures or acts; pter-
atic, when by wing vibrations; spir ocular, when made
known by sounds issuing from the breathing tubes or
spiracles; stridulatofy, when conveyed by the friction of
one organ against another; and antenna! , when the an-
tennae, or " feelers," are the media of communication.
Insects express emotion mimetically that is, by
bodily gestures. Mimetic language, though more limit-
ed in its ability to convey ideas, is not less intelligible
than vocal speech. Indeed, a glance of the eye, a move-
ment of the hand, a shrug of the shoulder, a stamp of the
foot, a toss of the head, may betray in man the true
thought or feeling within him, even when spoken language
is used to conceal it. How apt a medium mimetic
language may become for expressing clearly a wide
range of ideas one may see among the inmates of in-
stitutions for the deaf and dumb. We may find, per-
haps, that this medium serves insects no less effectively
for communication within that limited range of ideas,
shall we say? to which their faculties are confined.
Let us stand before this oak-tree and watch a double
stream of mound-making ants thronging up and passing
down the well-marked trail that leads to a herd of
aphides upon some oak-tree branches. The motion of a
finger near the trunk attracts the attention of a sentinel,
one of a number that seem to be guarding the flanks of
THE LANGUAGE OF ANTS
the column. It halts,
thrusts out its anten-
nae, and shows signs
of excitement. As an
experiment, the fin-
ger is moved slowly
within an inch or
more of the ant. Its
antennae wave rapid-
ly. Its head and body
jerk with eager in-
tentness. It stretches
forth its head, and
reaches out its fore
legs, with jaws eager-
ly agape and antennae
quivering (Fig. 60).
The whole attitude
and every bodily de-
tail clearly express to
the observer the idea
of vigilance, of suspi-
cion, of a challenge,
of a purpose to repel.
As plainly as if it had
spoken, the sentinel ! ;
has said: "I suspect
you! I test you! I
bid you begone !" We Fig. 60 MIMETIC LANGUAGE IN ANTS
Onlookers understand The S esture of repulsion and defence
this. Is it supposable that the ants themselves do not
From the tree-path we turn to the conical mound
whence these ants are issuing. It stands silent in the
shadow of the tall surrounding trees, its quietude broken
only by the movements of a few worker-ants, who are
lazily dumping pellets of soil from one of the few upper
ports. At the base of the cone, where most of the
gates are located, the column stretches across the grove
to the aphis-covered oak. Give the mound a sharp blow
with foot or hand. What a change! Instantly the
whole community is aroused. From every gate pours
forth a surging torrent of irate sentinels, followed bv
C_> O / /
other inmates, until, in an incredibly brief time, the
mound is covered with angry insects. They run to and
fro, their bodies a-quiver as they go. They challenge
one another with crossed antennae. They peer at every
unusual object in their way. They startle, and stand
rampant at the vibration of every sharp sound. The
surface fairly buzzes with the excited creaturelings,
their whole mien and attitude saying, unmistakably:
"Our home has been attacked! We are in danger! Rally
to the defence! Death to our enemies!" (Fig. 61).
We change the field of observation. The writer was
once standing before the great round web of a female
Orange Argiope (Argiope aurantium), a large and hand-
some orbweaving spider [McG. 24, pp. 97, 98, vol. i] ?
testing with a tuning-fork its sense of hearing, when a
bee flew by in exploitation of a flowering honeysuckle
vine that covered an arbor on which the web was hung.
The droning of its wing-strokes as it flitted from flower
to flower fell upon the ear as a token of content. To
all and sundry it said, or seemed to say, what no doubt it
felt: "I am a well-satisfied bee!"
But in a hapless moment it touched the spider's orb.
Its feet were entangled in the sticky threads. Straight-
THE LANGUAGE OF ANTS
way its wings began to move violently, and their buzz-
ing, together with the attitude of the body and of every
member thereof, expressed the creature's fear. These
varied as the bee became more thoroughly entangled,
< ,. ._.. v - *fij
Fig. 61 ANTS IN ATTITUDE OF
COURAGE, ANGER, AND ALAR.M
now waxing, now r waning, until the audible notes pro-
duced by its enfeebled movements seemed to utter its
growing sense of danger and dread.
Meanwhile, and the interval was rarely brief, Madam
Argiope underwent a striking change. She had been
enjoying peacefully a dejeune of cold Diptera, taken
through the fibres of a silken saclet in which her prey
was encased, and which hung upon the upper part of the
oval rug that overlay her central seat. But instantly
the bee struck the web her whole being was transformed.
She dropped her lunch-bag. Her reposeful attitude was
changed into one of eager animation and intense ferocity.
Every spine and bristle upon her legs, her body armature,
seemed to be erect, and her fangs were open. With a
rush, like the vault of a cat upon a sparrow, she charged
over her web, and, seizing the bee with her fore paws,
shot forth upon it from her expanded spinnerets a band
of silk. All the while revolving the insect between her
swiftly moving fore feet, she soon had it swathed as
closely as a mummy.
This done, her outward seeming of tense energy re-
laxed, and having suspended her empouched captive
to a twisted strand of her broken snare, she left it hang-
ing there, like a cured ham to the rafters of an old-
fashioned smoke-house, and quietly resumed her inter-
rupted luncheon on the fly. In all this, Argiope was
without speech; yet her varying emotions were plainly
and, one may conclude, not incorrectly, read by the
observer from her motions. As for the bee, its last
audible utterance was a low and broken hum that sound-
ed like the expiring wail of apian despair.
Now entered upon this tragic scene of animated nature,
man the philanthropist, his pity crossed, let it be con-
fessed, with a strain of curiosity to know the condition
of the prisoner. "Poor bee/ 3 quoth the observer,
"this great and greedy spicier has quite enough food
without you!" With a pair of pocket-scissors an open-
ing was made into the swathing-sac. Not without pro-
test from Argiope, however, who, feeling these move-
ments through the delicately strung meshes of her web,
feared that her prey was escaping, and rushed upon it.
THE LANGUAGE OF ANTS
She was turned back by a smart tap upon the head;
whereat she seized the tip of her rug and began to
oscillate her snare, as though to shake off an intruder.
When this diversion had quieted down, the scissors
were plied again. As the rent lengthened, the bee
seemed to awake and began to stir. One leg appeared,
then a wing. Thereupon issued a low hum of satisfac-
tion, which rose into a higher note, apparently of vexa-
tion, as the body gradually appeared. At length, with
a burst of limbs and wings, the insect was free. There
was no mistaking the character of her emotions now;
they were not jubilant. She was mad! and was waxing
madder in remembrance of the indignity put upon her.
Her wings vibrated with a velocity that raised their
responding sound to a high note which plainly signified
wrath and vengeance. The observer, at least, under-
stood; for instead of turning its wrath upon its captor,
the bee made straightway for its liberator with sting
outthrust, and with that peculiar buzz which bee-
familiars know as a war-note.
Discretion in that case "the better part of valor"
justified retreat. Moreover, the quest was not quite
ended. It had been determined that an insect can be
captured and swathed and trussed up by a spider with-
out impairment of aught but her temper. But it re-
mained to see what her beeship would do ; and that soon
appeared. Its pursuit of its back-stepping deliverer
ended, it turned again to the honeysuckle vine, and took
up its search for pollen and nectar as though life had
known no "hairbreadth 'scapes' 1 from deadly peril, and
timely rescue therefrom. Her war-note died awav into
the old droning hum of peaceful industry and busy
Here we have a series of actions by which two in-
vertebrates clearly communicated their emotions. The
spider passed rapidly through stages reaching from
quiet enjoyment of food to intense passion of the chase
and ferocity in capture, and to the repose of success
when the prey was secured. Thence the course swung
to rearoused energies under apprehension of loss, and
to fear of some unknown superior foe when rapped by
the observer, and anxiety to defend herself therefrom,
as shown by shaking her web.
The bee, too, had swift transitions: from her hum of
contented industry to the subdued note of resignation
to her fate when shut up in her silken sarcophagus;
thence to vivid reawakening to life, with her sense of
injury, her blind wrath and revenge, the wish to strike
at something; and so back to where the cycle began:
at the song of peaceful labor. In all these stages these
children of the wild betraved their current moods to
man. "There was speech in their dumbness, language
in their very gestures. " No careful observer of their
natural actions and of the field-life of their kind can
doubt that, within limits indefinite and difficult to define,
like actions among the more highly organized insects
are understood bv one another.
Still further, it does not seem probable that the ability
thus to make known their emotions is limited to such
modes of expression as human intelligence can interpret.
Beyond the sphere of ideas and sentiments whose sym-
bols men can discern, there doubtless are others peculiar
to themselves, and therewith due methods of inter-
In the cases above cited the actions may be said to
have been simply the unconscious physical expression
THE LANGUAGE OF ANTS
of natural animal impulses, without any purpose to
communicate the same to another, such as language
implies. Even so, it should be considered, first, that
these examples are given as types of other uses of
mimetic language behind which lies the undoubted pur-
pose to communicate. And, second, that the rude
evolutionary germs of language in primitive man may
have been the utterance of just such impulses; and little
more need be claimed for insects. It marks the im-
passable difference between the psychic powers of man
and those of insects that human language, spoken and
written, has developed into its marvellous proportions,
while the symbolism of insects, and of animals generally,
retains the crudity of ancestral types, and apparently
can never pass beyond this bar of nature.
There was something more in this typical living
tableau of the spider and the bee than " gesture lan-
guage"; for the wing movements of the bee, as we shall
presently note, were special media of communication.
But the language of natural bodily motions may claim
some further attention here. If an unarmed man be
threatened by his fellow, his almost unconscious mode
of expressing his feelings will be to dodge or crouch or
flee, if he be afraid; or if he be brave and his combative-
ness be aroused, to throw himself back upon one leg
and put up his fists in self-defense. Under like condi-
tions a bear will rise upon its hams and extend its fore
paws, and a horse will rear upon his hind legs and strike
out with the fore legs and hoofs.
It is a long step from the primate, the ungulate, and
the ruminant to the invertebrate. But let us present sim-
ilar conditions to certain spiders say, the " tarantula' 1
of the southwestern United States. It takes a rampant
position, resting upon its two pairs of hind legs, while
its two front pairs, palps, and fangs are thrown up in
striking posture. [McC. 24, vol. ii, p. 320.] The same
attitude may be seen in the little jumping spiders (At-
tidae) around our house walls and vines.
From the tarantula turn to the stream of agricultural
ants of Texas, pouring over the roads that lead into their
harvest fields. Fix your eye upon this worker, returning
home carrying a grain of ant rice. Every motion of her
body, which fairly palpitates as she hastens on, shows
her sense of importance and satisfaction in service.
Now tap her with your pencil-point. What a transition !
She instantly stops, drops her burden, and rises rampant,
the fore part of her erect person declaring unmistakably
that she is startled, angry, and means to fight. She
thus takes her place as a link in the chain of life leading
down from man, among the creatures that communicate
their belligerent mood and purpose by bodily attitude
But something more than signals and gestures appeal-
ing to the eyes met the observer of that affair between
the orb weaver and the bee among the hone3 r suckle
blooms. The bee's wings made effective appeal to his
ears, and by their varying vibrations gave a fair token
of her tempers. This was " pteratic language.' 1 The
droning among the flowers, the quivering amid the
spider's meshes, the sharp buzzing of flight after release
sounded in unmistakable notes the insect's amiability,
anxiety, or anger. One can detect these varying notes
as he walks his garden and field while the bees are forag-
ing among the flowers or while one watches by his bee-
hives. So, mayhap, Shakespeare did near by Anne
Hat ha way's door, or while treading the pathway across
THE LANGUAGE OF ANTS
the fields from Avon to her cottage gate, and saw the
busy workers, like raiding soldiers,
"Make boot upon the summer's velvet buds;
The singing masons building roofs of gold."
Gardner, an English writer on the Music of Nature
(1832), makes the curious statement that he was once
in the gallery of the Royal Exchange to view the money-
dealers in the court below. He was struck not only by
the likeness of the scene to the interior of a beehive,
but by the similarity of the sound, the buzz of the two
thousand voices being perceptibly amalgamated into the
"key of F." This is the key, the author concluded, to
which the most prevalent sounds of nature may be re-
ferred a fact by which musicians have unconsciously
been influenced; for scarcely an ancient composition ap-
pears in any other key, except its relative minor, for
the first hundred years of the art. In Queen Elizabeth's
Virginal Book of four hundred folio pages nearly all the
pieces are confined to this key. There is not an instance
of a sharp being placed at the clef. 1
According to the same author, the house-fly and the
honey-bee hum in F on the first space. The bumble-
bee, the contra-basso of the tribe, performs the same
note, but an octave lower. The present writer is able
to confirm this conclusion only in part. F seems to
him to be a nearly true note for the common fly as
tested by his ear, unaided by an instrument. But the
wing-note of bees and the general tone of a large mis-
cellaneous company of insects humming above a bed
of flowers hydrangeas, for example seemed to him
1 I have not the opportunity to verify this statement, which I
make on Gardner's authority.
to bo A, us tested by the flute as well as by the ear
Recently the vibration of insects' wings their
pteratic language has boon studied from the character
of the note caused thereby, the pitch determining the
Fig;. 62 PTERATIC LANGUAGE
The author, with his flute, testing the keynote of wing-strokes
as insects are humming over flowers
THE LANGUAGE OF ANTS
number of vibrations on the basis of two hundred and
fifty-six per second for the note C. 1 Tuning-forks are
perhaps the most convenient instruments for such ex-
periments, which may be made by any one who has an
ordinarily ^ood ear for musical sounds. The writer
has used his flute with some measure of success. The
house-fly has a wing-tone of F, or three hundred and fifty-
two vibrations per second. The honey-bee strikes A,
which means that it moves its wings at the rate of four
hundred and forty times a second. When, burdened
with its weight of pollen, the bee is on its homestretch,
its wing-tone falls to E, indicating three hundred and
thirty vibrations a second.
An interesting confirmation of these results has been
made by fixing a fly within a carbonized cylinder re-
volved by clock-work. The tips of the fly's agitated
wings left at every stroke a slight mark upon the smoked
surface of the glass, which, being counted, gave sub-
stantially the same result as above viz., three hundred
and thirty wing-strokes a second.
To be sure, such tones as these may be held to be a
mere mechanical product or reflex; yet that they have
the power to express certain ideas will be clear to one
who will observe the effect produced upon a community
of bees or hornets by the buzzing of one of their number
when angry. The excitement runs rapidly from one to
another, until many members are visibly affected. The
original irate had certainly communicated her mood
to her fellows.
Even insects of alien species seem to understand such
wing-stroke language. Let an angry hornet or yellow T -
1 The most extensive studies in this interesting field are those of
the veteran entomoloo-ist, Samuel C. Scudder.
jacket course the suburbs of a populous ant-hill, and the
knowledge of her temper will be conveyed to the ants,
who apparently understand that a highly keyed note is
a threat which they must needs resent. It is a question
how such information is conveyed; but perhaps, like a
coursing motor-car, the intruder may give forth not only
a hostile note but a pernicious smell !
Before entering further upon this theme, it behooves
both writer and reader to remember not only the vast
gulf which separates us from insects as well as the com-
mon bonds of nature that unite us to them. Ento-
mologists have already disclosed much of the real life
of the lowly creatures that share with us the earth;
but we have as yet scarcely passed beyond the threshold
of the temple of knowledge that Nature has reared around
us. Many problems that have barely been stated re-
main unsolved or partly solved, though our scant
knowledge might be far more complete "would men
observingly distil it out." Innumerable other problems
doubtless are beyond the screen, duly to rise as the
horizon of discovery shall enlarge.
What know we, for example, beyond the narrowest
bounds, of the senses of ants of their number, their
quality, their range? What know we of the endless
degrees of sounds and shades of color that may form the
world within which insects move, familiar to them, but
a terra incognita to us? May there not be a Nature
within our known Nature, worlds within our knowable
world like the successive enclosures within a Chinese 1
"nest* of boxes of which insects know, and wherein
may be their largest moiety of life? To them a wild
meadow, a flower-garden, a grove, or a brook-side may
be a boundless scene of beauty and activity, friendly
THE LANGUAGE OF ANTS
and hostile, such as we might depict as a fairyland.
Therein may be landscapes hidden from our eyes, with
many grades of color, fair or grewsome, and octaves
of sounds, pleasant or fearsome, that lie beyond human
senses or even human fancy. Of this world, or these
world-spheres, much must remain unknown to us at
least, in this cumbered state of existence. But to pene-
trate it further and further, to unlock its secret doors,
to explore and disenchant its chambers of mysteries,
and to interpret to mankind its inarticulate symbols
this is the highest function of the true natural history.
HOW AXTS COMMUNICATE
TT^E are not yet done with our typical bee's capability
f T to express her current feelings. She can resort
to spiracular language. When deprived of the use of
wings by the spider's entanglements, she still made her
plaintive, or petulant, or wrathful protest through the
spiracles. These are breathing organs arranged in pairs
along the abdomen and thorax of insects. Behind each
spiracle is a membrane, or chitinous projection, which
is agitated during breathing, and may be set vibrating
so rapidly as to produce a sound. This, for lack of a
fitter word, has been called a "voice," and certainly
suggests the product of the vocal chords in man. It
comes as near to being a true voice as we are likely to
find among insects, and perhaps the diminutive "voice-
let" might be applied to it not inaptly.
One need not be a naturalist to satisfy himself of its
presence. Let a house-fly be held by its two wings-
our bee being hardly available for such an experiment
for ordinary observers and there will be heard a high-
toned buzzing which manifestly is not made by the
wings. It issues from the spiracles, and is the insect
voicelet. The same note inav be heard from the un-
happy victims of fly-paper, who, though their wings
are held in the grip of the sticky compound, continue to
send out a pitiful cry from their spiracles. The same wail
HOW ANTS COMMUNICATE
or shriek may be heard from the unfortunate creatures
whose wings have been burned off in a lamp or candle.
Another familiar member of the Diptera has the
faculty of voicing the mosquito, with " blood-extract-
ing bill and filmy wing." The " honest" mosquito, that
blows her shrill pipette to warn of her approach, pro-
duces her peculiar note by the use of her spiracles. It
is doubtful if her chivalry in giving her chosen prey
a chance for defence is appreciated. To many, the
soundless sort that fall with the silence of death or
" sable- vested night," and go straight to their phle-
botomy and make no fuss thereabout, are the less
pernicious of the two.
The bee's spiracular voicing is known among bee-
keepers as " piping." The senior Huber [Hu. 1, p. 157]
published the first intelligent account of it. When the
old queen of a colony has left with a swarm, the new
queen is sometimes seized with a fancy to sound her
pipes, standing, while doing so, with her thorax against
a honeycomb, and her wings crossed on her back in
motion, but without being unfolded. The sound has a
remarkable effect upon the workers, who, with their
faces toward the queen, lower their heads and remain
motionless, as though smitten by some strange charm,
and listen intently. The young queenlings, still within
their cells, perceive the sound through the waxen walls
that confine them, and respond thereto with what seem
to be notes of defiance and challenge.
Even without such stimulus, the queenlings within the
royal cells, while waiting to be freed by the workers,
will play their pipes. The sound emitted Huber de-
scribed as very distinct, a sort of clacking, consisting of
several monotonous notes in rapid succession. He con-
jectures that the use of this piping, in the economy
of Nature, is to give notice that the young queen is ready
to be released an office which the workers keep well in
their own hands, in view of the instinctive tendency of
all apian royalties to destroy one another, and enjoy,
through regicide, an undisputed reign. Whatever be
the purpose of this ceremonial song, the fact is patent
that piping is a mode of communicating certain emotions
well understood by both queens and workers, and there-
fore serves the end of language. The worker-bees, upon
the adoption of a stranger queen, will gather in a series
of circles around the newly installed sovereign, and,
staidly vibrating their wings, sound a sort of coronation
anthem, which, as it appears to issue from the spiracles
as well as wings, may be classed with the piping of
queens. [Hu. 1, p. 107.]
It is strange that an act which should have the bene-
fit of the community in view should open a way to dis-
aster. Yet so it appears. Huber was greatly disturbed
by the ravages of an unknown enemy among his hive-
bees. At last the invader was found to be a large moth
(Sphinx atropos], popularly known as the " death's-head
moth " (Fig. 63), from certain body-markings that rudely
resemble a skull and cross-bones. Experiments showed
that the bees have ample power to defend themselves
against this moth. In the case of the bumble-bee the
power is used to sting it to death, those children of the
wild being less open perhaps to the seductions of musical
enchantments than hive-bees with their more artificial
habits. How could a moth, destitute of natural force
and weapons fit to cope with throngs of insects which
can repulse a man, manage to cow them or charm them,
and thus safely plunder their homes?
HOW ANTS COMMUNICATE
Huber's suggestion, which has been supported by
other observers, was that Atropos has the gift of making
a sound so like the " piping ' : of queen bees that the
workers are deceived thereby, and stand inactive and
Fig. 63 DEATH'S-HEAD MOTH WHICH MIMICS THE PIPING
OF A QUEEN BEE
seemingly fascinated, as is their wont under the real
royal notes, while the moth works its will in their house-
hold. [Hu. 1, p. 312.] How strangely this bit of natural
history suggests the stories of witch and enchantress
that were wont to awe our credulous ancestors! Or,
much to its advantage, did Atropos thus prove that
"music hath charms to soothe the savage breast"? But
the "music," like that which satisfies the average savage,
would hardly charm an Asaph,or aHaydn,or aBeethoven;
for it is simply the grating sound produced by rubbing
the palps against the base of the proboscis. But, then,
the bees are not dainty in their musical taste, as witness
our boyhood's recollection of a throng of excited villagers
following a swarm of bees across the fields, with jangling
of cow-bells and clanging of tin pans, moved by the tradi-
tional faith that bees would thus be charmed to " settle."
Thus we are brought to another form of insect lan-
guage stridulation. Our typical bee, unlike the Atropos
moth, is not gifted in this wise. But the art is possessed
by some spiders, and one species, akin to the tarantula,
gets therefrom her specific name stridulans. The in-
sect music with which we are most familiar is thus
caused. The organs which produce the various notes
are built on the principle of the violin and mandolin.
In other words, they are the result of regulated friction,
though the degree of regulation is crude and limited.
Take, for example, the grasshopper, whose shrilling
is one of our well-known autumn field-notes. On the
inner side of the thigh is a series of fine cogs, or teeth,
which one can see with the naked eye or with a hand-
lens. These, rubbed rapidly against the wing-covers, as
one might rub a file against a goose-quill, cause the grass-
hopper's rather cheerful chirrup (Fig. 64).
Brunelli, an observer of the eighteenth century, con-
fined in a closet a bevy of male grasshoppers (Gryllus
viridissimus) , who proved quite philosophical prisoners;
for instead of sulking, they kept up a merry fiddling all
the day. A rap at the door at once stopped their note;
but an imitation of their chirruping, which the naturalist
managed to make fairly well, brought a low response
from a few, which soon swelled into a chorus by the
HOW ANTS COMMUNICATE
whole group. One of the males was shut up in a cage
in the garden, and a female captive was set at liberty
near by. Soon the male put his mandolin into play,
whereat Madame Gry 11 us flew to his side. " Barkis was
woolin' -and Peggotty, too ! Certainly here was a case
c - c CU ( :.( ..( ( (( f
Fig. 64 STRIDULATING ORGANS OF LOCUST AND CRICKET
RED-LEGGED LOCUST (MELANOPLUS FEMUR-RUBRUM)
Below it is shown a greatly enlarged section of the file on wing-cover of male cricket, against
which the scraper on opposite wing-cover is rubbed to produce its call
of intelligent communication between two lovers, and
that by means of sound, and not by scent alone. And
it may be that for the most part this form of insect lan-
guage is amative. So, doubtless, much of human speech
was evolved around sexual and parental loves.
But Brunelli was preceded at least eight centuries in
discovering that caged grasshoppers will utter their
stridulant notes. According to the late Prof. Lafcadio
Hearn, 1 the Japanese, as long ago as the tenth century,
were addicted to their interesting habit of confining
insects in cages for the sake of their music. To-day the
sale of these insects and the dainty cages 2 in which they
are kept is a large and lucrative business in Tokio and
1 Exotics and Retrospectives, pp. 39-79.
2 The writer is indebted for the two ingenious specimens here
figured to Mr, Lucien Sharpe.
other Japanese towns (Fig. 65). To that remarkable
people the shrilling of crickets and grasshoppers seems
to be as sweet a sound as the song of canaries to us.
One who deems this a barbarous fancy may be remind-
ed that the men of classical Greece held the cicada to
be sacred to the deitv of music.
One finds such insect musicians as charm the Japanese
everywhere around him in the fields during late summer
and early autumn. Sitting here, writing, on the open
porch of his country home, the author hears the notes
of hosts of insects beating upon the hot noon air. Wild
bees, yellow-jackets, brown wasps, and blue mud-daubers
keep up a ceaseless hum as they hover over a flowering
Fig. 65 JAPANESE CAGES FOR STRIDULATING INSECTS
vine that drapes and shades the railing. Just overhead
hangs a fragrant clematis, among whose leaves a tree-
cricket plays hide-and-seek with the writer, and inter-
jects an occasional high-keyed Krea k I kr-reak! Out of
HOW ANTS COMMUNICATE
the grove issues the cicada's rolling call, swelling in
volume and dying away, and not well ended till an
answering or another trill is heard. And so, on and on-
beech-tree responding to maple and chestnut to w^hite
oak, with hardly an interval of silence. When night
falls "the katvdid works her chromatic reed," not
indeed "on the walnut-tree over the well/ 3 but on the
beeches and oaks, beneath whose branches wind the wood-
drive and the ramble. All these and others, with organs
varying in structure, as is the wont of versatile nature, are
the product of insect stridulation. 1 And could one tune
his ear to the finer sounds with which the occult spheres
of cosmos are full, he would hear many like sounds.
Ants, for instance, are supplied with stridulating
organs, which, reasoning from analogy, they must use
as means of expressing certain feelings. Yet one of the
rarest events in insect ethology is the record of an emmet
stridulation unless, indeed, the rasping noise one
hears issuing from the excited hordes of a disturbed
ant-hill may be the aggregate of many stridulators
instead (as conjectured) of the clatter of numerous
mandibles and the grating of chitinous body shells as
they rub against one another. The writer was long in-
clined to the latter view, although more than a quarter
of a century ago, in his studies of the honey-ants of the
Garden of the Gods, he showed that ants possess organs
well fitted to produce stridulatory sounds, and cited
at least one case that seemed to prove such use thereof.
[McC. 4, p. 07.] But the evidence now in hand puts
beyond doubt the existence of the habit.
1 Rubbing the femora or the wing-covers together, and rubbing
the bases of the two wing-covers (tegmina) together, are the chief
modes of stridulating among locusts, grasshoppers, and crickets.
However, the ordinary listener should not be dis-
appointed if he fail to hear these stridulatory notes, so
delicate and faint are they. They belong to that occult
realm of sights and sounds into which few of the more
highly organized forms of life are privileged to enter,
and not to the great company of insect musicians who
fill our summer and early autumn fields and woods with
their varied orchestration.
One who carefully observes the abdomen of a large
ant, even with the naked eye, can see that it is made up
of segmental plates, five above (dorsal) and five below
(ventral). These plates are imbricated that is, they
overlap one another, like tiles on a house-roof. They are
composed of epithelial scales, hexagonal in form, which
present a beautiful appearance, as of delicate mosaics,
when viewed through a microscope. When a profile
view of one of these scales is exposed to the lens the
serrate edge is clearly seen. Thus it is plain that a
backward and forward motion of the plates upon one
another might produce a faint rasping sound. All that
is required for the complete conditions for stridulation is
the muscular ability to perform this action rapidly (Fig.
06). Ants certainly possess this; and, in fact, they may
be seen thus moving the abdominal plates in and out,
back and forth, with a rapidity that seems to increase
with their excitement. The many faint sounds thus
made, inaudible in the individual, but audible in the aggre-
gate, would account in part, at least for the peculiar
hiss-z-z-z which arises from an excited column or colony
of ants. It will also help to explain the popular belief
that one sometimes picks up in rural parts, that u ants
sing." Besides this grating of the abdominal plates over
one another, there is a rotary movement of the base of
HOW ANTS COMMUNICATE
the abdomen upon the post-petiole which produces the
same effect, probably even more generally than the
Professor Wheeler believes that stridulation is an
important means of communication ---at least among
Fig. 66 PROBABLE STRIDULATING ORGANS ON ABDOMINAL
PLATES OF ANTS. (MAGNIFIED SECTIONAL VIEWS)
E. i. .<?. Epithelium; imbricated, serrate edge. i. ab. pi. Interior
of abdominal plate. e. ab. pi. Exterior of abdominal plate
such ant families as the Mrymicince, Ponerinse, and
Dorylina3. To this he attributes the rapid congregation
of ants when particles of food are discovered by errant
members of their community. The pleased sensation
of falling upon food is apt to start an ant a-stridulating,
and thus other foragers abroad in the vicinage are
attracted by the food-call. This also explains, in part,
the rapid spread of the heroic rage to defend their home
which runs through a populous ant city and calls out a
legion of eager sentinels and workers.
Stridulation also accounts for the ease with which
members of such species as the agricultural ant of Texas
are trapped by sinking a glass jar or bottle on or near their
formicary. One ant falls in, and begins to stridulate.
The sound attracts passing comrades, who throw them-
selves over the rim to the rescue, and in turn, finding
themselves imprisoned, begin to stridulate, until at last
so many are sounding the alarm that the chorus is
audible even to the human ear.
If, now, the jar be corked and shaken to further excite
the inmates, and then held over another Pogonomyr-
mex commune, whose members are peacefully sauntering
about, the wildest excitement suddenly seizes them, as
though there had been a call to arms. The writer has
collected these Texas ants by this method, but such a
reasonable explanation for the clatter within the bottle
did not occur to him until suggested by Professor
More decisive than the above, and it is conclusive, is
the description of the remarkable stridulation practised
by the leaf-cutting ants ( A tta fervens) of Texas. Herein
the different forms from the huge females, through
males, the large-headed soldiers, and the diminishing
castes of workers, down to the tiny minims present a
sliding scale of audibility. The rasping stridulation of
the queen can be heard when she is held a foot or more
HOW A.XTS COMMUNICATE
from the ear. The male and soldier, to be heard, must be
held somewhat closer, and the worker-majors still closer.
The smaller workers and minims, though stridulating,
as may be seen from the movements of the abdomen on
the post-petiole, are quite inaudible to the human ear.
We may safely join in the inference that "it is not at
all improbable that all this differentiation in pitch,
correlated as it is with a differentiation in the size and
functions of the various members of the colony, is a very
important factor in the co-operation of these insects,
and of ants in general. " [W. 4, p. 11.]
Such a condition, of course, implies that this stridula-
tory language is "heard" -that is, produces an effect
analogous to that of hearing. No auditory organs have
yet been discovered in ants with such positiveness as to
establish their existence beyond question, although
minute bodies within the tip of the antennre are believed
by some observers to serve as such in part. But that
their behavior under certain conditions is quite con-
stantly precisely what one would expect were ants known
to have ears, or their equivalent, is easily seen. Whether
they hear or not, they respond to sounds in a way cor-
responding to the acts of creatures that have ears, and
do undoubtedly hear. It is manifest that this must
have a vital effect upon the government of ant communes
to which, as in all other governments, some means of
intercommunication are essential.
Last of all, and perhaps most important of all, as a
means of intercommunication, is antermal language.
Ants, in common with most insects, are provided with a
pair of peculiar organs known as antennne, located upon
the face, above the mouth and midway bet \veen the eyes.
Externally these are thread-like rods of greater or less
length and thickness, jointed, and articulated upon the
face to increase their flexibility. In ants they consist
of two parts: the scape, a single piece that unites them
to the head, and the rlagellum, composed of a number
of segments ending ordinarily in a bulbous tip.
The olfactory sense has its seat in the antennae, usually
in the flagellum or the pore-plates and olfactory rods
thereof. While ants are sleeping, as observed in my
artificial formicaries, the antenna) have a gentle, quiver-
ing, apparently involuntary movement almost like the
regularity of breathing. [McC. 3, p. 134.] It seems as
if these sentinel organs keep on duty even during sleep,
guarding the approaches to their unconscious possessor
Livingstone [Li. 1, p. 576] gives a good example of the
dependence of ants upon the sense of smell as lodged in
Fig. 67 THE FACE OF AN ANT, SHOWING THE FLEXIBLE ANTENNA
the antennaB. He states that certain African species,
which he designates as " soldier ants," when on their
pillaging excursions, if their trail be covered with soap
and water or with fresh earth, will halt in apparent
confusion, and the succeeding ranks will mass in great
numbers at the point of stoppage. Meanwhile their
HOW ANTS COMMUNICATE
"leaders," who seem to act as scouts in scenting the
trail, will diverge from the column, flank the obstruction,
and recover the trail. Whereupon the main body move
around the tainted section and proceed upon their
In the antennae of ants are concentrated a great degree
of diverse sensibilities. The sense of hearing (probably
in whole); a large measure of the function of sight as it
exists in other insects and higher animals; the faculty
of communication (language) all seem to be located in
the antenme. They probably surpass in sensibility any-
thing at the command of higher animals, or even of man.
[C. 1, p. 210.] They are not only the prominent guiding
organs, as in insects generally, but are rendered peculiar-
ly sensitive by the addition of delicate hairs, some of
them highly specialized, spread over their surface.
Besides, they are articulated to a degree which gives
great flexibility and permits a variety of movements in
The removal of the antenme produces an extraordinary
disturbance in an ant's intelligence. It can no longer
find its way or recognize companions. It ceases from
its usual tasks, from seeking food, and from caring for
the larvae. Its condition recalls descriptions of the
consequences of removing the frontal lobes from the
brain of higher animals. The analogy is so close as to
suggest that the antennae of ants, with a brain adequate
for their functional requirements, are equivalent to the
most important parts of the brain of higher animals
working with inferior sensory organs. Such organs of
sensibility as those possessed by ants are a specialty in
sensory organism a conspicuous illustration of sensory
power concentrated in an external organ. "Their loss
is equivalent to the destruction of the mental faculties."
[Ro. 1, p. 142.]
Darwin's remark, that u the brain of the ant is one
of the most marvellous atoms in the world, perhaps
more so than the brain of a man/ must be taken
with a measure of qualification; at least, the word
" brain/' as used by him, must be understood in con-
nection with the antennae, the special organs of the
sensori - motor system of which the brain is only
the governing centre.
We may remark in passing that, in considering the
intelligence of ants, the sensory organs afford the key
to the situation. They are the real test of intelligence
or power of discrimination. They are a specialty in
animal life, and, as remarked by Professor Calderwood
"the ant's intelligence is in keeping with the recognized
functions of the organs of discrimination at its com-
mand." [C. 1, p. 216.]
The popular name of antenna? is " feelers," and it is a
quite fitting one; for when ants are awake and in
action these organs are kept continually revolving in
front of them and on either side of their path, touching
the various objects met, and sweeping the foreground
as though to feel the way. That, in fact, is their chief
use; they are feelers. With ants they are, perhaps,
even more important than the eyes for personal locomo-
tion and service and communal action. They determine
the forms of objects; they locate the individual trail
and the path of its fellows ; they distinguish foes from
friends; they test the quality of food and of all other
bodies, both by their odor and by their tactual reflex;
they give definite perceptions of space relations, and
thus enable insects, while moving over the ground sur-
HOW ANTS COMMUNICATE
face and through the ground closure, to orient them-
selves, and make sense record for subsequent use as a
rudimentary sort of memory.
It is this remarkable structure that so highly qualifies
the antennae for the function as the chief organ of com-
Fig. 68 THE CHALLENGE WITH CROSSED ANTENNAE
munication between ants. One cannot observe a colony
or a moving column of ants for any length of time with-
out seeing how constantly the crossed antennae are
used, obviously for communication. On the great dome
of the Alleghany mound-makers, and on their tree-paths
and the trails leading thereto, sentinels reach out their
antennae in challenge, and receive in the same way a
response. Two errant foragers meeting on the hunting-
field invariably cross antennae (Fig. 68). Going or com-
ing, leaving home or returning, on the city premises or
afield, it is always the same. One reads at once from
the manner the mutual "All right !" which passes.
Whether conveyed by odor or by contact or by both,
one may not affirm. But that it is conveyed, he read-
Two battling armies of the pavement ant (Tetramorium
coespitum) are massed on the edge of a flagstone walk.
A column of highly excited warriors is streaming from
the scene of action to the home nest of one of the com-
batants, from which issues a group of peaceful citizens.
The two lines meet. Antenna are crossed, a quick,
sharp action, and the messengers for such they are-
hasten on homeward. Note the result. Some myste-
rious influence has passed from one to another. The
peaceful citizens are transformed into combatants raging
with the lust of war, and with every bodily member
a-quiver, rush into the thick of the battle hurly-burly
to reinforce their comrades. Was there not antennal
communication between couriers and recruits?
Here is a case recorded by Lord Avebury [Av. 1, p. 75],
which strikingly illustrates this function of antenna?.
A worker of the dark Lasius (Lasius niger) was occupied
in carrying off larvae to her nest. At night she was
imprisoned, and, being released at 6.15 A.M., immediately
resumed her occupation. At 9 A.M. she was again im-
prisoned until 4.40 P.M., when she was put once more
to the larvae. She examined them carefully, but went
home empty handed. At this time no other ants Were
outside the nests. In less than a minute the original
worker, which had been marked with a dot of paint,
came out with eight friends, and all trooped off toward
the heap of larvae.
When they had gone two-thirds of the way the
marked Lasius was imprisoned, whereupon the others
hesitated for a few minutes and then returned home.
They evidently missed their leader's guidance. At
HOW ANTS COMMUNICATE
5.15 P.M. the marked ant was again put to the larvae.
Once more she went home empty handed, and, after only
a few seconds' stay, came out of the nest with thirteen
friends, and all went toward the larvae.
In this case the twenty -one ants must have been
brought out by the marked one, for they came exactly
with her, and no others were out. Moreover, they must
have been told, since in neither trip did she bring a
larva with her, and the sight thereof could not have led to
her being trailed, though of course it is just possible that
a faint odor clinging to her from contact with the larvae
may have given a signal that caused her to be followed.
Good examples of antennal parley occur in the preda-
tory expeditions of slave-making ants. From the nests
of the sanguine slave-maker (Formica sanguinea-rubi-
cunda), studied in New Jersey, scouts were seen to be sent
out to discover and locate the nests .of the SchaufTuss
and Fuscous ants (Formica Schauflusi and Formica
subsericea). When these \vere successful, they hastened
home with their antennal message, and soon the plun-
dering raid was in full heat. Forel describes the same
conduct as habitual with the amazon ants (Polyergus
rufescens) of Switzerland. Moreover, he relates that in
case of uncertainty as to the right route, the column
will halt, and wait \vhile the scouts go forward and locate
the nest of the predestined victims. Returning, the
antennae play vigorously between couriers and column,
and the piratical excursion advances.
One example more. At Faisons, North Carolina, Doc-
tor Forel, during his visit to the United States, found in a
rotten log a nest of the totally blind little Eciton Carolin-
ense. He captured the colony, and put it under observa-
tion. The ability of these ants to find their way about
rapidly and unanimously in new territory without one
estray seemed almost incredible. They were transport-
ed to Washington, where a handful of them, with their
young, was thrown into an open garden. Without losing
a moment's time the wee blind creatures began to form
in files, which were fully organized in five minutes.
Tapping the ground continually with their antennae,
they took up their larvae and moved away in good order,
like well -drilled soldiers, reconnoitring the strange
region into which they had been cast. Every pebble,
plant, crevice was tested with the antennae, and the
place best suited for concealing their young was soon
found. Most other ants would probably have been an
hour in accomplishing this. The experiment was re-
peated twice with the same results.
The marvellous fact in this action is the certainty and
quickness with which the "topochemical trail' -to use
ForePs phrase and the company relationships were
recognized. The groping about, and wandering to and
fro, and hesitant mien, common to ants when first placed
in strange sites, were wanting. We are prepared, there-
fore, to learn that the antennae of these Ecitons are highly
developed. And we share the interest of the learned
observer who watched the perpetuum mobile of those
organs, as in the most lively manner they kept titillating
the ground, their companions, and all surrounding
FEMALE GOVERNMENT IN ANT COMMUNITIES
THUS far our studies have been chiefly of the ex-
terior of the ant commune. We are now to take
a view of its internal economy. The reader may have
noticed that the author, in referring to the insects under
observation, gradually passed from pronouns of the
neuter to those of the feminine gender or, rather, has
used the two interchangeably. This accords with facts.
The worker ant, although in common parlance a "neuter,"
is structurally a female. In her the special function of
the female, to produce the eggs from which the young
are reared, has been subordinated, though not wholly
atrophied ; for workers occasionally drop eggs, which are
cared for, and which yield males.
Other faculties have been developed or have appeared
needful for communal safety and prosperity, and thus
it has come about that the government of these emmet
societies, as with bees, hornets, and wasps, is really a
gynarchy, or government by females. Our worker
ants are veritable Amazons. Not only does the entire
domestic control and service of community fall to them,
but also those more virile acts (according to human
standards) of war and public discipline and defence.
To the fact that the female temperament dominates
affairs we may perhaps attribute many of the charac-
teristics of public administration among social insects.
That there is a "female temperament," sharply dis-
tinguished from that of the male, is obvious enough to
the student of emmet habits. That its dominance is
advantageous to these organizations the natural history
of the Hymenoptera attests.
What would be the effect upon human societies should
similar conditions prevail among them? As a specula-
tive theory it is worth discussing, and one would hardly
err in thinking that our public and official affairs would
be greatly bettered could woman's temperamental view
of things have wider influence therein, especially in their
relations to the young. Our civil governments and their
administration, from the township to the national capital,
are almost wholly products of the male element of the
race. The predominance of the female element, which
one sees in ant communes, might not be desirable in our
present stage of civilization, although it would be an
interesting experiment in a county or even in a State.
Such illustrations as the United States presents throw
little light upon the problem, for the general conditions
of society in the States that give woman the suffrage
really differ little from those prevailing elsewhere.
They certainly fall far short of the female status in an
ant commune. One may safely think that a great deal
more of it would be to our advantage. The fact to be
especially noted is that among ants, as also among other
insects, nature has built up upon the female organiza-
tion, and not upon the male, the most remarkable and
successful examples of social life and government known
to natural science --the ant commune, the beehive, and
the hornet's nest.
In the internal view of an ant commune's affairs the
most striking facts are the relations of the queen mother.
Her queenhood is wholly fanciful, except in the first
stages of her independent career. Her motherhood is
the great fact of life to her and her fellows. It is as a
mother that she is the destined foundress of a new com-
munity. After her isolation or adoption into an es-
tablished commune, which follows the marriage swarm,
she begins to lay eggs which are developed into workers
in due time.
If she goes solitary, her larger size and generous
nurture have accumulated enough substance to supply
Fig. 69 A QUEEN ANT SURROUNDED BY
HER COURTIER GUARD
food to the initial colony with little or no outside forag-
ing, and this is imparted, after the manner of her kind,
by regurgitation. All the duties of nurture, nursing,
washing, keeping up and keeping clean the premises, are
wrought by her within her secluded and protected quar-
ters until a little band of helpers has been reared around
her. These at once begin to share labors with the
queen mother (Fig. 69).
When they have passed their callow period, they break
the original bounds and venture forth in search of food.
Day b}^ day the number of inhabitants increases; the
formicary is enlarged by cutting out and building up new
rooms and galleries; perhaps a new site may be chosen.
A wider range of foraging is compelled by the needs of
the growing community. The various labors, carried on
at first by the queen, and afterward by the few pio-
neers, become more and more specialized, until at
last are developed the vast and divided industries of a
large and fully organized ant commune.
Meanwhile a remarkable change has befallen the queen
mother. The workers, as their number grows, have taken
on more and more the responsibility of labor and adminis-
tration, until at last the whole burden thereof is lifted
from the queen, and she is limited to the function of
motherhood (Fig. 70). She lays the eggs from which
new citizens must be recruited, a service which increases
in importance with the expansion of the community.
Not only have the labors enlarged, but the wastage of
life has greatly increased through accidents by flood and
field, and perils of farther adventure into a world full of
strange creatures who prey upon them, as do birds and
divers beasts; and who war on them, as do sundry in-
sects and ants of alien tribes. Across their trails come
cattle and men, whose ponderous feet crush them un-
wittingly or carelessly.
Every day has its list of casualties, very large at times.
Every morning sees many who venture forth in quest
of food supplies for dependents and home-workers,
bounding with vigorous life and highly intent upon use-
ful service, who never come back. When evening comes,
at the gates of their loved citv no watchful sentinels
greet them. No eager nurses, or hungry antlings, or
comrades weary with toiling on the \vorks, shall lift up
lips for sweet refection, the garnering of the day's ad-
venture. Somewhere outside the city bounds, it may
be near by, it may be afar, there has been a tragedy
that no annals shall record and no ballad sing, but which
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robs community of a useful life, and cuts down a happy
worker in the midst of a wholesome career. "Only one
ant!" Yes, it is not much in the vast fecundity of
nature, and is easily replaced. But it is an atom in the
world's order that no human power can restore.
So, in the larger field of industry and research, our
race has its daily tragedies, not wholly unlike these
which befall the citizens of an ant-hill. The martyrs
of industry, heroes, and heroines; the fallen soldiers of
the great army of labor, are thus going forth daily to
perish in the path of duty. It is said that one out of
every eight adult persons dying yearly in Pennsylvania
dies a violent death. Not wholly, but too much, far
too much, like the indifference of ant communities is the
indifference of human society to these industrial trage-
dies. And the rational excuse of the ant is not for us,
for nature has not made us that way. With us it is
mainly wicked hardening of heart. How shall we cut
down, if we may not wholly cut out, the long list of such
wasted lives? Meanwhile, how shall we provide for the
maimed, and for the dependents of the slain? Are not
these soldiers of industry also worthy of communal care?
a Breed more workers !' is the answer of the ants.
The circle of their instinct has no wider swing. But at
least they will make sure that the supply is sufficient,
and the standard of wholesomeness and efficiency is kept
up. And so the queen mother must be encouraged to
her utmost productivity, and every egg dropped must
be preserved and reared with utmost care. 1 A peep
within the city walls will show a rare condition. The
whilom sole potentate who, first in solitary power, and
then in maternal sovereignty, and next in undisputed
matriarchate, held unchallenged authority and the ex-
clusive right to labor, is seen in a large vaulted chamber
in the heart of the galleried cone. She is not alone, but
is surrounded by a circle of workers. Is she a prisoner?
1 In my work, Nature's Craftsmen, chaps, i and ii, readers will
find a rather full and connected account of a queen ant's life.
-a sovereign deprived of queenhood, and in the hands of
regicides? Not so bad as that! The offices of the
guardians are at least friendly. They are a body-guard ;
in fact, the so-called "courtiers" of the ant queen a
phrase of courtesy, as is the word "queen," for the days
of sovereignty are over, and these keepers of the sacred
person are saying to her, in this decided way :
" God did anoint you with His odorous oil
To wrestle, not to reign!"
: \ ^^^^^M-^ f f^^^^m^^ : ^
Fig. 71 WORKERS TAKING EGGS FROM THE QUEEN MOTHER
And here you may see a fair example of the sort of
" wrestling ' expected of her. She has paused in her
march around the room. She raises her body well upon
the hind pair of legs. See ! From the oviduct beneath
her abdomen she forces a minute, white, ovoid object
which has no sooner dropped than one of the body-
guard rushes from the inner circle, seizes it in her jaws,
and hurries therewith from the chamber (Fig. 71). It
is an egg the norm of a future citizen! The process
described will be repeated over and over again, many
thousand times, until the ovaries are exhausted or death
It is to save these precious particles of living matter
for the community that this circle of watchers con-
tinually surrounds the queen mother's person. Theirs
is a tribute to motherhood, not to queenhood. Cer-
tainly they have not reasoned it out, but instinctively
they know that the prosperity, the very life, of the
commonwealth depends upon the maintenance of that
fecundity whose cessation would be "race suicide."
Popular fancy has brought to the explanation of this
"royal body-guard'' the familiar lines:
"There's such divinity doth hedge a king
That treason can but peep to what it would."
It will be seen, however, that this "hedge" about our
ant queen amounts simply to a case of communal
vigilance, represented by watchers set by the self-govern-
ing majesty of the commune to save all the ant eggs
possible. Doubtless there is "divinity" in it, as there is
in all honest discharge of duty and outworking of
nature's laws. But anything like regard to sovereign
state, or purpose to give or maintain royal honors, is
wholly foreign from the situation. Reverence for
motherhood is there, however wholesome and protected
motherhood, the essential fountain of communal virtue,
vigor, and perpetuity. Are we losing from our own race
the due reverence of that "divinity doth hedge about' 1
maternity? Woe to the nations or peoples, be they ants
or men, in such estate!
The body-guard of an ant queen is an elastic ring that
expands and contracts with her movements. If she
move around the room they move with her. If she seek
an adjoining apartment, the ring precedes, accompanies,
pursues, but never breaks up. Sometimes the guard
conceives that her maternal majesty needs special guid-
ance, a very courtier-like and cabinet-like conception.
Then one will see her bulky body gently solicited by a
pull upon her sensitive antenme made by a worker-
minim, or a tug at a leg by a worker-minor, or a push
or pinch upon the abdomen by a worker-major (Fig. 72).
A sort of volunteer steering committee are these; func-
Fig. 72 A TRUANT QUEEN BROUGHT HOME
tionaries apparently needed, or at least present, in all
organized governments, but coming as near to the van-
ishing point in ant cities as is conceivable.
Leaving the queen and her body-guard, let us follow
the fortune of the egg. From the queen mother it is
carried into a separate room, presided over by attendants
who have received the not inapt name of " nurses."
There is nothing to distinguish them as a separate class.
They are on duty at that point for reasons satisfactory
to themselves and to the secret but all-sovereign Spirit
of the Commune, whose mysterious sway all freely obey.
It does not appear that there has developed a special
class of workers with the charge of the communal young
as their chief function. Nor are such duties assigned
to the maimed, or to the toothless members, whose jaws
have been worn clown by age and by the gritty and
stubborn material upon which they must labor. The
nurses seem to be in the prime of anthood, vigorous and
It does appear, however, that the callow antlings, just
out of their cocoon cases, are found among the nursing
squads. They lose no time in taking up their life's work
as helpful citizens for which they enter imagohood full
panoplied but fall -to where opportunity first serves,
and take care of their larval fellows. Until their shelly
bodies become well indurated, they do not usually vent-
ure out-of-doors, but engage in tasks to which they are
physically better adapted. This is a part of such educa-
tion as they are to receive; for ants, like children, profit
more by examples than by precept and criticism. The
models of active public servitors are before them, and
they simply do what all around them are doing. But
the whole field of labor lies open to these prentices of
the State, with no restrictions thereon.
The eggs soon develop into minute larva 1 , fragile and
helpless things that need close and constant care to pre-
serve them in life. Owing to the social conditions of
their being, they do not have that sturdy hold on exist-
ence, and power to care for themselves that mark
solitary larva?, or such as those of moths, that are gre-
garious in their larval or caterpillar stage. Thus from
the beginning and throughout their growth and they
grow rapidly they must be fed and cared for. Their
care is always a first consideration. In the wreck of
an ant city the workers may be seen to grasp the eggs
and the young, and, careless of themselves, run to and
fro, seeking places of refuge for their helpless charges.
One will see them in little heaps, often graded accord-
ing to size, scattered throughout the nurseries. No
observer has yet conclusively noted such treatment as
prevails in beehives, where male and female eggs are
separated from ordinary workers, and a queen can be
developed from a worker larva by enlarged quarters
and specially enriched food. The larva? of all ant castes
and sexes seem to be kept in common and to receive like
attention. The nurses continually hover over them.
They lick them as a cat does her kittens. The larvae
learn to perk up their wee black heads and open their
mouths, into which the nurses place food and drink.
They shift their positions from side to side, sometimes
from room to room, sometimes with apparent good rea-
son often, one fancies, simply from the overflow and
outgo of such maternal sentiment as leads a young
mother to dandle and fondle her infant offspring, cooing
the while her love-phrases or love-songs; a spectacle
truly pleasing to the observer at least, and doubtless
often to the infant. Certainly herein the female temper-
ament shows its supremacy (Fig. 73).
Some readers who, like the author, have seen service
in active military campaigns, know that male soldiers
can be organized into a hospital corps for effective nurs-
ing of sick and wounded comrades. Many of us carry
remembrances of how bravely and well, with what devo-
tion and fidelity, this duty has often been done. Here
and there, too, men have developed special qualities
that have made them pre-eminent in the delicate and
But what veteran, who has had the opportunity to
observe, has not noticed the vast change that the en-
trance of trained female nurses has brought into the
field? There are deftness and sympathy and tender-
ness a something indescribable, but most potent,
which women bring with them ancl which men have
not -that work a transformation in scenes where human
dependents are to be ministered to. These qualities
are the fruitage of the female temperament. They
spring out of physical organism, which so largely in-
So, too, we all have known men whose love of children
Fig. 73 AN EMMET NURSERY FOR THE YOUNG
has been strong as death, and has made them good and
careful nurses in an emergency, and for a limited period.
But that man is indeed a rarity who is a proficient in the
care of immature infants. Men may dandle young
babes with delight and even success for a while. But
the tact, patience, enduring fondness, and instinctive
knowledge of the real natural nurse of infants are en-
dowments of the female temperament alone.
It is certainly so among ants. The males are simply
nonentities in the care of the commune's dependents.
They are themselves dependents of the most absolute
sort. Nature has denied them the gifts requisite for
effective service. To one who knows them well, and
their temperament and ways, it would never occur to
think of them as caretakers for the nurslings of the
commune. And this judgment is not affected by the
occasional and very rare instances in which male ants
have been seen to make some slight and awkward ap-
proaches toward a seeming part in the ordinary worker's
For the most part, nurse ants take up and go through
their duties in a business-like spirit and way. It is done
thoroughly, and does not cease until the larvae have
spun up around them their silken pupa-cases. Nor
then; for these cocoons are constantly watched, cleansed
and cared for, and when the time comes for the young
imago to escape, it is aided by the scissors-like jaws of the
nurses, whose obstetrical services are aided by the efforts
of the outcoming nymph.
Did Lycurgus get from the ants among his Spartan
hills a first suggestion of his theory that children are a
communal possession, to be reared at the charge and
with the oversight of the State from the earliest age
practicable? Certainly, our American Republic is well
impregnated with the germ of that theory. Its essential
spirit largely controls the subject of education. True,
we have not yet reached the high stage of ant govern-
ment, in which the whole aim and activities of the com-
monwealth pivot upon and move around the rearing
and care of the young. But, at least, it is a ruling theory
of our people that organized society owes every child a
common-school education. At a tender age our children
are separated from their homes for a part of the day,
and placed by the State by legal compulsion, if need be
-where they get training and instruction without re-
gard of social distinctions.
Herein is the common meeting-ground of all classes at
the most impressionable period of life, and the main-
tenance of the true democracy of our republic depends
largely upon that fact. We carry the principle so far
that we not only provide school-houses, teachers, school
apparatus, fuel, light, and janitor service, but we supply
text-books for the scholars. In many sections their
car-fare to and from school is paid; or, as in a district
school hard by the writer's country home, a big omnibus
goes the rounds o' mornings and gathers up the pupils,
and again at evening calls for and distributes them to
We dare not have it otherwise. Government must
continue to be responsible for the education of its young
citizens. For, however willing, individual families are
not able to do this unaided by the State. Even the
wealthy must submit to something like the same law.
Said a multi-millionaire to the writer: "We shall go to
Southern Italy to spend the winter. We might as well
close our house. To-morrow our son goes to (naming a
college for men), and our daughter returns from Europe
to go to (naming a college for women). Our only other
child is married. Our home will be empty. We will
go abroad." Thus the wealthy parent is not exempt
from the necessity of committing his children to society
to educate and train for future citizenship.
So it is in all higher education classical, scientific,
mechanical, professional, military, and naval. Or-
ganized society becomes, and must become, a nursing
mother to the youth from whom, for the most part, her
future rulers and most useful servants must come. As
for the young waifs of society the flotsam and jetsam
of child-life, continually tossed amid the wreckage of the
world's great social sea long since government has seen,
and sees it more and more, that they are in an especial
sense the children of the State, and must be adopted
and trained into citizenship by the State. Thus far, at
least, our commonwealths are swayed by theories and
have taken up practices long ago prevalent in ant
Unhappily, our system breaks down where that of the
ants proves splendidly effective: by our absence of
system in providing work for young citizens as soon as
their working powers are mature. In the ant commune
every individual passes at once from pupahood to the
status of a laborer. In a human community the
citizen's work, in both fact and form, is left chiefly at
haphazard. It must, indeed, be that with us, as with
hymenopters, the Spirit of the Commune has some subtle
potency in directing unconscious youth to the choice of
occupations and keeping the working mass in activity.
But the State as a State eschews the matter, and there
is no sense of communal responsibility that every citizen
throughout life should be employed steadily and usefully.
Might it not be that if there were more of the female
element in the governing of our communes, there would
be far less of that waste of steadfast and regulated in-
dustry, through lack of early discipline of citizens, which
is such a fertile source of loss of character and profitable
THE PROBLEM OF COMMUNAL DEPENDENTS
WHEN one considers the incalculable hosts of ants
that inhabit all parts of the earth, and that every
individual thereof has been reared, from egg to imago,
by the direct personal care and toil of adult members of
ant communes, he may have some conception of the
immensity of the labors involved therein.
It would, perhaps, be pushing metaphors to an un-
warranted extreme to speak of "dignity of labor" in
connection with the occupations of ants. But if by
the phrase we mean that labor is the honorable lot of all
citizens, and that all labors of whatever sort are upon
the same level of respectability, then we might venture
to apply the saying even to the labors of an ant-hill.
For therein all are workers from the newly fledged
callow to the veteran of a second summer.
Therein is no taboo upon "hand toil." All forms
thereof are equally creditable. We are reminded of the
simpler state of society in the pioneer days of the
United States and Canada and the British colonies.
Indeed, it is the natural social order of human com-
munities, until great possessions, earned and inherited,
or usurped, or fortuitously acquired through communal
increment, create a favored class. Surely this is an ideal
republic no idlers, no tramps, no citizen-parasites, no
misers, no spendthrifts, no paupers !
This inviolable law of the emmet republic needs to be
restated when we come now to consider what seems to
be an exception thereto. We have seen that the popu-
lation of ant communities is largely composed of the
larvae and pupre, the helpless younglings from whom the
future citizens must corne, and whose nurture is the
chief aim of the active commonwealth.
These immature dependents are so numerous that one
would think that they alone might tax the resources of
any society. Nor is it simply a problem of crude labor,
quantitative energies, herein involved. As an outside
intelligence views the situation, there is a large field
for the exercise of qualitative energies, also, in the rearing
of these youngling ants.
We have already seen how the squad of so-called
"courtiers," in a circle of ceaseless vigilance around
the fecund queen, manages to secure the eggs and trans-
fer them to the charge of the nursing detail. It is mani-
fest that the process by which these minute specks of
vitality, that carry within them the future of the com-
mune, are tended cleaned, fed, shielded from changes
of weather and all hostile influences must involve a
good deal of delicate and discriminating care.
The eggs soon become Iarv83, small, soft, and ex-
tremely fragile objects, which need dainty handling to
nurse into vigorous life. They grow rapidly, and one
must suppose that the portioning of food to the chang-
ing grades of age and strength requires such qualities
as we are wont to ascribe to a considerate mind. Again,
as the larva? pass into the pupa stage they demand a
different character of treatment, which must call into
play faculties, or at least activities, that with men would
imply reflection and wise selection and decision.
PROBLEM OF COMMUNAL DEPENDENTS
Besides the fertile queen, or queens, and the host of
brooding larva', our ant community is taxed with the
support of the winged virgin queens and males (Fig. 74).
For ants are not apterous insects, unless we take the
worker as the original type of the order. The parents of
Jg 74 - YOUNG WINGED QUEEN OF HONEY-ANT (SIDE VIEW)
nearly all known species have, and from a remote period
have had, wings. These have been lost to the maternal
stocks through the exigencies of an underground or
interarboreal habitat; and the winged forms have been
preserved in females and males to favor that flight and
commerce in the air by which species have been pre-
served and distributed. The swarming of winged ants
on a soft September day is a sight not easily forgotten
by a new 7 observer, and which is not apt to lose its in-
terest to the adept, As often as the writer has seen it,
he still feels the thrill of excitement that pervades the
commune, as he sees the hosts of winged creatures pour
out of the formicary gates.
Here, beneath a young apple-tree, is a nest of Lasius
flavus, whose existence had not been suspected until,
in passing it, the free soil around the trunk was seen to
be alive with a seething mass of yellow ants males,
females, and workers intermingled. They ascend the
tree, whose surface is fairly covered with them. The
gauzy wings of the sexed forms glisten in the sunlight
as they march along. The workers hurry back and
forth among the hordes upon the ground. Some join
the column upon the tree trunk. They seem to en-
courage their winged proteges to take flight, even nip-
ping them at times with their jaws to hasten depart-
ure. They are in a fever of excitement.
And well they may be; for this is the grand event to
which a good half of the summer's work has steadily led.
Thenceforth the commune shall be free from the immense
burden of supporting this army of non-workers. How
many of them there are! Numbers are continually
taking flight. Away into the brilliant sunshine they
soar, until they are lost to sight, the females alone and
not accompanied by mates, as I have observed in other
cases. Their fecundation had been accomplished with-
in the nest. Others still are streaming out of the city
gates to join their winged comrades on the tree.
Like scenes are enacted at two other points the
farthest twelve feet distant, under a young pear-tree: .
the nearer in a shaven orchard sod. midwav between the
two. These three centres of agitation seem to be parts
of a common movement of one great community, whose
subterranean quarters intercommunicate across the
intervening space. Ere nightfall the crowds of winged
forms have disappeared and the city gates are solitary.
And this throng of creatures, many hundreds of them,
PROBLEM OF COMMUNAL DEPENDENTS
had been wholly dependent for food and care upon the
workers of the colony during the entire summer.
Again, on a warm day late in June or early in July,
one may see the air, at a short distance above the ground
and for many square yards around, filled so thickly with
flying insects that they seem like a thin cloud of quiver-
ing mist. They are the sexed forms of a small species
of Lasius, whose inconspicuous nests are spread numer-
ously over the lawn and field.
Many of these make their exit and marriage-flight at
the same time (Fig. 75). They rise and fall, and weave
in and out through the quivering air in their mating evo-
lutions, sporting in the sunlight. They fill one with won-
der that such a feeble folk as rule the weak communities
whence they issue could bear the burden of nurturing
into maturity such swarms of dependents.
But considerable as are these outputs of non- workers,
they do not strike the imagination so forcibly as some
of the well-authenticated accounts of immense marriage-
flights of ants that have been published. 1 It seems
incredible that the whole surface of a lake of two
lakes, in fact should be covered so thickly with these
winged creatures that they could be pushed up by
passing boats into windrows several inches high and
extending from shore to shore on all sides, as in the
observations of Mr. W. C. Prime on Lake Lonesome.
It is interesting to note that this is not a novel oc-
currence. Such disasters have marked the history of
flying ants from the earliest ages. Professor Wheeler
spent the summer of 1906 collecting in the Florissant
fields of Colorado, noted for their rich yields of fossil
1 For details, see author's Nature's Craftsmen, chap, ii, Harper
& Brothers, New York.
insects and spiders. This ancient Florissant lake-basin
lies among a series of low- wooded hills and ravines. At
the period of the Oligocene division of the Tertiary geo-
logical era this elevated lake must have been a beautiful
sheet of fresh water. It
was hemmed in on all
sides by granitic hills,
whose wooded slopes came
to the water's edge in
this phase not unlike
Lake Lonesome, among
the White Mountains.
Professor Wheeler's col-
lections, which I have
been permitted to exam-
ine, show the fossil ants
to be more abundant than
any other insects. But
only males and females
Fig. 75 A MARRIAGE-FLIGHT, OR " SWARM," OF WINGED MALE
AND FEMALE ANTS
are represented, indicating that these had been sub-
merged in the lake during marriage-flight, precisely like
those reported by Mr. Prime.
PROBLEM OF COMMUNAL DEPENDENTS
Thus the vast interval between the present and the
Tertiary eras is bridged by a continuity of habit which
joins in substantial unity of social behavior the ants of
to-day with those of far geological antiquity. In
harmony with this is the statement that all of the eight
hundred specimens secured belong to extinct and un-
described species, and are wonderfully like existing
forms. It is substantially the same story that one reads
in the even better preserved ant forms of the fossil
amber of Europe. The Formica fusca of the Baltic
amber, for example, appears to be entirely identical
with that of the present. Much the same general con-
clusion arises from a study of the fossil spiders. 1
In every such case as the fossilizing of the Florissant
ants and the swarms of Lonesome Lake, the innumer-
able hosts of insects massed within a comparatively
limited field must have come from a great number of
nests dispersed throughout the general locality. We
may conclude that the cycle of maturity was completed
simultaneously in all these communities, and that
similar favorable conditions united to induce con-
temporaneous flight. The intermingling of the various
individual swarms, as they were borne along by the wind,
sufficiently accounts for the extraordinary massing of
winged creatures which were swept over and into the
White Mountain lakes. This will not wholly explain the
phenomena; for the virgin queens and their male
partners, in full maturity, have often been seen to be
inhabitants of the commune for a considerable period
before marriage-flight. Evidently they are prepared
for the exit long before it comes, and await therefor
1 See author's American Spiders and Their Spinning-Work, vol. ii,
chap, xv, " Ancestral Spiders and Their Habits."
some signal from nature, some potent impulse or con-
Setting aside, then, the completed cycle of maturity
as the sole cause of this remarkable assembly, it is in-
teresting and not improbable to suppose that a wave of
sympathetic excitement issuing from a few nests may
have infected all the surrounding section until, by a
common impulse, the entire emmet population of the
mountain-side was astir with the fever of flight. We
know how, in human societies, neighboring families,
towns, and cities are apt to be seized almost simultane-
ously with a political or patriotic or religious fervor, or
revival, that spreads with a swiftness and complete-
ness that are so remarkable as to appear to many quite
beyond known causes. With equal celerity and uni-
versality, and equal mystery of psychological cause,
will panics spread among armies and communities of
Psychic contagions are not confined to men. The
animal world, in some of its races, at least, is subject
thereto ; and to these, in some measure, we may attribute
the impulse that seizes at once the myriads of winged
ants, and sets them forth together. This impulse must
be felt by the workers also, the rulers of the communes,
if indeed it does not originate with them: for their de-
pendents are not always willing exiles from the favorel
precincts of the home nest. I have seen them turning
back at first with manifest reluctance, and seeking to
enter the city gates against the ungentle persuasion of
the workers' sharp mandibles. It requires such dis-
cipline and the allied mighty force of a natural instinct
to banish them from their sheltered life of ease in their
happy native homes.
PROBLEM OF COMMUNAL DEPENDENTS
It is appalling to think that upon the industrious work-
ers devolves the task of providing food (Fig. 76), home
quarters, and protection for the many millions of robust
creatures that were overwhelmed in the waters of Lone-
Fig. 76 ARRIVAL OF THE FOOD-BEARERS AMONG THE WINGED
some Lake, together with the multitudes that must have
escaped. And all this in addition to the nurture and
care of an equal or even greater number of immature
citizens in the form of eggs, larva?, and pupa?! That
this is done, and done effectively, is a marvel of industry
and devotion probably unsurpassed in the records of
Why is this service undertaken? What is the im-
pelling force to such labors and sacrifice? The answer
is not far to find. It is the inborn and ingrained instinct
to preserve the species and the commune. For that
ants live, and for that they die. Their life is ideally
altruistic. Nature has so deeply fixed upon their or-
ganism the love of their own community and their own
kind that there seems to be no room for mere selfish
pleasure of any sort. The necessity to maintain by
their labors the host of males and virgin queens raises
no opposition, and apparently excites no ill will. It is a
communal necessity. It is exacted by nature. That
is enough for an ant citizen.
I have never noticed in the working castes the faintest
ripple of anger or rude treatment toward these adult
dependents, suggestive of envy or of impatience under
their heavy burdens, and reacting in violence. Their
attitude is invariably helpful when help is needed, and
tolerant and good-tempered at all times. Not until the
crisis moment of the commune has come, when the great
exodus of the sexes is to begin, is there any show of wish
to be rid of their charges. And that is controlled by
the same imperative spirit of altruism toward the future
of the race, and has in it no trace of personal cruelty or
Doubtless, in their brief and strenuous life, the pleas-
ures of appetite have some place, although indulged with
exemplary moderation. Theirs, too, must be the satis-
faction of all normal healthy organisms in natural work
and in the achievement of daily rounds of service. What
PROBLEM OF COMMUNAL DEPENDENTS
may be the depth or quality of such feelings in ants we
may not know, but surely kindly nature has not denied
some just measure thereof to these faithful and laborious
creatures. But, as far as the observer can note, these
are small factors in determining emmet behavior. And,
withal, work is work, in an ant commune as elsewhere.
Its burdens are often severe, its risks are great, and the
number of workers daily maimed and slain in the course
of duty is a heavy drain upon the vital resources of all
Yet, how diligently their task is wrought, how cheer-
fully, how patiently, how bravely, how well ! Silent citi-
zens of the ant city! With all his God-like endowments,
man may well consider your ways herein as worthy
models for his own relations to the commonwealth and
the common weal. It may be true that all this admi-
rable conduct is wrought without moral consciousness
and freewill, such as mark " articulate speaking men,"
undesignedly, instinctively, automatically, if you please.
But .there it is. And it is there by that Over-thought
and Over-force who has appointed destiny and basal
character for communes of ants as well as for cities of
men. And by this bond and fellowship we may find
a common ground for our admiration and for imitation.
We have seen that the first eggs laid by the ant queen
are embryo workers. This follows necessarily from the
fact that new communities arise from single fertile
females. The existence and growth of the society re-
quire that its first members should be helpers and not
dependents. Only when the pioneer colony is strong
enough in workers to add to the needful conditions of
ordinary life and growth the burden of supporting the
males and females do these sexed forms appear.
The author has not noted in newly dropped eggs any
marks indicative of differences between sex-eggs and
caste-eggs, nor does he know of observations by other
connoisseurs to that effect. If such exist they are of a
subtle character and escape ordinary observation. But
as the eggs develop into larvre and begin to grow, they
are easily separated into groups by their sizes, according
to the nature of castes, in any specific nest. So, also,
when the larvae have spun themselves into their
cocoons, the workers and the females issue from the large
cocoons, and both appear with their own distinctive
characters. There appears to be no seclusion of workers
for special feeding and care in order to produce queens,
as with bees. The larvae lie in common heaps, and share,
as far as can be noted, precisely the same amount of
feeding and attention. The worker castes, as well as
the males and females, show at once after emergence
from the pupal stage their distinctive characteristics, not
only in size, but in such a striking peculiarity as the
unusual development of the head of the soldier caste in
genera like Pheidole and Atta, although this is liable to
When the imago life is achieved, the radical difference
between the sexes and the worker forms soon appears.
The workers excel in complex instincts, and as they turn
to their various duties as nurses, builders, miners, for-
agers, sentinels, warriors, sanitarians, etc., they display
a plasticity of temperament that suggests the possession
of marked qualities. These are much less apparent in
the virgin queens, where, indeed, they scarcely appear.
But after fecundation, deflation, and entrance upon
nest-founding there is a rapid development of latent
qualities into action which their important role requires.
PROBLEM OF COMMUNAL DEPENDENTS
On the other hand, the males are phenomenally stupid.
They are unable to distinguish friends from foes, or to
find their way back home when they wander from their
nests. The points in which they are richly endowed are
the eyes and antennae,
the two sense organs
which are connected
with the brain, and
give that keenness of
sight and smell re-
quired for their espe-
cial function in life-
to possess themselves
of the female during
or before their nuptial
With these etholog-
ical facts closely cor-
responds the structure
of the brain in the
three forms that con-
stitute an ant commu-
nity. This has been
admirably shown by
Doctor Forel in his fig-
ures of the brains of
the worker, queen and
Fig 7 7_ TH E BRAIXS OF ANTS
Worker. F Female. M Male.
male of LasillS fldigi-
nosus (Fig. 77). The
brain is relatively large
in the worker, the cortical portion extremely rich in
cellular elements. It is much smaller in the female,
and is almost vestigial in the male, although in the
latter the optic and olfactory lobes are large. [F. 5,
Life within the precincts of ant communes is largely
hidden from the outside world. However, one mav get
fairly truthful glimpses thereof from studies of formi-
caries arranged in glass vessels. Many such, which were
artificial only in their limited spheres and furnished
food, for they were built up by the inmates wholly upon
their own lines, have yielded the author numerous facts
and hints from which he has pictured images of interior
life that cannot be far from correct. Observations of
actions on and around the nest exteriors, and analysis
of the mounds themselves, have added to the accuracy
of such inferences.
But much remains unknown, and we are left largely
to conjecture in representing the life of the winged males
and females that fill up the cavernous rooms and crowd
the galleries of the Alleghany mound-makers and similar
emmet architects. We can fancy the industrious work-
ers passing from one to another among these throngs
of winged dependents, feeding them from the liquid
sweets stored within their crops during foraging trips.
How eagerly are welcomed arrivals from the outer world,
of these voyagers! And how zealously the incomers
hasten to their task! A bevy of boarding-school boys
could not give heartier greeting in their living-quarters
to the latest arrival from home, laden with spoils of
storeroom and kitchen, than the}* receive.
We see the crowding and the general stir as the food-
bearers come round : the flutter of wings, the haste and
hustling of greedy ones after undue portions, since even
an ant-hill is not exempt from such traits, especially
(one might almost say, exclusively) among the idlers.
ROBLEM OF COMMUNAL DEPENDENTS
We note the agitation that follows in the trail of the
ministering ants as they push their way from point to
point, until their exhausted supply warns them to retire
from the scene.
What other pleasures than those of appetite are open
to these winged dependents? The pleasure of work is
denied them by nature. The natural history of social
insects gives no examples of more absolute idlers than
they. Does time hang heavy as they plunge through
the galleries, jostled by the miners and builders, who pay
little heed to them as thev run to and fro with their
burdens? In the domed chambers wherein they con-
gregate, and the swelling bays that relieve the strain of
traffic upon the galleries and gangways, they huddle and
preen their coats and sleep, and in some species, perhaps,
pay and receive sexual court. What other activities
engage their attention in this listless life, in the midst
of their strenuous supporters, it were vain further to
conjecture. Future observers may have something more
Such a subterranean career is, from our standpoint,
passed in darkness. But we are not to conclude that
the same or even an analogous condition exists for our
emmet cave-dwellers. There may reach them vibratory
remnants of light-rays, in measure and quality quite
beyond human appreciation, but which suffice for ants.
Moreover, those remarkable olfactory organs, the an-
tenna 1 , are so extended and flexible, so sensitive and so
capable of conveying a knowledge of environing condi-
tions and relations, that they may easily supplement or
even supply the seeming deficiency of light.
Be that as it mav, the writer, after the most careful
attention of which he is capable, has never been able to
note, in any species, the slightest shock or shrinking
when ants issue from their formicary gates into the sun-
light, such as one would expect in beings organized after
our human fashion. Of course, the passage through
the vestibule of the gateway, where it exists, affords
an opportunity (were such needed) to adapt the eye to
such an extreme change. But in our mountain mound-
builders, and other species of like habit, there is no
measurable vestibule. Besides, the movements of the
ants are so rapid that their plunge out of darkness into
full light seems to be instantaneous. As all the varied
labors of the workers are carried on within the sub-
terranean passages and rooms without the least em-
barrassment, those places cannot be so cheerless to the
winged idlers as one might fancy. Perhaps the monot-
ony of their inactive career, in such sharp contrast
with that of their protectors, may be the chief factor
in their discomfort, if any such there be.
Breaks in the monotony of this underground life come
to the virgin queens and males in occasional visits to
the outer air. These were especially observed during
studies of the agricultural ants of Texas, in the neigh-
borhood of Austin. Such excursions were frequent, and
were evidently made simply to enjoy a bit of sport in
the sunshine (Fig. 78). Both sexes were seen bobbing
in and out of the gates, peeping forth and quickly with-
drawing, and again venturing one or two feet distant
from the entrance upon the smooth disk that surrounds
it. However, they rarely went far beyond the gate, and
were quick to retire within at any sound or suspicion of
danger. [McC. 3, p. 141.]
One female reached a grass-stalk near the pavement's
edge, and amused herself by swinging upon the blade.
PROBLEM OF COMMUNAL DEPENDENTS
On the broad plaza of one city half a dozen or more
young queens were out at the same time. Their play
took the form of running up a large pebble near the gate,
facing the wind, rising to a rampant posture, and so
down again. Several having ascended the stone at one
time, there ensued a playful passage-at-arms for position.
Fig. 78 WINGED FEMALE ANTS AT PLAY ON THE PLAZA
They nipped one another gently with their mandibles,
and chased one another from favorite spots. Their
whole demeanor was that of a party of romping youth
playing "tag" or "hold the fort'' upon a big rock.
[McC. 7, pp. 4, 22.]
While the young queens lightly nipped one another in
their game, as dogs at play will do, it was noticed that
they never took such liberty with the workers. The
latter evidently kept close watch upon the sporting
princesses. They occasionally saluted them with their
antenme in the usual way, or touched them at the
abdomen, but did not interfere with the sport. Their
attitude reminded one of that of an under-teacher, or
usher, charged with the duty of conducting, or oversee-
ing a bevy of seminary girls in their daily exercise in
the open air.
In order to test the strictness of this watch, one of
the group was thrown, by a quick motion of the hand,
from the vicinage of the gate to the verge of the plaza.
She was instantly surrounded by several workers, who
began a determined effort to control her action, trying
to compel her to return toward the gate. The queenling
was confused or stubborn, and opposed her strength
quite vigorously to the purpose of the guard. For some
time the party floundered among the stumps of grass-
stalks in the little clearing on the margin of the plaza,
the bulky form of the one stubbornly set against the
quiet persistence of the others. It was noticeable that
the guards carefully abstained from anything like hurt-
ful violence to their charge, and that she did not attempt
to escape by flight. The issue of this trial of will-power
was not determined, for the refractory queenling was
needed as a specimen.
It is perhaps worth noting that the worker castes were
never seen at play. If records have been made by other
observers of such light behavior on their part, the author
has not noted them. The truth seems to be that their
life is so strenuous from its first experiences of imago-
hood to the end of their career, that there is no time for
PROBLEM OF COMMUNAL DEPENDENTS
recreations of any sort. Work ! work ! ceaseless work on
their endless round of duty is their lot, varied only
by scant periods for eating, for sleep, for personal cleans-
ing, and occasional mutual ''shampooing." The amuse-
ments of ant communities, such as they are, are limited
to the dependent leisure classes. However, it must be
remembered that all of the routine labor is not of the
exacting sort, like mining and nursing. Moreover, as
we have seen, the liberty to " knock off work" at will is
one of the inalienable privileges of the workers one that
is freely used, but apparently never abused. No doubt,
under such a rule, they get more satisfaction one might
even say more enjoyment out of life than winged idlers
whose career is shut in and restricted at so many points
that they seem to be little more than privileged prisoners
WARRIOR ANTS, AND THEIR EQUIPMENT FOR WAR
"VVJT AR, it is said, is a brutal way of settling differences
T T among men. That is true; and therein lies the
fact which gives most serious pause to one who would
study the subject philosophically, with an outlook upon
nature at large. War is brutal a natural habit of
brutes, and of the whole realm of organized life below
them, that wage war upon one another instinctively.
Their natural life is one of endless conflict. They who
justify war do so on the ground of its universal preva-
lence among creatures in a state of nature. It is brutal
but natural, arid man, being of nature, has his physical
kinships with brutes and their lower allies.
Doubtless those who base their opposition to war on
the divine precepts of the Prince of Peace have here no
difficulty. They admit the premise, but claim that
Jesus Christ, whose laws they obey, came to abrogate
the evil in the old, and to establish a new and spiritual
kingdom in Nature. He brought into human discipline
a new development, a higher stage of life, wherein war is
a discordant element. This is the new Nature, the spirit-
ual kingdom. It is the dawn of an ever-deepening Day
after a birth-Night wherein wild things ruled, but Life
and Light were born. The spiritual man, not the natural,
is now supreme, and under Christ the nations are to
learn war no more.
WARRIOR ANTS, AND THEIR EQUIPMENT
The writer accepts this view. He has had personal
experience in two wars the American Civil War and the
Spanish-American, in Cuba. He knows well its worst
features and its best. He believes that universal peace
and fraternity ought to be the ultimate aim of our race,
and that armies and navies are justified simply as na-
tional police forces for the administration of those be-
nevolent functions for which governments should exist
among men. Nevertheless, he recognizes that to many
minds the force of the facts, as seen in nature, is not
readily put aside; and that the universal war habit of or-
ganized beings, as it appears to have existed in all time,
seems to place upon a higher plane, as in harmony with
natural laws, those war-like habits and acts that have
dominated human history. This, at least, gives an ex-
ceptional interest to a study, for the sake of comparison,
of the war methods of those lower orders of living beings
whose social organizations strongly suggest our own.
Among the foremost of these are ants, and ants, as an
order, are war-like insects. The foragers carry their
natural pugnacity into the field as isolated individuals,
and show decided courage in the quest of food. Therein
they are freebooters. Whatever falls in their way and
they are able to possess, they take. This, as in the case
of human brigands, often requires an appeal to force.
An ant commune is as fair a scene of peaceful industry
as a beehive; but everywhere in its vicinage "doth
dogged war bristle his angry crest, and snarleth in the
gentle eyes of peace."
This readiness for hostilities and ferocity in attack
have been noted and recorded often of the hosts of
true ants that swarm along the pathways of travellers
in the tropics. For example, Stanley speaks of the
'belligerent warriors" among the innumerable species
of various colors that filled the African forests; of the
"hot- water ants/' as his men not inaptly named them,
from the smarting pain of their stings; and of the minute
red ants that everywhere covered the forest leaves and
attacked his pioneers so viciously that their backs were
soon blistered. These creatures doubtless acted from
a principle of self-defence that led them to hurl their
fighting myriads upon everything that crossed their
way and disturbed their solitudes, though with no
hostile intent. It was an act of natural bclligerencv, and
no doubt was protective, in the aggregate, of life. It
certainly seemed as little reasonable as were the un-
provoked attacks of the human hordes of cannibal
savages that assailed his expedition in their crowded
boats, as he made his way through the heart of the Dark
Continent, along the mighty Livingstone River. The
tribes of ants and the tribes of men were not unlike in
the native cornbativeness that animated them. [St.
vol. ii, pp. 138, 225.]
The woods within whose open spaces the mound-
making ants rear their conical cities are also hospitable
to the carpenter ants (Camponotus pennsylvanicus) ,
and the two species are natural enemies. Wherever
they chance to meet a combat is inevitable, in which
numbers sometimes become involved, and always death
and wounds succeed. Should one of these errant
Camponoti, from a near-by nest in a white-oak tree,
chance to cross a mound-builder's bounds, its tread,
light as it is, affects the commune like a signal-shot or a
fire -alarm. From the nearest gates issue squads of
sentinels, who fling themselves in mass upon the intruder.
Flight is thus hindered, even if it were considered, and,
WARRIOR ANTS, AND THEIR EQUIPMENT
despite the overwhelming odds, Camponotus joins battle,
and only succumbs, and is dragged within the walls, after
a number of its assailants have been maimed or slain.
The agitation in such a case is limited to a narrow
sphere, for somehow the commune knows that the dan-
ger is merely local. Therefore, outside of that circle,
the various duties of the government go quietly on.
But it is a notable feature of this commune that upon
a general alarm the whole citizenship rises up to meet
the threatening peril. Many times in many ways has
the author tested this. A few pats of the foot or strokes
of a stick upon the surface would call out a host of
sentinels and workers. The interior construction of the
mound is well adapted to communicate sound or vibra-
tory movements rapidly. Through the conical mass of
intercommunicating galleries and rooms the agitation
at the surface appeared to be quickly carried to all parts
of the mound.
At all events, it reached enough to call out, almost in-
stantaneously, a multitude of insects. With antennae
erect and quivering, with abdomens well raised from the
ground, with legs a jerk and heads aloft, they circled
about and rushed to and fro, their whole mien showing
keen excitement. With them, assuredly, "the toil of
war" is "a pain that only seems to seek out danger."
It is not a question of who has made the attack, or why
made, or whether one or another should come to the
rescue. At once the republic is ready to launch forth
its entire force, if need be, against real or imaginary
foes. This perfect unison in resisting the assault of an
enemy is surely an element of civic strength and per-
manence. During my boyhood a saying of one of our
naval heroes was widely current, and was a theme for
discussion in some of our Ohio debating societies:
"My country: may she always be right; but, right or
wrong, my country!" Xo budding ant citizen would
need to debate that question. The commune with ants
always has absolute priority with all its citizens.
Their supreme law is its demands, for life or for death.
History and, indeed, our own observation have shown
among men examples of somewhat smiliar communal
unison under the impulse of great social movements.
A wave of patriotic feeling will sweep over city or State
or nation, and carry it swiftly along until the purpose
or sentiment or emotion that inspired the movement
shall be spent in achievement or hopeless failure. Such
movements are more unanimous, and so more harmoni-
ous, in ant than in human communes. There is absolute
good temper and unanimity of feeling among the myriads
of inhabitants of our emmet mound city in all move-
ments noted, whether peaceful or warlike. Of course,
one does not expect such complete fraternity among
men, even in far less widely extended citizenships.
Whether in this the bipeds or the sexipeds are better
off and nearer to nature, let the reader query. If one
\vere to indulge such a fancy as that human civics have
developed from such lower and simpler forms as ants
exhibit, it would seem that in the evolution they have
been carried a long way (in some respects) from the
Xo trait in emmet character is more interesting than
this entire devotion of every individual, even unto death,
to the welfare of the community. The uprising of a
threatened ant city is a remarkable exhibition. The
peaceful commune is instantly transformed into an armed
camp. There is not the slightest delay or hesitation in
WARRIOR ANTS, AND THEIR EQUIPMENT
the response. \Yith utter abandon the little creatures
hurl themselves upon their assailants. No question
seems to arise. Shall we abstain? Shall we retreat?
" Or shall we on the helmet of our foes
Tell our devotion with revengeful arms?"
No condition of size or character in the adversary has
the least influence upon their action. There is no trace
of personal fear, no regard for life, no balancing of prob-
abilities as to victorv or defeat, but with the most
formidable as with the feeblest enemy the ants join
eager issue. There is no f melangering." None hangs
back waiting for others to take the brunt of battle.
In our mound-making ants, cowardice is an unknown
vice. I do not recall a clear case of poltroonery. They
are as valiant as they are industrious. In many cases
the destruction of the defenders is foregone, and the
foremost in the column are certain to perish. That may
not be understood by them; but were it so, it would not
make any difference with these citizen warriors, with
whom labor, health, unlimited service life itself are
held as the unreserved heritage of the commune.
There have been times in the history of human com-
monwealths when a large portion of the citizenships
reached as high a standard of patriotism. At all times
there are some who, in the surrender of their substance,
their service, themselves, and yet higher sacrifice-
their sons to the nation, show like devotion. But history
would surely falter if challenged to find among men a
case of unanimity in devotion to the commune in time
of danger equal to that of the mound-making ants of
A good example of the pugnacity and courage of ants
is a small species (Dorymyrmex flarm). that digs its
little nests upon the great open spaces surrounding the
central mound of the Occident ant of Colorado. A
large commune of the latter which had been badly
damaged by the wash of heavy rains was a scene of
active rebuilding. Four moundlets of Dorvmvrmex
had been reared upon the pavement, one of them quite
near a centre of operations in one of the main tracks by
which the workers had ingress and egress. Here an
incessant warfare was being waged by the dwarfs upon
their big neighbors. Every Occident that essayed the
passage to or from the ground was attacked. Squads
of Dorymyrmex surrounded their single gate, and on
the approach of one of the Occidents the nearest war-
rior flung herself upon the unconscious intruder. That
she was alone, that there
was such disparity in size
between her and her adver-
sary were facts that plainly
had no part in her calcula-
It was curious to note
the effect upon Occident/alls.
She stopped instantly ; drew
her feet closer together;
stiffened the legs, thus rais-
ing her body well above the
earth ; bowed her back ; ele-
vated her head; stretched
out the sensitive antennae,
as though to guard them
aspecially from harm; opened the mandibles; and, in fact,
presented an amusing likeness to the pose of a cat at the
Fig. 79 WAR-LIKE DWARFS
ATTACKING AN OCCIDENT ANT
WARRIOR ANTS, AND THEIR EQUIPMENT
first onset of a dog. The fore leg upon which Dorymyrmex
had seized, and which had instantly been raised, was then
shaken violently, and the little assailant rolled upon the
ground (Fig. 80).
Thereupon Occident unbent herself and resumed her
way. She scarcely had started ere her tormentor
Fig. 80 A PLUCKY LILIPUTIAN ATTACKING AN OCCIDENT ANT
again was upon her, followed by another and another,
until her body was dotted with the little vixens. They
grasped her feet, fastened upon the under parts of the
abdomen, mounted her back, seized her antenna. They
could not be shaken off. She snapped at them with her
strong jaws; struck at them with her claws; doubled her
abdomen under her body, and thrust at them her barbed
sting. Some were crushed, some were thrown off, but
others came to the assault. Anon the warring mass
rolled upon the ground, a whirling ball of red and dark
yellow, of quivering legs and antenna?. At last the
aggressors were driven off, or released their hold, and
Occident retired to a safe distance, combed her ruffled
hair, and passed by on the other side (Fig. 79).
Some of the Occidents, as soon as they ncared the
Dorymyrmex bounds, paused, and stood quite still, as
though reconnoitring the hostile quarters. The pause
was fatal, for they were attacked at once by the vigilant
sentinels, who sallied forth to a goodly distance upon
the avenue. Others seemed to recognize that discretion
is the better part of valor, and made a wide detour of the
skirmish line of the little vixenish raiders. It was plain
that the Occidents thoroughly knew the qualities and
temper of their involuntary guests, and regarded them
with wholesome distrust, not to say fear.
The result of the guerilla warfare above described was
rather remarkable. The next morning, upon visiting
the ground, I found that the Occidents had abandoned
their old avenue, had cut down and around the Dory-
myrmex colony, and made an opening on the edge of a
slight ridge several inches beyond the disputed terri-
tory, but still in the line of the avenue they had been
using. A little of the pains required for this last would
have cut out and carried away the whole Dorymyrmex
/ V V
nest space, whose contingent of diminutive warriors
could have been overwhelmed in a moment bv the
legions of their huge hosts. Subsequently the Occidents
made an amusing retaliation upon their wee tormentors,
for I found their nest literally buried under the dirt
excavated from the new gangway, and dumped upon
their gate and moundlet. It was a fitting and laugh-
able punishment for the little churls, who, however,
would probably cut their way out, unless the process
These incidents will suffice to show that courage is a
general characteristic of ants. But one finds herein the
same temperamental differences that mark the tribes
of men. As there are nations justly described as war-
WARRIOR ANTS, AND THEIR EQUIPMENT
like, and others as peaceful, so among ants there are
species whose natural disposition is bold, fierce, com-
bative, and species that, in comparison, are timid and un-
warlike. An apt illustration of this is given by Professor
Wheeler [W. 1, p. 527], who found lodged in the leaves
of a Mexican tillandsia, colonies of ants of several dif-
ferent species. They seemed to be living on good terms
with one another, but were not so complacent toward
their human observer. While tearing the leaves asunder
the little Cremastogasters (brevispinosa) attacked him
vigorously, though their lilliputian stings and mandibles
hardly got through his outer skin. But the huge
Camponotus abdominalis rushed out in a body, and the
powerful jaws of the soldiers, reinforced by the copious
formic acid batteries of the whole company, compelled
him to give up his investigations. Two species of the
grotesque genus Cryptocerus were as gentle as lambs,
resting quietly on his hands and clothing. A group of
timid little Leptothorax petiolatus took to their legs;
while the superb, wasp-like Pseudomyrmas (gracilis) made
dashes at the investigator from among their glistening
larvse and pupa?, but hastened back, as if afraid to leave
their young. Here, at one view, could be noted the dif-
ferences in natural combativeness which one sees among
creatures of a larger sort.
Professor Forel, during his visit to the United States,
observed a small troop of sanguine ants attack a com-
munity of Formica siibsericea. At the mere assault
the latter betook themselves to flight, carrying their
larva? and pupse. [F. 6.] Even these they permitted
the slave-making marauders to snatch away without
serious resistance. In all his extensive and varied ex-
perience, Forel had not seen such complete and absurd
x * 199
cowardice. And yet the Subsericeas observed by me
and there are numerous colonies on my own country
place while they are not conspicuous for warlikeness,
are quite normal in their general courageousness. But
there are no slave -making species in their vicinage.
Their spirit has not been cowed through successive con-
quests by stronger and fiercer foes (Fig. 81). The}^ are,
therefore, not open to that strange seizure which may
beset ants as well as men, even the stoutest-hearted men,
known as a panic.
With ants, too, as with men, circumstances influence
both personal and communal courage. Formica fusca,
Fig. 81 SLAVE-HOLDING ANT
Going home with a plundered cocoon and a dissevered
head of an ant clinging to leg
one of the favorite auxiliaries or so-called slaves of the
sanguine slave-makers, has been observed by Wassinann
and Forel to be more courageous with them than when
nesting alone. Backed up by their redoubtable leaders
and associates, thevact as do men under similar circum-
stances, and stand to their weapons when otherwise
they would retreat. Independent of other and tactical
WARRIOR ANTS, AND THEIR EQUIPMENT
considerations, military men know the value of "rc-
seives" in bracing up the troops on the firing-line by
an appeal to the confidence that springs from conscious-
ness of an efficient support to fall back upon. It is
curious to note an experience somewhat like this in the
above reflex as operative among ants.
Naturally our thoughts, when turned to the subject
of war, call up the forms of males as the sole or chief
actors. So it is among men and the wild and domestic
animals that we know best. Certainly there are women
not lacking in belligerency, and when congregated in
riotous mobs they have at times been fiercely combative.
But in organized and communal fighting females rarely
have had a part. War is a male occupation.
In ant communes we strike another atmosphere. The
difference between the male temperament in ants and
in the human species is so great that we cannot frame
a just basis for comparison. In man the masterful
qualities have developed in the male and the dependent
ones in the female. It is the reverse of this among ants.
The male is a dependent, in whom capability of self-
support and self-defence has been obliterated. The
female and it must be remembered that all workers
are females in a stage of incomplete development is
the sole efficient servitor of the commune in peace and
The queen, or productive female, is The Commune in
germ. She is THE AXT, in whom are centred all the
qualities and functions of the whole race, save the power
to quicken the eggs. She is the true "war lord' in the
early stages of establishing a nation, and only when
the foundations are securely fixed does she abdicate, and,
effacing herself as a chief, retire from the toils and ex-
posures of leadership, and give herself up to the duty of
adding to the communal numbers, in and for which she
is sheltered and protected by the force of the entire
Her original offices as builder, purveyor, caretaker,
defender, pass over to the workers, and therewith the
war-like spirit which communal safety requires. When
the young males come upon the scene they are treasured
for their one necessary function, and then cast off. They
are not specialized as soldiers; nature has forbidden that
by denying them defensive weapons; for a like reason
they cannot work. They are simply dependents-
nature's beau ideal of the "gentleman loafer." Thus
it comes about that all emmet warriors are veritable
It has been said that an ant commune presents the
phenomenon of a social government where every active
citizen or "citizeness/ if one fancies that term is a
warrior. Yet herein, also, we may record an exception
that approximates them more closely to our own con-
ception of a military organization wherein the fighting
members are a separate class. One might use a stronger
word: for in the course of human history soldiers have
been not rarely a real caste. And, in truth, it amounts
to that even now in some nations. Perhaps the "anti-
militarism" that marks social agitations in certain quar-
ters is simply a modern industrial swing away from the
old bondage to a soldier caste.
Be that as it may, nature shows us in ant communes
examples, in various stages of development, of the dif-
ferentiation of ordinary workers into soldiers. In many
species the line of demarcation is not sharply drawn,
but appears in the greater size, especially of the head
WARRIOR ANTS, AND THEIR EQUIPMENT
and jaws, the important parts in attack and defence.
But in other species the process has advanced to a
distinct soldier caste whose characteristics are most
In the genus Pheidole we have species in which the
functions of workers and soldiers are clearly separate.
Of a Texas species, Pheidole instabilis, the workers alone,
though of comparatively diminutive size, manage the
colony's affairs. Thev forage for seeds or dead insects,
which they drag to the nest. They dig out the galleries,
carry about and tend the larvae and pupae, even the huge
female ones, and assist to cut out the callows from their
pupal envelopes. On the other hand, the soldiers have
an office for which their abnormally large heads and
strong jaws peculiarly fit them: they are the guardians
of the nest. They form a sentinel cordon around the
young brood and the callow antlings. They are stolid
and inactive, keepers at home, but eschew the task of
tending the communal babies. Their heads are so large
proportionately to their bodies, that if turned upon
their backs they are often unable to right themselves,
and if not relieved may die practically standing on their
This big-headedness, with its corresponding develop-
ment of the jaws, however, has led to a peculiar service.
The soldiers act as the communal carvers or trenchers,
and crack the shells of the oily seeds and the tough,
chitinous cases of the insects which the foragers collect
for the commissary department (see chap. iv). Some-
what oddly, they abstain from levying toll upon the
food supplies thus made ready for general use, but draw
their rations directly from the comminuted stock in the
workers' crops. [W. 6, p. 4.]
This genus has representatives in the Eastern United
States Pheidole pennsylvanicus and Ph. meyacephala.
The author has made some notes of the habits and nest
architecture of the former in his studies of the Occident
ant. [McC. 4, p. 140.] As it is a harvesting ant, the
Fig. 82 THE FLORIDA HARVESTER (POGONOMYRMEX CRUDELIS)
1. Soldier worker. 2. Uiiwinged queen. 3. Worker major.
4. Worker minor.
massive heads of the soldiers may be utilized for the
same service as those of P. instabilis. Another ant,
the Florida harvester, Pogonomyrmex crudelis (Fig. 82),
which garners seeds, has large-headed soldiers, which
perhaps may be found also to unite the function of po-
licemen with that of trenchers. [McC. 3, pi. ix, Fig. 41.]
The genus Atta contributes examples of species pro-
vided with soldiers with exagge r ated cephalic enlarge-
WARRIOR ANTS, AND THEIR EQUIPMENT
ment. In a Texas species, A tta fervens, the duties of
these soldiers appear to have been specialized until they
are as distinct as are their forms. This species is the
well-known cutting or parasol ant of Texas, whose de-
foliation of .trees, for the enrichment of their cavernous
"mushroom gardens," makes them a decided pest to
farmers. In my studies of their habits it was noted
that during their foraging excursions the vast columns
of leaf-cutters and carriers were marshalled by the big-
headed soldiers. Both on the raid and on the return
they accompanied the marching ranks, appearing to take
no part in the actual work of the expedition, but moving
back and forth along the flanks, after the fashion of
scouts and pioneers, or of an official guard and escort.
[McC. 10, p. 36.]
Other examples might be cited, 1 but the above suf-
ficiently illustrate the fact that in the military govern-
ment of ant communes we meet with a feature analogous
to that well-known characteristic of human societies:
the differentiation of the functions of police and defence
into a special class, or caste, known as soldiers and
How far in tnis natural arrangement the industrial
element among the ant citizens is dominated bv the
soldiers is not known- -at least, to the present writer.
Within certain lines as, for example, submission to their
soldier escort by the leaf-cutters of Atta fervens they
permit the exercise of legitimate authority. But they
seem able to control the situation when so inclined.
Professor Wheeler has seen the workers of Camponotus
ferrugineus kill and dismember their soldiers in a case
1 A remarkable case is that of Poh r ergus, which is referred to in
chap, xiv "The Founding of Slave-Making Ant Communes."
where the food supply of the commune had become in-
sufficient. They thus at once showed that their guards
were not their masters, and that workers held the first
rank in their social order, though of course they in-
cidentally demonstrated their lack of anything like "the
quality of mercy' 1 in their relations to their associates.
In short, we may conclude, with reasonable assurance,
that the government of ant communes is not a military
despotism, and that soldiers, when specially differenti-
ated, are simply a co-ordinate and subordinate part
of the social organization.
The weapons (Fig. 83) with which ants carry on their
wars are placed at the extremities of the body. A pair
Fig. 83 MANDIBLES WHICH ARE USED AS WEAPONS BY WORKERS
OF AGRICULTURAL ANTS
of movable jaws, or mandibles, are attached by strong
muscles to the face. They are palmate, toothed along
the receding edges, terminating on the inside margin in
a large pointed tooth or tusk. These two opposed in-
struments, working against each other, form the compo-
site tool and war-weapon of ants. With these they dig
their galleries in the earth, or carve them out of wood,
cut down grass, defoliate trees, seize and cut up food of
all sorts. Being palm-shaped as a rule, the gathered
and comminuted material can be compressed into their
hollows, and so carried as conveniently as in a basket or
barrow. As the muscles permit the application of much
or little force at the insect's will, the mandibles can be
WARRIOR ANTS, AND THEIR EQUIPMENT
clamped together with power enough to break and tear
tough fibres, or approximated so gently that the soft
eggs and tender larva? can be borne about as daintily
as an infant in a mother's arms. Thus they aptly com-
bine some of the qualities of the human hand with those
of a beast's jaws.
It is this instrument for the two mandibles work
together as one organ that serves ants effectively as
the chief weapon in their various combats; it is at once
war-club, battle-axe, and sword; it will decapitate a
foe with the facility of a sabre or guillotine, will sever
a leg or antenna as deftly as a scimetar, or crush a skull
in its formidable vise as would tomahawk or club. It
is terrible to see, in the fierce encounter of emmet war-
riors, the cruel havoc wrought by this implement.
As effective, perhaps, and fatal, but less apparent in
its operation, is the weapon attached to the opposite
Fig. 84 USING THE STING IN FLIGHT
Occident ant in duel with fetid ant. A comrade Occident
looks complacently on
extremity. Enclosed within the vertex of the abdomen
is an arrangement of organs known as the sting (Fig.
84). In one great division of the ant genera these are
veritable stinging organs, like those of bees and wasps.
For example, in the agricultural ant, in which the author
has studied them most carefully, they consist of the
poison gland and sac, the accessory organ or oil sac, and
the stinging apparatus. These are all situated in the
lower portion of the apex of the abdomen, close to the
ventral surface, and are covered by the final ventral plates.
The word "sting" as commonly used cannot be applied
to any one organ, but expresses rather a combination of
three organs, one of which, the sting-case, is single; the
others, the stinging-prickles and the out-sheath which
encloses them, are double. They are supported within
the apex of the abdomen, and are operated by a most
ingenious system of levers and muscles. The sting-case
is somewhat curved toward its chiselled point, which
resembles a carpenter's gouge. In the act of stinging
this gouge makes the first incision.
The two shafts of the stinging-prickles in repose are
contained within the sting-case, but are thrust out
alternately when the ant stings, entering the wound
made by the gouge, aggravating it, and injecting the
poison. The prickles are slender, sharp, hollow tri-
angular chitinous rods with barbed points. The pos-
terior parts, or shafts, which lie alongside each other
within the sting-case, are straight below, but at the top,
or anterior part, are bent away from each other, form-
ing the bows. Each stinging-prickle thus consists of a
shaft and bow which, as operated in action, serves the
purpose of a spear, or lance, and bow and arrow. The
force of human muscles by which the ancient artillery
was made effective has its analogue in the protruder
and retractor muscles of the ant, attached to the bow
of the prickles, by which the shafts, with their pair of
six-barbed needles, are forced out and drawn back.
WARRIOR ANTS, AND THEIR EQUIPMENT
The above forms substantially what is the piercing
mechanism of the harvesting ant's sting. 1 But the
ant warrior does not depend upon the simple thrust of
its lance to place its antagonist out of action. The
poisoned arrows and the chemical projectiles of human
warriors have also their representatives in the equip-
ment of emmet soldiers. Situated above the stinging
mechanism, and communicating therewith by a conduit,
is the poison sac with its included gland. Herein is
secreted a virulent acid which, being forced by muscular
pressure into the hollow prickles, is carried down and
into the incision made by the point, perhaps through
an orifice in the barbs.
Associated with this is the accessory organ or oil sac,
located also just above the sting-bow. Its duct, through
which issues an oilv secretion, enters the throat of the
sting-case close beside the opening of the conduit of the
poison sac. Both ducts pass for some distance into
the case, separated only by a delicate chitinous fold,
finally to terminate together. The oily secretion, min-
gling with the acid poison, probably tends to distribute
it over a larger surface, with corresponding ability to
injure; and may add to its power to adhere to and pene-
trate the attacked surface. Perhaps, also, it serves as a
lubricant to the sting.
In a large number of ant genera, including many with
which we are most familiar, as Formica, Lasius, and
Camponotus, the stinging organs are rudimentary; that
is, they are without the sting proper. They have no
lance or arrow to thrust into their foes. Their stinging
1 A more detailed description would be out of place here, but
special students will find a complete histological description in
the author's Agricultural Ant of Texas, pp. 171-102 and plates.
organs, otherwise complete, arc operated as acid batteries
from which shoot out poison streams. These enter the
system of antagonists by the joints of limbs or other
unarmored parts, and produce paralysis and death.
Camponotus will eject this formic acid in such quan-
tities as to be visible to the naked eye. When large
numbers of the Alleghany Mountain mound-makers are
irritated and given some object to attack, the fumes of
the strong acid emissions are soon perceived. Lord
Avebury found, after disturbing the nest of a species of
Formica in Switzerland, that a hand held as much as
ten inches above the ants was covered with acid. Their
mode of punishing a human victim is to scrape away the
outer skin with their mandibles and eject their poison
upon the abrasion, which causes a painful smart. In
combat thev drench their adversaries from these
formidable acid batteries. Thus ants are effectively
equipped for both defensive and aggressive war.
HOW ANTS CARRY OX WAR
NO living creatures known to the writer so closely
resemble man in the tendency to wage pitched
battles as do ants. Vast numbers of separate species,
or of hostile factions of the same species, may be seen
massed in combat, which is continued for hours, days, or,
in at least one case noted, for over a week. Some of
the most extensive battles observed have been fought
between neighboring communes of Tetramoriunica'spituni,
a small dark-brown species common to America and
Europe. It abounds in and around Philadelphia, where
it is popularly known as the " pavement ant, " on account
of its habit of making its nest under the bricks and flags
I have often seen them engaged upon the large paving-
flags that cover the walk from the manse through the
grassy terrace fronting the church at Chestnut and
Thirty-seventh Street. They fairly blackened con-
siderable spaces of the gray stones with the vast numbers
of the combatants. Some details of one of these fights
will give a fair type of all. In the centre the warriors
were heaped several ranks high. The mass seemed to
boil with the intensity of the action. There was no
appearance of orderly array or "line of battle ' ; forma-
tion. It was literally a melee, recalling descriptions of
battles in the clays of chivalry, when armored warriors
fought hand to hand.
From the central mass the numbers gradually dimin-
ished until, as spaces opened in the surrounding fringe
of the fight, one could see small groups of combatants
scattered over several square feet of surface. Most of
them were duels; but trios, quartets, quintets abound-
ed. In one case six ants were engaged with one; in the
centre, two were tugging with interlocked mandibles,
and five others were grouped around, like spokes in a
wheel, each sawing or pulling at a limb of the un-
fortunate central integer, who was being torn to pieces.
Here and there a larger group would be piled upon one
another, heaving, pushing, tugging, like the athletes of a
football rush, but with mortal intent.
The duellists seized each other by the head, frequent-
ly interclasping mandibles, and pulling backward or
swaying back and forth. It was literally a "tug of
war." Again, one would have her antagonist grasped
by the face above the mandibles, which placed the
latter at a great disadvantage. In such and other cases
both ants would often be reared upon the hind and
middle legs, with abdomens turned under and stinging
organs out-thrust, making vicious stabs at one another.
All over the field disengaged ants were running about,
excitedly seeking a foeman, incessantly stopping to
challenge with antenna?, then hastening on until a hostile
party was met, when at once the two locked mandibles
and fell to. Many ran to and fro, stopping now at one
group, now at another, to nip an abdomen, gnaw a leg,
or snap at face or antenna , and then would rush away
to some more promising service.
Meantime, from the gates of the warring communes
HOW ANTS CARRY ON WAR
-small openings on the edge of the paved walk two
streams of recruits were pouring toward the scene of
strife. Their bodies fairly quivered under the in-
tensity of their emotion as they ran along, reminding one
of human crowds hurrying to a fire or a fight. As the
two opposing streams met and intermingled, ant tackled
ant in deathly grapple, and thus the fury of the battle
Of one party, distinguished as "Alpha," a long file of
warriors was running from the field along the trail to
the home nest. They challenged briefly every passing
fellow, and pushed on. I conceived, as a solution of
this conduct, that this was a file of messengers bearing
from the field an appeal for recruits. They certainly
were not running away. All appearances and all ex-
perience were against that inference. At all events, the
ideas of a recruiting detail, a call for relief, fell in with
the analogy of a human battle-field so strongly suggested
by the scene before me.
From the central point of the fight, as first seen at the
edge of the walk nearest the "Alphas," the vortex of the
combat gradually shifted toward the gate of their an-
tagonists, the "Gammas." At first it seemed as though
that army were being slowly pushed from the field.
But if so, the tide of battle afterward turned; for victory
finally remained with them, as far as it could be adjudged
to either party. At this period the field of battle was
spread over a space two feet long by six inches wide, the
fighters grouped most thickly about two centres, be-
yond and around which the walk was dotted with many
duellists and small contending groups.
At 12.30 P.M. the battle, which had begun at 8.30 A.M.,
was practically over. The "rear guard" of the Alphas
were continually dropping into their home trail, and
numbers of Gammas were filing to their gate in a sluggish
way. Not a recruit from either side was coming to the
field. The dead lay in little windrows where the tide of
battle had left them, or whither they had crawled to die,
or the rising breeze had borne them. Here and there
among them were ants still living but fatally hurt,
struggling to drag their mutilated bodies from the mass.
Even so, two enemies, when forced together in this grim
fellowship, would grip one another and roll and strain,
giving their waning strength to a last hostile tug.
It was a not inapt reminder of after -battle scenes
among men. Only, there was no hospital corps separat-
ing the dead and bearing off the wounded; no surgeons
plying their ministry of bodily help and repair, nor
chaplains their ministry of spiritual consolation. Dead,
dying, and wounded were all alike abandoned by their
late comrades, a number of whom, on both sides, were
now gathered around the pats of butter and sugar which
I had vainly placed in hope to lure them from fighting.
The refection which they refused during the heat of
combat was eagerly accepted to refresh themselves after
the toils of strife. That, too, was a quite human-like
scene, for soldiers must eat and drink when the dreadful
stress of battle is eased. However, there was no at-
tempt by the living ants to feed upon the dead, as one
sees under other conditions.
The state of the wounded was pitiful, an exhibit in
miniature of the dreadful aftermath of human battles.
For example, here was a warrior whose middle leg on one
side was sound, the hind leg cut off at the thigh, the
front leg at the trochanter a mere stump. On the
opposite side the hind and middle legs retained all the
HOW ANTS CARRY ON WAR
parts, but were broken, curved, useless, like paralyzed
limbs, the joint effect of its enemies' mandibles and acid
batteries. Its antenme were both paralyzed, bent up,
and motionless. It was thus bereft of all sense of direc-
tion, and all power of communication and progressive
motion. It lifted up its head again and again in vain
efforts to rise. It shook its stumps of legs, rolled upon
its side, rested a moment, and then with ruling passion
of emmet tidiness, strong even in death, struggled to
support itself upon its abdomen, and tried to cleanse
(perhaps to heal) with its tongue a fore leg.
Its adversary had not a whole leg left, its most per-
fect one being a middle leg that had lost the foot. All
the others were torn off to the thigh, or the tibia, or close
to the body, and one antenna was gone. There the
two foes floundered close together, dismembered and
dying, left to their fate by the comrades who had
mutually helped in the achievement of this great victory.
Like examples were scattered over the field, from which
the rage of conflict had died away, except as it lingered
here and there in duels or small groups of combatants
doggedly fighting out their controversy to the death.
From time to time various groups had been removed
from the mass, and placed in artificial nests prepared
with a view to special experiments. Among these was
a pair whose fate I wished to follow separately. One
ant, that seemed to be quite sound, was interlocked with
an antagonist much damaged, having lost several legs
and an antenna. But it had tightly gripped in its jaws
a leg of its adversary, who snapped at its antagonist's
neck and face, and squirmed and doubled, and strove,
with many contortions but in vain, to disable its oppo-
nent and get free.
As it promised to be a long engagement, I left them
alone in their box and turned to view the battle. When
I next saw the pair the duel was finished. The maimed
warrior lay dead and near by the victor was seated
upon a pebble nonchalantly preening her ruffled coat,
and with comb and tongue and spined limbs was re-
pairing the damage of battle.
I placed her near the Gamma gate, wishing to see if
she could find her way home, and what would be her
conduct and reception. She ran about in an involved
path for nearly fifteen minutes, covering a great space,
and at last fell upon the regular trail to the nest used by
the ants of that commune. But as she showed no
familiarity with the field, I concluded that she be-
longed elsewhere, and transferred her to the vicinage of
gate Beta, one of the outlets in the territory of the Alpha
She circled around in an irregular course, always
drawing a little nearer to Beta. In her march she met
a pair of combatants, exchanged antennal salutations,
and passed on. Presently she came upon another duel,
again challenged, and again passed on. She acted as if
lost, but kept bearing gradually toward Alpha gate.
Now she met several scouts who challenged her with
some evident doubt as to her status, but let her go.
Next she was stopped by a group with whom, plainly
enough, was exchanged a satisfactory password and
"How d'e do!" and then she was off with a joyous trot.
She had struck the home trail ! In a moment she dived
into the gate. Home at last home from the wars!
Doubtless there may have been, on her part, a passing
satisfaction like that which Burns sang in The Soldier's
HOW ANTS CARRY ON WAR
" When wild war's deadly blast was blawn,
An' gentle peace returning,
Wi' mony a sweet babe fatherless,
An' mony a widow mourning."
But we may be sure it was but a fleeting emotion,
and that on the part of the commune there was neither
for her nor for any other returning braves a civic
demonstration of "Welcome home from war." They
glided simply and naturally, as though from a night's
rest, into the regular routine of communal duty, and
there was no more to-do about it. Every active member
of society stood ready to take the same risk, do the same
service, make the same sacrifice. What occasion was
there for special hero-mongering ?
Verily; and when human commonwealths have reach-
ed the same level of patriotism and civil devotion, citizens
may fairly take a like attitude. But until then grati-
tude for and due recognition of true heroism in army and
navy must be held as a civic virtue, and the poet's,
admonition be in place:
"The brave, poor soldier ne'er despise,
Nor count him for a stranger;
Remember, he's his country's stay
In day and hour of danger."
What was the cause of these conflicts between insects
that apparently ought to have been close friends? In
at least one case noted the quarrel clearly arose over a
find of rations. The centre of the warring mass was
some fatty matter which had been thrown on and around
the seams of a brick pavement through which a large
formicary had cut its gates. From the battle-field a col-
umn of Tetramoriums three or four lines deep stretched
along a depression made by a shallow surface drain to a
second nest under a gate that led through a party- wall
into a house yard.
Apparently, the ants from the curb colony had fallen
upon the unctuous treasure which had dropped by their
door, but had been disturbed in their " feast of fat things"
by stragglers from the gate nest. These were attacked;
others came, and were also attacked. Messengers ran
to the gate nest for reinforcements; fresh squadrons
issued from the curb colony, and so the battle grew.
[McC. 11, p. 158.] It is probable that many like con-
flicts arise from rivalries for the possession of food ; and,
as in the above case, it is almost sure that a communal
war springs out of a quarrel between a few', who, appeal-
ing to civic partisanship, finally enlist in their contention
the two communities represented. Of course, conflicts
between separate genera and species are readily ex-
plained by race antipathy.
Perhaps the most usual cause for the wars waged be-
tween our city Tetramoriums is the irritation produced
by the encroachment of the mining workers upon their
neighbors in the enlargement of their living-quarters.
This is the more likely, as the most common period for
the battles is the early spring, when the demand for
larger room is greatest for the accommodation of the
rapidly increasing young of the commune. The galleries,
nurseries, and living-rooms for the numerous males and
females are pushed out with such fervor that the ex-
cavated pellets rise into heaps and moundlets around
the nest gates. In such conditions the overlapping
of the new boundaries is inevitable, and in the tense
nervous strain and high communal pressure under
which the work is being pushed, the contact between
HOW ANTS CARRY ON WAR
the rival parties is almost sure to be hostile. [McC. 3,
As the season advances, and the excitement of home-
building and the keen fervor of communal parentalism
abate, the war fever cools down, and peace prevails.
Whatever be thought of the above as an explanation of
the wars of our city Tetramoriums, it at least opens to us
a secret chapter in the life of ant communities that
awakens unusual interest. It is the story of under-
ground wars. The surface combats are sufficiently intense
and tragical. But there is a mystery about the battles
waged within the dark caverns of the communes beneath
the surface that clothes them with an air of romance.
Here are mining and countermining, just as one sees
it in engineering campaigns of men, without the horrible
accessories of explosives. Here a gallery is broken
through; a sharp engagement follows; the assaulted
party rallies to the defence of the works; the victors
have pushed their way in; the vanquished fall back.
But behind them a working detail has thrown up a strong
barricade, behind which the besieged rallv. and the bat-
TJ */ /
tie goes on anew. In the case of such a "thief ant >;
as Solenopsis fugax, whose diminutive commune is
constructed within that of some far larger host, the
mining tactics and the spirited resistance may be ob-
served in artificial glass nests, and they are extremely
interesting to watch. A rather remarkable feature of
the communal habits of this ant is that its swarming
does not occur, as Forel observes, until September, long
after that of its host ants (July, August). Thus they
can get to the surface safely and swarm undisturbed,
that belligerent period of their huge neighbors being
overpast. [F. 5, p. 499.]
But in most cases no sufficient reason appeared for
the frequent wars between the pavement ants. They
are of one species, and in some cases, as it seemed to me,
of one commune. Why should they fight? To be sure,
civil wars are, unhappily, not unnatural to human
societies, and indeed to social aggregations of humbler
creatures. But somehow one expects better things of
ants, even though their "ways'' may not be held as
"wise' ; in all things as those of Solomon's harvesters.
Yet almost the first act of our city Tetramoriums, upon
issuing from their winter quarters, is to engage in fierce
war with their neighbors or fellow-formicarians. At
times throughout the season these hostilities were re-
If, as we conjecture, the individuals be of one nest, is
this nature's mode of distributing the species from the
home centre, by causing the worsted party to emigrate?
Or, supposing the combatants to be of separate ad-
joining communities, is this wasting pugnacity a sort
of emmetonian malthusianism by which the surplus
population is reduced and kept within due bounds, much
to the comfort of survivors, and more to the satisfaction
of man? Whatever theory or conjecture one adopts,
he is apt to conclude that it is well-nigh as hard to find
a really good reason for wars of ants as for many wars
Another perplexing problem here arises: How do
these ant warriors recognize friend from foe? The
device of variant uniforms does not serve in this case,
for they are all alike. Take a group of combatants in
the hand and put them under a magnifier, as one can
readily do, so intent are they upon mutual destruction.
The most careful observer can note no difference be-
HOW ANTS CARRY ON WAR
tvvccn individuals of the two factions, yet they do
infallibly and instantly distinguish their nest-fellows
from the enemy. This is done by the antenna), which
are kept in constant motion, the tips describing sundry
curves. At a meeting between ants these organs touch
and embrace the face; if the parties be friends, they pass
on; if foes, they straightway begin to fight. The new-
comers, thronging to the battle-centre, where hundreds
are struggling in a heap that is chaos to human eyes,
but presents no difficulty to emmet senses, plunge into
the seething mass and instantly recognize and join com-
bat with their enemies. How is it done?
Thirty-two years ago, during the summer of 1877,
while pondering this problem, it occurred to the writer
that this recognition was based upon a certain odor,
emitted in different degrees of intensity by the respective
factions, or upon two distinct characteristic party odors.
The degree of odor or difference in odors, he thought,
might be dependent upon some peculiarity in the phys-
ical condition or environment of the antagonists. Sup-
posing that there were any truth in this theory, it further
occurred to him that the presence of an artificial and
alien perfume strong enough to neutralize the distinctive
animal odors, or degrees of odor, and environ the com-
batants with a foreign and common odor, would have a
tendency to confuse the ants, and disturb or destroy
their recognition of the distasteful and exciting element.
In which case he conjectured that the result might be
their pacification and reconciliation. Experiments were
made to test this hypothesis. [McC. 17, p. 17.]
A number of warring Tetramoriums, taken upon a
flower border, were placed together in a large glass
vessel upon some soil. The jar was vigorously shaken
so that, if possible, the mechanical agitation might
separate the combatants. The ants emerged quite un-
affected by the miniature earthquake, to continue or
recommence the fight. When the surface was well
covered with them, and the battle was again at its
height, a ball of paper saturated with cologne water was
introduced into the jar. The ants showed no signs of
pain, displeasure, or intoxication under the strong fumes.
Some ran freely over the paper. But in a few seconds
the warriors had unclasped mandibles, released their
hold of enemies' legs, antennae, and bodies, and, after a
brief interval of seeming confusion, began to burrow
galleries in the earth with the utmost harmony. There
was no renewal of the battle. The quondam foes dwelt
together for several days in absolute unity and fraternity,
amicably feeding, burrowing, and building.
This experiment was followed by others, varying the
conditions and the individuals, but holding to the
species. The result was always the same with Tetra-
morium ccespitum. The perfume of the cologne proved
a complete pacificator of the contending parties, and so
far verified the theory. The alien odor neutralized the
distnctive nest odors which had served to identify
friends and foes, permitting them thus to return to their
normal neighborliness ; or in some way had mollified the
hostile parties, and transformed them from enemies into
Similar experiments were tried with colonies of car-
penter ants taken from the Alleghany Mountains and
from Logan Square, Philadelphia. These pointed to a
conclusion just the reverse of the above. Whatever the
cause a failure of the experimenter in arranging his
conditions, or the presence of some disturbing element
HOW ANTS CARRY ON WAR
that was overlooked, or because one or both parties were
too far saturated and seasoned in their own native nest-
odor to respond to the cologne treatment the fact was
that the experiments led to opposite conclusions.
However, I had little doubt then, and have none now,
that the original inference was substantially true in the
case of wars between separate communes. The ants
were recognized by a special odor which they absorbed
during residence, and which was stronger or weaker
according to age and environment and conditions un-
known. How acute and delicate and accurate must
be the sense organs seated in the antenna?, which are in-
struments of recognition, the facts related will show.
"He does not carry the odor of my species, my com-
mune, or my caste. Therefore, we will fight!" To a
human philosopher meditating upon these things, it
seems a small difference on which to divide two such
closely related creatures into hostile camps. But may-
hap he who counts this for abatement of the common
fame of ants for wisdom might find, in the history of
human wars, originating causes as insignificant and
ALIEN ASSOCIATES AND AFFINITIES IN ANT
THAT "no man liveth to himself is an aphorism
not to be questioned in human communes. That
no community lives to itself is equally true. And this
applies to ants. Their societies are established in the
vicinage or in the midst of numberless creatures, most
of them, like themselves, free citizens of that wild life
which nature has organized and maintains in the cul-
tivated parks and fields of men no less than in a wilder-
ness. He deludes himself who thinks that he ever is
delivered from the environment of wild things. Of the
large and grosser sorts, it may be; but civilization never
will tame or exterminate the innumerable hosts of minor
creatures, seemingly as wild now as in the primitive
Eden, that inhabit our day-world and, even more, our
The ants are examples of this. They find and keep
a foothold everywhere. I have surprised immense com-
munities in the heart of great cities. I have shown an
American farmer, who boasted in the tilth of his acres-
under the plough since the first English settlements-
that he could scarcely put down a foot in a walk through
a field without placing it upon a little commune of
These cases do not stand alone. A naturalist would
ALIEN ASSOCIATES AND AFFINITIES
soon point out to our farmer that many other living
things have possession of his domain whose ancestors
were probably here before Columbus, and whose de-
scendants will doubtless outlive the Republic. These
are the creatures with which ants have to neighbor.
Close neighbors they are at times; sometimes hostile,
sometimes indifferent, sometimes friendly. In the course
of ages of neighboring experience, strange inter-relation-
ships have been established, presenting some of the most
interesting and puzzling features of emmet communal
life. To a few of these our attention will now be turned.
Taking up once more our mountain mound-builders,
we note certain loose relationships established between
them and some other insects in cold weather. Winter
deadens energy and subdues combativeness, and, when
severe, suspends activities. One will then come across
colonies of our common white ant (Termes flavipes) im-
bedded within the great cones of Formica exsectoides
Bunches of cockroaches are found, and sundry beetles,
with other insects, that in the adult or larval stage nat-
urally domicile in the ground.
Most of this sort of neighboring is the result of that
truce which Jack Frost enforces, and will largely dis-
appear when spring relaxes nature and insects come to
their normal antagonisms. But it shows how certain
companionships may have been formed which, at first
accidental and temporary, were found to be harmless,
more or less helpful, and in some cases highly beneficial.
Use and heredity, operating upon casual affinities and
the acquisition of a common nest-odor, may have thus
brought about those examples of symbiosis, or sym-
pathetic companionship, which exist among ants, and
between them and other creatures.
Let us consider a little more in detail this theory that
winter conditions may have influenced the formation
of communal affinities and associations between ants
and alien insects, as well as between separate species and
genera of ants. Do the facts seem to justify it? One
night, while encamped among the ant-hills of Brush
Mountain, Pennsylvania, late in August, 1876, there
fell a heavy frost that well disclosed the effect upon ants
of such temperature changes. [McC. 2, p. 284.] At
3.45 A.M. I made the round of the hills, and found their
inmates in a state of semi-torpidity. Tapping the sur-
face and stamping upon the surrounding stones, which
heretofore had always brought out a host of workers,
failed to arouse a single sentinel. I dug into one mound
eight inches before finding ants, and these showed little
activity a marked contrast with their usual mode.
Then the aphis feeding-grounds were inspected. A
white-oak tree near a stone wall, whereon numbers of
aphids were domiciled, was a popular emmet resort.
Mounting the wall, I turned the lantern light upon the
overhanging boughs. The aphids were in their places
on the leaves and branches, surrounded and covered bv
groups of ants. But all were semi-torpid. The frost
had surprised them at their feast, and left them frigid
upon the spot. Many of them had abdomens distended
by crops gorged with honey-dew, which showed trans-
lucent as the light fell upon them. In my long ex-
perience of a full generation in observing emmet ways, I
recall few more striking visions than that. If one could
only have preserved those congealed specimens for the
But as the sun returned with his wonted August
fervor, the statuesque groups began gradually to dissolve.
ALIEN ASSOCIATES AND AFFINITIES
First, with sluggish movements, slowly stirring; then
more vigorously, as the sunshine fell upon the branches;
until by nine o'clock the tree-paths were thronged with
workers, most of them repletes, and homeward bound.
So also it was on the mounds. As the sunlight pierced
the woods, and fell upon them and warmed them up,
they resumed their normal activity. The benumbing
effect of the frost upon the insects had been no doubt
intensified by its suddenness, and the high temperature
that had preceded it.
The above facts led me to studies of the winter condi-
tion of the mound-makers, which were made late in
October, 1876, and the latter part of February, 1877.
[McC. 2, p. 286.] It was found that the winter tended
to drive alien insects to the formicaries for harborage.
Lodged in one nest was found a colony of our native
termites. They were in an unfrozen part, exposed to
the sun, occupied a space of about four inches square,
and were then (February 14th) quite lively. Near them
was a large herd of roaches, a hundred or more. The
ants in the mound were not torpid, although their
characteristic vigor and activity were suspended. It
would not have been possible for the termites to hold
such a position in midsummer; they would have been
eaten. Such a cluster of cockroaches would have been
equally impossible; it would have been scattered and
destroyed. This is doubtless the general experience.
Wheeler [W. 1, p. 30] found that in Texas, during
autumn and winter, the nests of Formica gnava teem
with alien insect guests of various orders, larvae and
adult, that are rarelv seen in summer.
How shall we account for this? In the case of the
mound-making ants, there seem to be two factors, one
negative and one positive, in drawing termites and
roaches to the nests. The first is the benumbing effect
of cold, which suspends the emmet energies, and there-
with suspends hostile acts toward intruders upon their
domain. The second is the greater warmth and com-
fort of the mounds. These are built of a light com-
posite of soil-pellets and pine and other leaves, which
form more congenial quarters than the surrounding
earth. The galleries that honeycomb them are air-
chambers which mitigate the cold and conduce to natural
Besides, to errant insects abroad in the autumn in
search of winter quarters, the upraised cones of the ants
are prominent and inviting objects, the most available
for them in the vicinage. So there the rovers settle
and stay until, in the revived activity of returning spring,
the ants make the premises entirely too warm for them.
These facts have at least a conjectural bearing upon
the origin of some ant affinities and associations. The
importance of the local nest-odor, and its intimate re-
lationships with the friendly or hostile attitude of ants
toward their fellows, has already been pointed out in
chapter xi. May it not follow that the temporary
and accidental lodgment of these alien insects upon the
ants' nests may have led, in occasional cases, to the
acquisition of so much of the local nest-odor as partly to
conciliate the ants? This complaisance may have been
increased by the inactive condition of the ants in early
spring, and at least made them tolerant of the presence
of their guests. This condition, acting upon tempera-
ments specially adapted to such an estate, together with
the discovery of some mutual advantage in nourishment
or massagerie, through the shampoo dejeune or other-
ALIEN ASSOCIATES AND AFFINITIES
wise, may have developed at last into the habits of the
permanent myrmecophile. This may be suggested, at
least, as a contributory factor in the natural evolution
of a remarkable feature of ant communes.
A brief observation will illustrate the advantage
which some of the alien ant-guests find in the connec-
tion, and which must strongly tend to hold them to it
when once formed. Certain little crickets of the genus
Myrmecophila live with species of Formica and Cam-
ponotus, and a diminutive, nearly blind cockroach (At-
taphila fungi-cola Wheeler) inhabits the nest of the
Texas cutting ant. The behavior of these myrmeco-
philes shows that the surface of the ant's body must be
covered with an unctuous, highly nutritious, and, it
may be, antiseptic secretion, probably derived from the
salivaiy glands of the host-ant or other members of the
colony. This secretion is also spread over the eggs,
larvae, and pupae, and it seems to retard the development
of pernicious moulds, since these tend to grow only on the
larvae and pupae that have been isolated for several days
from the workers and queens.
Both crickets and cockroaches live by licking the
surfaces of their hosts. The former remain on the
ground and reach up to lick the legs and bodies. The
latter climb upon the backs of the large Atta soldiers
and feed from that position. [W. 4, p. 14.] The ad-
vantage to the ants may be simpty the pleasure of the
massage and the satisfaction of being clean, although
there may be other advantages now unknown. How-
ever, we shall presently see that such affinities and
associations may exist even under strong disadvantages
apparent to human observers, at least.
Among the ant-loving (myrmecophylous) beetles
found with our Alle^hanv mound-builders is a Claviger
species (Tmesiphorus costatis) collected during the winter.
Doctor LeConte showed me (1876), in his rich collection
of Coleoptera, several of these taken at Bedford and Co-
lumbia, Pennsylvania, among which were Cedius ziegleri
LeConte, and others which he spoke of as "undescribed
specimens of Homolata and an unnamed species of
Oxyopoda." These were
small brownish insects
with slight pubescence.
The most interesting of
these ant -affinities (myr-
mecophiles) was his own
species, Xenodusa (Atame-
les) cava (Fig. 85). This
is a reddish-brown beetle,
about one-fifth of an inch
long, with tufts of yellow-
ish hair-like tubes on the
sides of the abdomen.
From these hairs exudes
(By courtesy of American Museum of Natural History) Q, SWCCt
Fig. 85 THE BEETLE XENODUSA which the ants feed, as
From a colony of Formica Schau- , . , i , ,1.
fussi-inscrta. (After Wheeler) aphides, and It IS tniS
fact which attracts ants
to them or assures their toleration of them. Specimens
of this beetle were also taken bv or for LeConte in
ant-nests of unknown species in Maryland, Illinois, and
Michigan. Among these was one still held in its host's
mandibles, as if taken while in flight from the disturbers
of its nest, and clung to with unrelaxed jaws in the
alcohol which killed it. Our American carpenter ants
ALIEN ASSOCIATES AND AFFINITIES
(Camponotus) in several species and varieties are often
the hosts of X. cava.
All the beetles of this group, the Lornechusa group of
Staphylinids, are true ant-guests. They are treated by
their hosts, both as adults and larvae, quite as their own
fellows, being fed, cleansed, and carried about. Indeed,
it is said that in case of real or fancied danger, the beetle
larvae and pupae have precedence of their own young
in the ants' attention.
This is all the more remarkable because, according to
Father Wasmann (S. J.), a devoted and distinguished
observer, and perhaps our highest authority on myr-
mecophilous insects, these adopted citizens repay the
host's care by ravenous assaults upon their own brood,
devouring numbers of eggs and larvae. The effects of
this, in weakening the commune, are apt to be serious.
It works toward deterioration, as Wasmann show^s, in
another way. This brood - parisitism appears to orig-
inate a curious form of abortive individuals inter-
mediate between the female and the worker, known as
pseudogynes. They are cowardly and indolent. They
decline to dig and nurse, and trot about the nest aim-
lessly. Thus, in sharp contrast with the valiant and
active workers, they hold a sort of "frustrate existence."
How comes this about? Wasmann believes, and
seems to prove, that it is caused by the diminished care
and diet due to the queen larvae for their full develop-
ment a case of restricted growth through defective
nourishment. A brood of beetles (Lomechusa) begin
life with a brood of worker-ants. The beetle larvae, as
they appear, are not only generously fed by the ants,
but begin to feed upon their eggs and larvae ; and as they
are extremely voracious and grow rapidly, they devour
enormous numbers. This makes a great breach in the
generation of coming ant-workers. These are essential
to the commune, and the adults aim to make up the lack
by converting into workers some of their larvae destined
for queens. This results in that intermediate form,
neither worker nor queen, but a spurious female a
At the same time the infatuated ants, under the im-
pression that their guest-larvae are valuable to the
commune, lavish on them care due to their own progeny.
Thus, again, arises a neglect of the young ant queens
which stays their growth, and diverts their develop-
ment toward the pseudogvne. It is the old storv of the
cuckoo among the birds, who thrusts her egg into an-
other bird's nest, and secures for her parasitic offspring
the nurture due the legitimate fledglings.
All this goes sadly against the general reputation of
ants for wisdom. But perhaps it might modify our
censure to mark our own history or survey existing
society. Would it not be found that we have not only
tolerated but have fondled and nurtured human
parasites in official, family, and private life, greatly to
the loss of the commune? Our parasites destroy the
virility and the very life of our young, and we endure
them. They waste our resources by graft and neglect
of duty and pernicious schemes and perverted policies,
and we give them our suffrages and support. We open
our homes and our harbors to guests who repay our
hospitality by implanting among us doctrines, practices,
and persons that carry the seeds of communal disorder
and decay. Misguided by such social and political
unwisdom, it fares with us, and will ever fare, as with
ant communes inoculated with Lomechusan beetles.
ALIEN ASSOCIATES AND AFFINITIES
It might therefore be maintained, with a good degree
of verity, that social men in their communal life show
no great superiority to social insects in dealing with the
R. JS. Howe del.
(By courtesy of the American Museum of Natural History)
Fig. 86 PHEIDOLE INSTABILIS AND ITS PARASITES
1. Big-headed soldier of Pheidole Kingi Andre var. instabilis Emery.
2. Typical worker of Pheidole Kingi instabilis. 3. Male of the
same. 4. Orasema riridis Ashmead, female. 5. Male. 6. Ora-
sema coloradensis Ashmead.
parasites that infest them (Fig. 86). Especially when
we consider the vast advantage of men over ants in nat-
ural endowments, the relative unwisdom of the latter
does not bulk so largely.
A Chalcid fly, Orasema viridis, is parasitic upon colonies
of the ant Pheidole instabilis (Fig. 87). The chalcid is a
beautiful insect, decorated with metallic green, and blue,
violet, yellow, and black, with iridescent wings. This
polychromatic creature, when seen among its ruddy hosts,
amid the shining red and black seeds stored for food (the
ant being a harvester), gives a brilliant appearance to
the nest. But it is a beauty which bears the germs of
death to those who cherish the possessors.
The mother Orasema posits her numerous eggs upon
the under surface of bodies of the young ant pupae,
(By courtesy of the American Museum of Natural History)
Fig. 87 ORASEMA VIRIDIS
A parasite in various stages of development on the surface of
the ant Pheidole instabilis
near the head. She chooses for this the pupse of the
large forms soldiers, females and males not the small
workers, as having the richest store of nutriment. Here
the parasites cling and grow rapidly, feeding upon the
juices of their host. When the parasites reach the pupal
stage within two or three days they are released by
the worker-ants from their host, now a lifeless mass.
Thenceforward they are objects of special care by the
Pheidole workers, who tend them as their own offspring,
not only in the pupal, but in the imago stage. Indeed,
so great is this infatuation or delusion that, as in the
case of the Lomechusan beetles, when a nest is disturbed
ALIEN ASSOCIATES AND AFFINITIES
the Orasemas are looked after before the ants' own
brood! [W. 6, p. 5.]
The life cycle of Orasema from egg to imago is a
week or ten days. Thereafter the adults are licked and
fondled and borne about by the ants, and fed by re-
gurgitation. The guests commonly take these atten-
tions passively; but sometimes just as growing boys
resist embraces seek to avoid them. One cause of
difference between hosts and parasites emanates from
the preference of parasites for free air, which, as soon
as they mature, both sexes aim to reach. Their hosts,
having different views as to the relative values of light
and darkness, guard the exit gates, and, seizing their
guests, drag them back to the dark inner rooms.
This tendency of the Orasemas results from their nat-
ural habit of mating in the open air, after which the
fertilized females seek a Pheidole nest wherein to start
a new brood upon the round of
parasitic life above described.
After pupation the mature Ora-
semas spend much of their time
lying on their sides among the
ant larvae and pupae. They con-
tribute in no manifest way to
the welfare of their hosts, their
only interest in them being the
selfish one of securing nurture
,, , - , . , . (By courtesy of Am. Museum of Nat. Hist.)
lor themselves and a brooding .
Fig. 88 PUPA OF ORASEMA
host for their offspring (Fig. 88). a _ Just before pigmenta _
One finds nothing in the life his- tion. 6 Pigmented pupa
. ,. ready to hatch
tory of insects more puzzling
than such seeming anomalies as the above associations
between ants and parasitic chalcidids and beetles. We
are used to some such social phenomena among men,
who seem to have a perverse strain that forces their de-
velopment along aberrant lines toward disadvantageous
and destructive ends. But in these simpler children of
nature such conditions surprise us as quite abnormal.
Once in a while, however, the ants do seem to shake
off the spell that binds them and awake to the true
nature of their guests. In one of Professor Wheeler's
artificial nests of Pheidole instabilis the workers rose
upon the adult Orasemas, after they had remained in
the nest several days, and killed and dismembered them.
But a doubt remains as to whether this was due to their
discovery that the victims were predatory aliens, or to
some special stress of hunger or other cause; for the
ants also killed and dismembered their own females,
and after that reared only their fellow - workers and
intermediates, as though they purposed to spare none but
the caste that furnishes the lightest consumers and the
helpers, and to free themselves from mere dependents.
Like action on the part of worker-ants of other species
has been known in times of special stringency in the
Another of the parasitic aliens that associate them-
selves with ants is a little Dipteron fly, Metopina pachy-
condylcc Brues. While sorting out a number of larva? of
a large black Ponerine ant, Pachycondyla harpax, several
were found to have larvae of the above insect attached to
the region of the first abdominal segment. It quite en-
circled the ant larvse, like a collar about the neck, "a
kind of Elizabethan ruff." The posterior end of the
parasite was provided with a sort of suction disk, by
which that part could clasp its host so tightly that the
fore part of the body could be released and swung out
ALIEN ASSOCIATES AND AFFINITIES
of position without the creature losing its hold. [W.
6, p. 45.]
The experimenter transferred a colony of the ants to
an artificial nest for observation, and fed them with a
number of young larva^ of the ant Camponotus maccooki.
These the Pachycondylse proceeded to tear to pieces,
freely lapping the exuding juices. Then they placed the
pulpy remainders in the ventral surface of their own
larvae (as on a serving-dish), which lay upon their backs
in a chamber dug in the earth of their nest. This
chamber was so situated under the glass cover that the
actions of both ants and larvae could be observed dis-
tinctly. The ant larva? thrust out their brown heads
and began to feed. The Dipteron larvae, by some un-
known sense made conscious of the presence of food,
unloosed their heads and necks without releasing their
caudal attachment, and dipped their beaks into the
mess. Thus the two young creatures so widely apart in
structure and destiny were here united in their cradle-
life and became fellow-trenchermen.
The experiment was repeated a number of times, and
with various sorts of food. The result was always the
same. To quote the picturesque language of the ob-
server, he was always "able to witness the strange
banquet the dwarf reaching from the shoulder of the
ogre, and helping himself from the charger formed by
the trough-like belly of his host." Pieces of ant larvae,
beetle larva 4 , myriapods, etc., when served up to the
Pachycondyla larvae, were partaken of with equal zest
by larval host and guest. The latter were thus shown
to be true commensals - ' perhaps the most perfect
commensals, in the original sense of the word, to be
found in the whole animal kingdom."
As a bit of by-play, we learn that when the ant larvae
lay close together a Metopina would reach over and
help itself from the portion of a neighbor, keeping the
while its rear attachment. Sometimes, when the ra-
tions were exhausted, the Dipteron would nip the tender
hide of a near-by ant larva till it squirmed with pain,
or it would tweak its own host. The suggested purpose
of this action was to attract the attention of the nursing
workers to the wriggling ant larva, and thus prompt
them to replenish the larder.
Both kinds of larvae were cleansed by the nurse-ants,
who, if they were conscious of the presence of the para-
sites, made no discrimination between them and their
hosts. Indeed, as this species of ant is almost blind, it
seemed doubtful if they really could distinguish larval
host from larval guest, the latter possibly being taken
for a mere enlargement of the former's neck.
One hesitates, however, to accept a theory which
implies such a lack of sensitiveness in the perceptive
organs of insects commonly so highly developed. How-
ever, as ants are notoriously devoted to the genuine
antennal "tone" of society, and as the Metopinae, from
the egg onward, are imbued with the true Pachycondyla
atmosphere, the distinction between the two larva?
might readily be lost in the common odor. Yet this
would equally account for their sparing the guest, even
though its nature were perceived.
The next stage of development in the life history of
these strange yoke-fellows is equally interesting. When
the ant larva is mature, and nature stirs within it the
great unrest that precedes transformation, it sets its
spinning glands in motion, and begins to weave around
itself the brown cocoon, or closed silken sac, within
ALIEN ASSOCIATES AND AFFINITIES
which the change occurs. It moves back and forth,
around and around, issuing from the mouth-parts the
liquid silk that hardens about its snug house of change
until that is complete, when it falls into the quiet of
What becomes of its Dipteron yoke-fellow during these
movements? Surely they could not be wrought with
that encumbrance upon it? No; it has disappeared.
Whither? The mystery was solved by opening an ant
cocoon. Therein lay the Metopina safely and snugly
tucked away in its own little puparium lodged in the
posterior pole of the cocoon. It had dropped off its
host's neck, had taken station close by the opposite end,
and had been wrapped within the silken sarcophagus.
Thereto it had attached itself, had wrought out of its
own larval skin an envelope (it is not a spinner like the
ant), and in that it pupated. The quarters were large
enough for both occupants.
Now follows another interesting chapter in our story
of these humble lives. Duly the time comes when Nature
bids the transformed antling break forth from its silken
coffin. It makes with its mandibles a rent at the an-
terior pole, favored, it may be, as with other species,
with the obstetrical aid of worker nurses. It creeps out,
and, though still a callow, is soon numbered among the
active members of the commune. The empty cocoon
case is carried by the workers to the common dumping-
ground for w r aste products of the commune.
But what, meanwhile, has befallen Metopina ? In the
struggles of the antling to get out, and from the cutting
and tearing of the nurses to deliver it, has the young
Dipteron escaped injury? Fortunately it so "happens"
-if that be the lawful word that its puparium is in-
variably formed at the posterior pole of the ant cocoon,
directly opposite the anterior pole from which, as the
point next to its jaws in its recumbent position, the
antling emerges, and to which, as the point of fracture,
the strain and force within and without are directed.
Thus the little squatter sovereign, in its tiny puparium,
goes scot-free and quite unharmed to the communal
kitchen-middens, along with the abandoned cocoon of
its voke-fellow. So it befalls that, as Professor Wheeler
quaintly puts it, " after a privileged existence as free
pensioner and bedfellow to a generous host, it is unwit-
tingly carried away in the worn - out bedclothes and
consigned to the family rag-pile."
Here one must note another admirable " happening."
The period for the Dipteron to emerge falls later than
that of the ant. Therefore its hatching - place is the
emmet dump where it has been deported by its foster-
mothers, the ants. Fortunately for the newly fledged
insect, since nature has not furnished it with fit imple-
ments to break through such formidable walls, it finds
a wide and effectual door already open in the tough
cocoon. It is once more debtor to its sometime host for
that hospitality which not only "welcomes the coming,"
but also "speeds the parting guest," and crawls out of
the rent made by the emerging antling.
Thenceforth its new world lies before it. It finds its
mate. It follows the mysterious impulse of its kind,
and returns to the commune whence it came, or flies to
some other colony of Pachycondyla harpax, and, mousing
among the robust larvae thereof, drops its minute egg,
and- But there our story of the cycle of her life must
And what a wonderful story it is ! Here, if ever, one
ALIEN ASSOCIATES AND AFFINITIES
may apply Marlowe's phrase: "Infinite riddles in a little
room." It has taken the patience, skill, and experience
of the trained naturalist to trace it and unfold it to us.
But it needs no expert to note the admirable adaptations
by which a minute fly has been borne on, step by step,
in utter helplessness, through the successive stages of a
dependent being, from a mere speck of vital matter to a
winged insect, armed with the instinct to invade an un-
unknown world and propagate its kind. How great
and how infinitely exact must be that Over-Force that
dominates nature, which can include within the compass
of laws that regulate the universe a series of adaptations
like these which guard the life of a two-millimetre para-
sitic fly! It amazes, while it perplexes one, to account
for it all. Yet, in the face of great Nature's workings,
one may venture to recall the proverb of Spenser (not
Herbert, but he of the Faerie Queene) :
' 111 can he rule the great who cannot reach the small."
In contrast with our studies of the chalcid Orasema
viridis and the beetle Xenodusa cam, it is pleasing to
record that the association between Metopina the fly
and Pachycondyla the ant is apparently wholly benign
at least, under ordinary conditions. The guest does not
prey upon its host; no physical injury seems to follow
its enforced companionship; and the bare particle of
food filched from the ant larva does not tax the supply-
department of the commune or cause its workers to
stint their own dependents. The larval hosts them-
selves are as large and healthy as others in the nest, and
produce normal pupae. It is a case of "all's well that
These are but types of numerous examples of those
strange and seemingly " unnatural" companionships
with alien creatures which have grown up in ant com-
munes. The literature thereof is already large, and is
continually growing as entomologists push their in-
vestigations more widely and carefully. It would be
impossible to present here an abstract of even a tithe of
the known facts, but from the typical ones which have
been chosen the reader may fairly judge of the general
tenor of the rest.
APHIS HERDS AND ANT ASSOCIATES
A PHIDES are the alien insects with which ants have
J\. the most intimate relations. The manner in which
ants search out, attend, protect, and domesticate these
creatures need not here be repeated ; l but as these con-
sociated relations are so widely and popularly known, it
seems well to distinguish them from other ant guests
and associates by a more detailed description than could
be given to other myrmecophiles.
The aphides, or plant-lice, belong to the order Hemip-
tera and the sub-order Homoptera, including such in-
sects as cicadas, or harvest flies, and the bark-lice. They
range from small to exceedingly minute, but make up
in numbers what they lack in size, and include some of
the most destructive pests known to the agriculturalist
and horticulturist. They are soft-bodied and gregarious,
and most numerous in the wingless forms. The eyes
are usually quite large and of a dark color, and the
antennae of many species long and threadlike. The
beak is two or three jointed, and in some cases as long as
or longer than the body. In the leaf-feeding species
the legs are rather long and slender. In the root-feeding
and gall-inhabiting forms the legs are short and stout.
1 See Nature's Craftsmen, chap. iii.
The wings are thin aud transparent, with dark veins on
the anterior margin.
The order Hemiptera is composed of beaked insects,
and the wings, where these organs are present, are of
the same texture throughout, and close, rooflike, over
the body. In several groups they are transparent, and
have many strong veins. In others they are tough and
opaque, and show many different colors. The head is
broad but usually short, without any neck, and has
the beak rising so far under the breast that it seems to be
attached thereto, and may be closely folded against it.
The honey-dew which aphids yield, and which gives
the occasion for their peculiar connection with ants, is
obtained by the insertion of the beak into the tender
bark of the plant. The pumping apparatus is then set
in play and the sap withdrawn into the body of the in-
sect. It is a minute type of the mode of getting sugar-
water by "tapping" the trees in a sugar-maple camp in
Vermont or Ohio. The sap thus withdrawn probably
undergoes some slight chemical change within the in-
sect. It differs in taste from the sap of the plant, having
an acrid flavor, in some degree resembling the taste of
honey, thus justifying the ordinary phrase " honey-dew."
It does not, at least as far as the author's observations
have gone, proceed from the nectaries, or nectar tubes,
as has been commonly supposed, but is a fluid excrement.
This, however, differs from the ordinary excrement, which
is a whitish, semi-solid substance, voided in long cylin-
drical strings or minute whitish balls, which roll up like
The injury produced upon the plants by this tapping
results from this attempt of the insect to procure its
natural food. The numerous punctures made within
APHIS HERDS AND ANT ASSOCIATES
a leaf cause it to shrink up, forming little rolls, or tents,
within which immense numbers of the insects dwell.
With them ants will commonly be found, attending them
simply for their honey-dew; but often they get the blame
of the damage done by their companions, an experience
that is apt to befall higher creatures. Minute as each
individual is, when multiplied by hundreds and thou-
sands the injury wrought upon the numerous leaves of
the plant is sufficient to affect their health.
When the punctures are made upon the roots they
result in little gall-like swellings, which harden, destroy
the natural function of the rootlets, and finally result in
death. When a large number of roots is thus affected,
the plant, of course, has lost its power of deriving sufficient
and wholesome nurture from the earth, and so falls into
decay. Other species of aphides secrete from a part
or the whole of the body a whitish powder or bloom, or
numerous filaments of fine cottony matter in which they
become completely enveloped.
In the various stages of development the nymphs of
some species secrete globules of honey-dew several times
larger than themselves. Sometimes the globule com-
pletely envelops the nymph. After they are moulted,
the nymphs usually find a new feeding-place, leaving
the old skin attached to the drops of honey-dew. The
moulted skins, the last moulted especially, often retain
their form so perfectly as to seem like a live nymph.
Professor Slingerland, in his account of the " pear-
tree psylla," 1 says that it ejects immense quantities
of honey-dew, which cover twigs, branches, and trunks
of the trees, and even the vegetation beneath. This
1 M. V. Slingerland, Bulletin No. 44, Cornell University Agricul-
tural Experiment Station, 1902: "The Pear-Tree Psylla."
appears soon after the 1 leaves expand, and is found
throughout the season. I have seen American for-
est trees attacked by aphids, from which the honey-
dew was flung out in such quantities that it sounded like
the patter of rain-drops as it fell upon the dry leaves and
grass beneath the infested branches. The whole surface
of the ground underneath was covered with the liquid
sweet, and thousands of ants, bees, wasps, and various
other insects had assembled to the forest feast.
The reproductive processes of aphids are extremely
complicated and remarkable, and have been the subject
of much careful study and experiment. At certain sea-
sons of the year, usually late in the summer or early
autumn, individuals of both sexes are produced, and the
females lay eggs which in some species hatch immedi-
ately. In others they remain over the winter. Sexed
aphids were formerly supposed to be the winged form,
but late discoveries show that there is not necessarily
any connection between the wings and the true sexual
organs, the wings being simply an adaptation for migra-
tion from one plant to another.
The form hatching from the egg is known as "the
stem-mother," and in the course of a few days begins
the peculiar process of reproduction known as parthe-
nogenesis, or agamic reproduction, bringing forth her
young alive and in rapid succession. This process has
been likened to the multiplication of certain kinds of
plants by slipping and budding. The offspring of the
stem-mother begin to produce viviparously in the course
of a few days, and in this way the multiplication of in-
dividuals proceeds at a most extraordinary rate, extend-
ing to several generations.
In the pear-tree psylla, for example, the hibernating
APHIS HERDS AND ANT ASSOCIATES
winter brood begin to pair and lay eggs in the first warm
days of April. The eggs are placed in the creases of the
bark, or in old leaves, or scars about the bases of the
terminal buds of the preceding year's growth some also
upon the side. They are usually laid singly, but rows
of eight or ten are sometimes found. The eggs are
scarcely visible to the unaided eye. It would take
eighty of them placed lengthwise to measure an inch.
A short stalk on the larger end attaches the egg to the
bark, and a long, thread-like process projects from the
About the middle of May most of the eggs are hatched
and the hibernating adults have disappeared. Imme-
diately after emerging from the egg the minute nymph
seeks a suitable feeding-place, and is soon at work suck-
ing the sap with its short beak, which appears to arise
from between its legs. The favorite feeding-places of
the nymph are in the axles of the leaf and the petioles
and stems of the fruit. Sometimes in early spring they
crawl into the buds. When the axles of the fruit-stems
and leaves are full, the nymphs gather in closely packed
clusters about the base of the petioles and stems. If
very numerous, they gather on the under side of the
leaves along the mid-rib. They move about but little,
and sometimes become covered with their own honey-
dew. If disturbed they crawl around rapidly.
The only time the nymphs seem to stop feeding is
during the casting of their own skins, which become
too small and give place to new and elastic skins formed
just beneath the old ones, in the ordinary method of
moulting insects. At the last moult, which occurs about
one month after the nymph emerges from the egg, the
adult insect appears.
The adult has habits quite different from those of the
nymphs. It has strong legs and wings, which permit it
to spring up and fly away quickly upon the slightest jar
of the plant or the near approach of the hand to its
resting-place. The hibernating forms gather, but quite
sluggishly, and are readily captured when found. The
summer forms fly from tree to tree, and can easily be
borne by the winds long distances, thus infesting neigh-
boring orchards and plants. The adults are also pro-
vided with a beak, with which they feed upon the tissues
of the leaves and the tender twigs of the trees. They
seem to have no favorite feeding-place.
Three or four days after transformation from the
nymph stage the adults of the spring and summer
broods pair, and egg-laying begins for another brood.
These eggs are usually laid singly, sometimes several
in a row or group, on the under side of the tenderest
leaves, among the hairs near the mid-rid or on the
petioles near the leaf. Sometimes the mother places
an egg or two in each notch of the toothed edge of the
leaf. These eggs of the summer brood resemble those
of the hibernating adults, but hatch in from eight to ten
days under more favorable conditions.
Fortunately for the safety of vegetation, aphids have
a number of natural enemies. Among the most ef-
fective of these are the well-known " lady birds," which
are beetles, belonging principally to the Coccinella. They
are small, roundish insects, generally yellow or red,
with black spots, or black with red or yellow spots.
There are many species, and they are generally distrib-
uted among the plants, and are familiar objects to those
who cultivate flowers. They live both in the perfect
and the immature state upon aphids, their natural food,
APHIS HERDS AND ANT ASSOCIATES
and their services are extremely valuable. Perhaps
it is this which has created the friendly popular senti-
ment toward these pretty beetles. It is rare to hear an
utterance of dislike toward them, and they fairly rival
butterflies as general favorites. Their young are small,
flattened grubs of a bluish or blue-black color, spotted
usually with red or yellow, and furnished with six legs
near the forepart of the body. They are hatched from
little yellow eggs laid in clusters among the plant-lice,
so that they find themselves at once within reach of their
prey, which from their superior strength they are able
to seize and slaughter in great numbers.
Another enemy of the aphis is the golden-eyed lace-
winged fly (Chrysopa perala), which is of a yellow-green
color, and has four wings resembling delicate lace. It
gives out an offensive odor. It suspends its eggs by
threads in clusters beneath the leaves or where plant-
lice abound. The larva is a long and cylindrical grub
provided with jaws moving laterally, which perforate the
body with a hole, through which it sucks the juice of its
victims. It requires only one minute to kill the largest
aphis and suck out the fluid contents of the body.
This sketch of the life history of aphids will be appre-
ciated by those who have learned the story of their rela-
tions to the ant. That their value as food producers
should have been discovered and utilized by such in-
veterate scouts and scavengers and cosmopolitan feeders
as ants is not strange. But that they should have ac-
quired the art of "milking" them; should have learned
to seclude them for their own use within walled en-
closures, as sheep within a fold ; even to rear them with-
in their own bounds and dwelling, tending them as a
herder or farmer does his domestic cattle, protecting
them and fleeing with them in predatory raids all this
forms a chapter of natural history so strangely analogous
to the ways of agricultural society as to link the ancient
and wide-spread human interest in ants with the life
of the aphids, which lend themselves so readily to their
remarkable uses. This natural adaptation is, of course, a
factor in the development of the habit which here has
commanded our attention.
That which makes ants particeps criminis in the de-
struction wrought by aphids is the remarkable habit
which they have acquired of deliberately housing the
aphid eggs and raising therefrom the adult forms. From
their breeding-camps and nurseries they transport these
domesticated herds to the plants which they elect to
attack, whether on root or stem. There the aphids
at once thrust in their beaks and begin to draw out
the sap, which the canny " proprietors " appropriate as
honey-dew. It is thus indirectly, and not by any direct
injury inflicted upon plants, that ants become at times
injurious insects. Otherwise they take rank with the
benign insects, by aiding the fertilizing of blossoms by
their frequent visit- for nectar, and the formation and
shifting of tillable soil by digging out their subterranean
Our attention must now pass from the communal
associations which have been established between ants
and alien insects to those existing between ants of sepa-
rate species. These are numerous and greatly varied,
and only a few typical cases can be considered here.
Among the consociated communes of ants there are few
whose relations are more interesting than those existing
between Myrmica brevinodis and Leptothorax emersoni.
The latter is a small species (much smaller than Myrmica),
APHIS HERDS AND ANT ASSOCIATES
and in forming its compound nest it shows a strong pur-
pose to keep its own living-quarters quite distinct from
its associates, although the workers mingle freely with
the Myrmicas in their larger galleries. As compared
with these, the gangways of Leptothorax are small, and
evidently are so made and kept to maintain their isola-
tion and hold their robust neighbors at a distance. It
is only by a deliberate onset of sappers and miners that
the larger ants can make way through their dwarfish
associates' narrow lanes; and this is occasionally done ;
although for the most part the Myrmicas seem content
to let the Leptothorax alone.
A sectional view of the joint underground commune of
the two species shows the parts occupied by Leptothorax
quite apart from Myrmica, but united thereto by their
narrow alleys. The " China town," or foreign quarters
of some of our cities, might be suggested, not inaptly,
as a somewhat analogous communal subdivision on
In spite of this maintenance of independent quarters
for themselves and their offspring, the two species are
truly symbiotic. They intermingle on the most amicable
terms; they run about together in the main galleries
and large myrmican chambers; and they exhibit, under
favorable circumstances, the chief bond of union that
holds this strange compound commune together; it is
the old, old " bread-and-butter" bond that draws to-
gether communities of men. The Leptothorax under
normal conditions obtain their food from the Myrmicas,
and from them alone. [W. 4, p. 14.]
A peep through the glass roof of an artificial nest
shows a Myrmica worker standing stock-still in a gallery
with a Leptothorax mounted upon its back. What does
it mean? The little creature is licking its great host,
who bows gently beneath the osculation as the manip-
ulator passes from back to neck, from neck to head,
from head to face. The recipient plainly enjoys the
operation, and reminds one of the domestic cat purring
under the stroking of a mistress.
Here and there other worker Myrmicas are undergoing
the same treatment, and the queens and the males, too.
In this corner a Myrmica queen, a giantess beside her
dwarfish guests, has four or five attendants, all mounted
in different positions upon their huge host, and working
away eagerly. Indeed, the process seems to be an excit-
ing one to them, as their abdomens are kept in almost
constant stridulatory movement. Doubtless it is agree-
able to both parties; but the act of Leptothorax is not
one of pure benevolence. This is the famous shampoo
dejeune the dinner shampoo of ants that we are seeing
through our trained observer's eyes. There can be no
doubt that the wee operators obtain some substance
from the body surface of their hosts, but what is its
nature, is not easily ascertained.
It has been suggested that it is a secretion from
cutaneous glands, faint but agreeable and edible, and
distributed over the body surface, or that it is a
salivary secretion spread over the Myrmicas by that
mutual licking in which they so often indulge (whose
prime motive would seem to be cleanliness), and of
which, minute as the quantity is, there is enough to
serve the diminutive leptothoracian appetite. This is
probable; for the salivary glands of ants are well devel-
oped, and, as in the case of the honey-bee, may be
good food-stuff, even as used in this indirect way. Here,
it may be, we have an explanation, in part at least, of
2. r >2
APHIS HERDS AND ANT ASSOCIATES
the strange companionships of myrmecophilous insects,
as beetles, crickets, and cockroaches, that become guests
of ant communes.
The Leptothorax have another source of refection, a
good example of which may be induced in an artifi-
cial nest by overfeeding the Myrmicas, which, if given
sugar and water after a fast, are apt to gorge themselves.
In this condition, as they wander about with half-open
jaws, minute drops of sweet liquor will be regurgitated.
These droplets, as they hang upon the maxillae and
lower mouth parts, attract the Leptothorax workers.
They mount the Myrmica's back and imbibe the pendent
droplet, at times sharing the confection with a hungry
myrmican worker ; or the little beggar, from its seat
atop of its host's head, will try the effect of the " dinner
shampoo," usually with the result that the over-full
Myrmica grows complacent and yields to her tiny but
canny solicitor the desired sweet.
It was noted that during all these interchanges be-
tween the bulky hosts and their tiny affinities the former
were continuously complaisant. There were no signs
of irritation or resentment at the officious and uninvited
solicitations of the guests. They were not menaced, nor
seized and held in the mandibles as a mild form of
protest or discipline. The Myrmicas rather seemed, as
their observer thought, to look upon the little creatures
with a gentle benevolence, much as human adults regard
children. The friendly antennal salute was always given
as they passed and repassed their guests. On the other
hand, the Leptothorax attended upon the Myrmicas
with a zeal that seemed almost comical. [W. 1, p. 442.]
Leptothorax is a genus of cosmopolitan distribution, of
heterogeneous instincts, and of catholic temperament,
and therefore well adapted to the varied roles for which
its peculiar size and plastic nature also fit it.
To the necessity for finding food combined with mi-
nute size we probably owe the origin of some remark-
able associations formed by sundry species. Some of
these which live in or near the nests of other species and
prey on their larva? and pupa?, or surreptitiously consume
certain substances in the nests of their hosts, have been
grouped together under the name of "Cleptobiotic" or
thieving ants. [\V. 1, p. 528-9.] As a distinctive name
the title is apt enough, but the lay reader may be ad-
vised that it is not meant to imply moral delinquency,
or that thief-ants are offenders above all others; for
the act of seizing food wherever it is found and can be
taken is common and natural to all ants.
Cleptobiotic ants are small in size and subterranean in
habit, and are persistent intruders upon the communes
of other and larger species. Their minuteness is their
security, and doubtless the source of their peculiar par-
asitic habit; for it enables them to steal into the gal-
leries and rooms of greater neighbors, and plunder their
Fig. 89 SOLENOPSIS FUGAX
a Male, b Dealated female, c Worker. (All magnified)
APHIS HERDS AND ANT ASSOCIATES
flocks of larvae and pupae. The habits of one species
will fairly illustrate the manner of life of thief-ants gen-
erally. Solenopsis fugax (Fig 89) is a European species
Fig. 90 SECTIONAL VIEW OF HOST - ANTS* NEST, SHOWING THE
GALLERIES WHICH THE THIEF - ANTS ENTER
(Wasmann, after Wheeler)
with minute yellow workers, and large black males and
females. The way in which it establishes itself within
the boundaries of a host is shown in Fig. 90 (reproduced
from Wheeler's copy of Father Wasmann's drawing),
where its minute galleries, in the sectional view, are seen
threading the storied commune of Formica pratensis,
and opening into the far broader chambers and tunnelled
roads of the host. Solenopsis is able to maintain an
independent colony and conduct her predatory raids from
a distance, but the greater ease of action doubtless attracts
it near and nearer and even within the bounds of its host.
Stealing along their tenuous subways, scarcely larger
than a lady's knitting-needle, they enter a Formica
nursery, where a bunch of cocoons has been stored in
fancied security. These they mount, perforate, cut the
included pupa? to pieces, and bear the parts away in
their mandibles or absorbed within their crops.
Janet observed their method in an artificial nest of
Solenopsis fugax and Formica rufibarbis. He fed the
former daily about ten cocoons of Lasius queens, placing
them near the formicary gate. Soon the little thief-
ants appeared. From ten to thirty so small they are
would climb upon a single cocoon, which ere long was
dotted with minute perforations that at last united in a
rift that exposed the contents. Then the fierce lilliputians
fell upon their victim, cut into it, sucked its vital juices,
and tore it into minute piecelets, which they bore into
the nest interior.
One wonders how all this thieving and killing can go
on unnoticed and unavenged by the Formicas ? A glance
at the diagram will show that the diminutive avenues
of the aggressors are a secure refuge for them, into which
the Formicas could not follow, even if they were detect-
ed. Moreover, such an experienced naturalist as Doctor
Forel is inclined to believe that, when the two species
chance to meet, the minute size of the thief-ants makes
them invisible to their hosts, so that the burglarizing
and murdering may go on unnoticed. How that could
seriously affect the situation, in view of the antennal
sensitiveness to other distinctions, does not clearly ap-
pear. Besides, small as they are, the thieves are armed
with formidable stings, and are so numerous that they
are antagonists not to be despised.
It must also be remembered that the secret and
APHIS HERDS AND ANT ASSOCIATES
stealthy method of these marauding mites is not cal-
culated to arouse the ire of the Formicas and marshal
them for resistance, as would, for example, the raid of a
battalion of slave-makers. They have the real kleptic
faculty of human robbers, and steal softly to their work.
Having acquired the protective nest-odor of their host,
they doubtless pass in and out, not unchallenged, but with
impunity. In America, Solenopsis fugax is represented by
S. molesta, a minute yellow ant with yellow queens and
dark-brown males. It is widely distributed, and Professor
Wheeler thinks that its habits are substantially the same
as those of its European congener. [W. 1, p. 533.]
It remains to speak of that form of consociation whicji
Wheeler has classified as Plesiobiosis, the "double nests "
of Forel, and which Wasmann has designated as acci-
dental forms of compound nests. This comprises cases
in which two, or rarely more, colonies of ants of different
species occupy galleries and seemingly have established
formicaries in close contact.
Among these ants Professor Wheeler groups several
species observed by this author. The foetid ant (Fore-
lius fcetidus Buckley---? 7 , maccooki Forel) is a small,
yellowish dolichoderine ant which lives amicably with-
in the nest boundaries of the Texan agricultural ant.
Numbers of these ants were seen frequently travelling
in long lines, in single or " Indian" file, across or near the
nests of the agriculturals. Usually their route was upon
blades of grass growing on those nests that were covered
with needle-grass (Aristida) , or along low tufts of grass
on the margin of the disk. The agriculturals took no
notice of their tiny neighbors at least, never interfered
with them- -and the two species seemed to be upon
the most friendly terms with each other. [McC. 3, p. 202.]
Another case of this sort of consociation is that es-
tablished between the occidental ant of Colorado and
a small species of Dorymyrmex (D. pyramicus Roger
var. flavus McCook). There was scarcely a formicary
of Occidentals that did not have upon its surrounding
clearing one or more species. Usually there were two
or three nests, sometimes four, located upon different
parts of the pavement. These were small moundlets of
fine soil, surrounding a central opening that led into an
irregular series of galleries and chambers. [McC. 5 ;
p. loo.] The insects are small, active, irritable, intense-
ly pugnacious, and courageous. The manner in which
these little fellows bullied and badgered their Occident
hosts was amusing and, indeed, amazing. Examples of
this belligerency are given and illustrated in Chapter
X " Warrior Ants and Their Equipment for War."
Of the nests of six species of true ants found parasitic
upon the nest of Occidentalis, I found colonies of the
Sanguine slave-maker on three separate pavements.
The gates were on the clearings not far from the central
mound, and on exploring one Occident nest the formican
galleries and rooms occupied a goodly part of the in-
terior space. The number of ants and slaves in these
compound nests was quite large, judging from those in
sight, yet there was seen no antagonism to these guests
on the part of the Occident hosts. The species were not
greatly unequal in size, but the disparity in numbers
and in belligerent efficiency was such that the Occidents
could have exterminated the Sanguines and their kid-
napped retainers. The auxiliaries of sanguined were
Formica Schaufussi and a small black ant which
Wheeler thinks was one of the Western varieties of
APHIS HERDS AND ANT ASSOCIATES
The most curious and interesting of these parasitic
formicaries was that of a large blue-black Formica. I
first found its galleries at four feet below the surface, and
thereafter traced them to within four inches thereof. The
Formicas occupied the central and eastern part of the ex-
cavation. The architecture was less regular than that
of Occidentalis, which may have been the result of
necessity rather than natural habit, but showed arrange-
ment in stories. The rooms communicated with one
another, and were crowded with workers, callows, and
grubs. The galleries and chambers were placed side
by side a ad in the midst of those of the Occident ant.
One of these, located in the heart of the nest and full of
larvre, was just above and flanked on each side by gran-
aries of Occidentalis packed with seeds. Another open-
ing higher up was surrounded by Occidents' rooms.
I have rarely seen so curious a study as that presented
by these interblended interiors. [McC. 5, p. 154.]
My miner assistant might well raise the query, "Which
of these fellows jumped the other's claim? 7 -the Col-
orado vernacular for, " Who was the intruder, and who
the original possessor ?' : The arrangement and relative
positions of rooms and galleries led me to infer that
there had been a contemporaneous growth. The two
queens established their original cells in vicinity. In
time, mutually expanding their bounds, they approached
each other, and thenceforward held the ground together.
By what peculiar gift or condition were the two species
able to so guide their engineering that they never con-
flicted? Or, did they conflict?
I could find no traces of intercrossing of openings
or impinging of chamber walls. Closely as these ap-
proached, they seemed to be structurally distinct. What
legal conflicts, what local battles and bloodshed, have
resulted from trespasses on boundaries made in the
gold and silver mines of the human neighbors and fellow-
miners of these insects, old-time Coloradoans know too
well. Were the emmets more peaceable and tolerant of
one another than the men? Or, would the secrets of
their subterranean abodes, if given to natural history,
uncover scenes of dreadful conflict and death?
As the excavations uncovered the interior of the
great nest, nothing appeared to indicate a state of war-
fare past or recent. As pick, trowel, and knife exposed
the rooms, both species were surprised in the midst of
their ordinary duties, and showed unmistakably that
they were wholly engrossed in peaceful industries.
But when, by some careless stroke of the tools, rooms or
galleries of the two species were forced together, or
when the crumbling earth precipitated the insects into a
common trench, then the polemic possibilities appeared.
Then blacks and reds grappled in hot strife and fought
with fury. The powerful sting of the Occidents was
brought into service, as the combatants rolled, strug-
gling, in the soil, and the sharp mandibles wrought like a
French guillotine, as witnessed by the decapitated trunks
of the Formicas quivering in the trench, leaving at times
the severed head still clinging to its antagonist by jaws
clasped in the rigor of death.
These battles seemed to confirm the fact indicated
by a study of the architecture, that the status of the
Formicas in this compound nest was one of peaceful
parasitism. The Occidents plainly tolerated their neigh-
bors, for manifestly they had the power, had they been
so inclined, to drive them out or destroy them.
THE FOUNDING OF SLAVE-MAKING ANT COMMUNES
NTS are unique among social insects in the prac-
tice of a form of slavery. Bees and wasps, as far
as known, show no tendency thereto. Indeed, their
physical condition and manner of life seem to bar the
way to the development of such a type of co-operative
citizenship, while, on the other hand, the habit of ants
rather invites it.
In this characteristic we have another suggestion
of those tendencies of human society which appear in
emmet life. As far back as run the authentic records
of our race, we trace some form of slave-holding. The
Abrahamic type, as uncovered in the Old Testament,
was little more than civic adoption, a kind of tribal
11 naturalization ' -to borrow a term from American
customs. Ancient Egypt had a far severer sort, as seen
in her remarkable mural history, preserved even to this
day in the inscriptions and paintings on the inner walls
of her tombs, and confirmed by the Bible story of the
The slavery of classic Rome and Greece, though most
cruel in many of its features, had some mitigations; at
least, it did not close and seal the door of hope, but
kept an open way for its "freedmen" to become honored
and influential citizens. It lacked, as did most early
forms of human bondage, that racial bar and taint which
was one of the worst features of American slavery.
Our British forebears, to whom we owe our views of
both civil liberty and chattel slavery, were at one with
all Europe in holding Africans as the lawful prey of
white men, and quite outside the pale of the common
right of man to liberty and independent life.
One needs this bird's-eye glance at this phase of hu-
man society as he takes up a somewhat analogous feat-
ure of certain ant communes; for our conception of ant
" slavery" is colored by the current meaning of the word
as derived from our own use and wont. It is not, indeed,
an inapt term as applied to emmet communes, if one
regard the usage of men in the whole course of social
history; but it is a different thing as interpreted by
one's preconceptions of slavery as lately existing in the
In point of fact, there is no trace of such slavery in the
relation. What one sees in a so-called slave-holding ant
commune shows no involuntary servitude, nor any con-
ditions substantially different from those obtaining in
ordinary ant communes, except the presence of two
distinct species. These, in their bearing toward each
other, give no signs of superiority or subordination.
It is a co-operative citizenship, whose duties, in one type
of commune, are more sharply differentiated between
the two classes of citizens than in the other, but wherein
all are apparently equal and free, although one class has
been kidnapped in infancy and reared in its abductor's
home. In so far, no further, they may be ranked as
There are several species in America that may be
classed as slave-holding but all may be ranged under two
types the SANGUINE and the POLYERGINE, so named
FOUNDING OF SLAVE-MAKING COMMUNES
from the species Formica sanguined and Polyergus ru-
fescens that respectively represent them. In communes
of the Sanguine type the dominant species preserves
all normal ant characteristics in full potency, and co-
operates with the auxiliary or " subject'' species in
civil responsibility and labor. In the Polyergine type
of colony the dominant species, Polyergus rufescens-
luciduSj the "Shining slave-maker/' has lost all disposi-
tion and even ability to any service but that of conduct-
ing periodic predatory raids and supplying the com-
mune with captives.
It seems an odd coincidence, in view of the preference
for African slaves among men, that the ants most af-
fected by the slave-makers are the dark species and
varieties, particularly Formica fusca and its glossy-black
American variety Formica subsericea. Both the San-
guine and the Shining slave-makers victimize these spe-
cies more freely, perhaps, than any others. This is due,
doubtless, to their greater feebleness and comparative
timidity, as well as their adaptability to associated
We are now to undertake an inquiry into the natural
conditions out of which this interesting phase of emmet
life may have arisen. 1 In so doing, it is well to remem-
ber that our search after the origin of habit must always
be more or less like progress up a blind alley wherein
we are sure to come to a point where a blank wall faces
us. For, follow back our inquiry as far as we may by
observation and experiment, and b}^ reasoning there-
upon, we come at last to the mystery unsolved, and
seemingly unsolvable by our natural methods how
1 For a study of the habit itself, see Nature's Craftsmen, chap. v.
1 8 263
arose the first individual and the first action of the series?
Nevertheless, one must push on, by virtue of his insatiable
thirst after the final cause of things, as far as he may.
The first decided step toward the truth in our study
of the phylogeny of the slave-making habit among ants
was made by Prof. William M. Wheeler. During the
summer of 1904, while studying ants among the Litch-
field Hills of Connecticut, Professor Wheeler made the
brilliant discovery that the female of Formica difficilis-
consodanSj after her marriage-flight, habitually seeks a
weak and probably queenless nest of Formica Schaufussi-
incerta, and thereupon founds a colony of her own species.
The host-commune, the Schaufuss ant, belongs to the
group whose native temperament seems to adapt them
to serve as auxiliaries, and a depauperate and queenless
condition favors the welcoming of a queen, even though
an alien. On the other hand, the Consocians female is of
characteristically diminutive stature, and thus physically
disqualified from the usual role of solitary queens the
rearing of an independent commune. Thus mutually
adapted for union, an alliance is formed, and the first
step of a mixed colony is made.
Now follows a strange and interesting history whose
bearing upon our subject readily appears. The Con-
socians queen drops her eggs. The Incerta workers,
true to their instinct, care for them and rear them to
maturity. Ere long they equal their nurses in number,
and soon exceed them. As there is no natural source
from which to recruit the ranks of the host-species, in
the ordinary course of communal life and service the
original founders gradually decrease, until all have died
out. There remains then Consocians commune pure and
FOUNDING OF SLAVE-MAKING COMMUNES
This method of founding a colony the discoverer called
"temporary social parasitism," and he inferred that a
number of these mixed colonies known to exist, and
which had been thought to be abnormal or accidental
consociations of two species, were in all probability
merely cases of temporary parasitism. And he pre-
dicted that various species of the Formica rufa group
would be found to establish their colonies after the
manner of F. consocians that is, by the aid of some one
of that group most affected as auxiliaries among slave-
It seemed to follow, as an almost necessary conjecture,
that this might give the clew to the true phylogeny of
the slave-making or dulotic habit first discovered by
Pierre Huber. He therefore entered upon a series of
remarkable experiments, from which we may conclude
that the method, as it occurs in nature, has been un-
covered. A strangely interesting story it is. [W. 2,
pp. 33-105.] Twenty-one experiments were made with
young queens of Formica sanguined (of the prevalent
American variety rubicunda) and artificial colonies of
siibsericea, a widely distributed American form of
Formica fusca, which is commonly found as an auxiliary
in slave-holding communes. Two of these were partially
and ten completely successful. The following accounts
of two experiments will show both the professor's meth-
ods and the results.
The artificial nest used was divided into two connected
chambers, one illuminated, the other darkened. Herein
was placed, within the dark chamber, a colony of twelve
large Siibsericea workers and a number of worker co-
coons. To these a female Rubicunda was introduced.
Some of the workers snatched up cocoons and fled
into the light chamber, while others fell upon the stranger
and began to tug at legs and antenme. The queen was
passive for a few minutes, then aroused herself, shook
off her assailants, and began to prance back and forth
in the chamber, pouncing on any worker within reach.
Having slain two of these in quick succession, she began
to collect cocoons and put them in a corner of the nest.
When eighteen had been assembled, she mounted the
pile and stood guard over it, with mandibles wide open
Meanwhile the Subsericeas had hastened with the
remaining cocoons into the light chamber and plugged
up the door with earth-pellets. For two days matters
thus stood, Rubicunda perched upon her looted cocoons,
and the black workers keeping to their own apartment.
At night, however, there must have been a sally and
a combat; for early next morning (July 9th) the queen
was dead, and her captured cocoons were replaced with
the others. The victors' formic -acid batteries had
wrought their subtle aim, for their adversary's large
body was not mutilated. Death resulted from poison.
Let us now mark a more successful experiment. Into
a colony made up of thirty-three Subsericeas, one hun-
dred and fifty cocoons, and a few larvae, a Rubicunda
female was placed. The workers were thereby intensely
excited, and, seizing their cocoons, rushed into the light
chamber. Two who advanced to assault the stranger,
as a sort of "forlorn hope," were shaken off and one
slain. During this conflict other workers stole back into
the dark chamber to secure more cocoons, which were
stowed in the remotest corner of the light chamber.
Meanwhile the queen's excitement had much increased.
In the interval of four hours she had killed five more
FOUNDING OF SLAVE-MAKING COMMUNES
workers. She entered the light room, raided the cocoon
stores, and captured and transferred them, thirty-six
in all, to the dark room. Between trips she stopped
twice to attack and kill workers that ventured near.
Now she retired to the dark chamber, and collected
her booty into a compact pile. This was not done with-
out some opposition, for two Subsericeas slipped by her,
deftly snatched up cocoons from the fringe of the pile,
and carried them back to their own quarters. Their
venture cost them dear, for Rubicunda in the end de-
tected them and slew them ruthlessly. She was highly
excited, and pranced vigorously about the floor that is,
she moved in a jerky way, taking a few steps in one
direction, then wheeling quite around, took a few steps
more, her antennae waving eagerly, and her whole body
seeming to throb with passion.
By the next morning (8 A.M.) only two Subsericeas
remained; but they had managed to regain thirty co-
coons, which the survivors were guarding in a remote
corner of the light chamber, while Rubicunda stood guard
over a bunch of them in the dark chamber. Two and a
half hours thereafter she sallied forth and recaptured all
but six of the workers' cocoons, and added them to her
own store. However, she had not forgotten the scant
remainder, for soon she secured four more, and early in
the afternoon another was captured.
The two workers wandered about forlornly, seemingly
dejected at their hard fate. One came into the dark
room and approached the queen, possibly to test her
willingness to come to terms; but she opened her mandi-
bles threateningly, and the peace ambassador fled. Dur-
ing the night this and one other survivor were killed,
and the last cocoon was added to Rubicunda's collection,
She was now, indeed, monarch of all she surveyed.
She had wholly extinguished the colony of adult Sub-
sericeas brought up under and devoted to the old regime,
and was in a position to rear around her from her kid-
napped pupae and larvae a commune to the manner
born, who would accept her headship and build up a
She seemed to taKe a greater interest in the pupae than
in the larvae naturally, perhaps, because from the former
would come her first effective helpers. And they soon
began to come. That night (July 16th) five callow ants
appeared, and one larva was partly eaten. Had it been
sacrificed to the queen's appetite, or had it died first?
That afternoon she was seen opening a cocoon to release
a mature pupa. She used her fore and middle feet to
hold the stiff silken cocoon-case, while with her mandibles
she tore it open. The youngling aided her by thrusting
out legs and antennae, and was soon drawn through the
hole, to begin the life of an imago novitiate.
Now the work of delivery went briskly on. When-
ever the nest was uncovered, Rubicunda might be seen
either opening a cocoon or removing the pupal envelope
from a new-born callow. A week thereafter the whole
brood of living pupae, one hundred and thirty, had been
set free, the older callows assisting in delivering their
sisters. The queen took the greatest interest in her
black family, and they in turn soon began to care for her.
They fed her and cleansed her, plying their tongues to her
body in the usual shampooing process.
Meanwhile a marked change occurred in her instincts.
Instead of resenting intrusion, and rushing to the de-
fence of her brood when the formicary w r as opened, she
slunk awav and tried to hide among the workers. She
FOUNDING OF SLAVE-MAKING COMMUNES
acted quite like the old queens, who at once fly to the
galleries and lower rooms when a nest is opened or
lighted up. On July 26th, eighteen days after its be-
ginning, this experiment was ended and the ants turned
loose in the garden to shift for themselves.
These two experimental observations which I have
thus presented in abstract fairly typify the results gain-
ed by the patient and ingenious observer. As he remarks,
" the reactions displayed are so definite, uniform, and pur-
poseful, even in artificial nests, that one can hardly doubt
that they are similarly manifested in a state of nature."
If, then, we will permit imagination sufficient play
to suppose our ant queen expatriated and wandering
solitary, we can fairly picture the process by which a
slave-holding commune may be established in natural
site. Through fa oring chance and native instinct she
falls upon a nest of some inquiline species Formica sub-
sericea, let us say. She pushes her way into the room,
vestibule, or hall, hoping, mayhap, for a welcome and an
amicable adoption. The amazed and alarmed inmates
seize their cocoons and larvae and fly before her into the
lower galleries and rooms. To cover their retreat, a few
devoted patriots advance to meet and attack the in-
truder. Her choler rises before this inhospitable re-
ception, and at once her latent war-like and predatory
instincts are aroused, and she flings herself upon her
assailants. Her superior size, strength, and martial spirit
make her a match for many of the unwarlike blacks,
and the home defenders are slain. There follows a pro-
longed quarrel, and the caverns of the Subsericean
commune witness a succession of conflicts and manoeu-
vres for the possession of the infant antlings and their
nurseries and home.
Rubicunda's maternal instincts have now awakened,
and Nature plies them side by side with her martial
powers. She seizes and assembles the cocoons and
larvae of her unwilling hosts, who, in turn, by violence
and stealth, seek to retain and recover them. Back and
forth from the care of the usurper to that of their natural
kin the tender and unconscious things are borne, until
the extraordinary conflict is closed by the conquest of
the Subsericean domain and the death of all its original
owners. Here, in this lowly sphere of life, as so often
it has been in human affairs, violence and usurpation
have prevailed, and a new commune is founded upon
war and robbery.
For this is indeed the foundation of a new emmet
commonwealth. Fierce arid remorseless as Rubicunda
has been to the adult blacks, she is not untender to their
offspring and kin. Her interest in the young brood
deepens with the advent of peace. She feeds the Iarva3 ;
cleanses them, dandles them, shifts the cocoons from
place to place, as though to give them exercise or to
better the location. At last and it is a rare event in
the history of this budding commune the queen-
mother marks the signs of maturity within the tough
silken case that encloses a pupa. With hereditary
gentleness and skill she plies her mandibles as facile
an implement for this delicate surgery as it is dreadful
in fight and delivers from the cocoon-case, Nature's
detached womb, a living imago ant! The delicate
membrane that still encloses it is removed with even a
daintier touch, and there appears the first auxiliary
citizen of the new commune, full grown and full panoplied
for civic duty. It is yet in callow antlinghood ; its shelly
"skin' will harden and darken; but its instinct for the
FOUNDING OF SLAVE-MAKING COMMUNES
service of citizenship is full blown, and at once it joins
its foster-mother and queen in helping into freedom its
Steadily the number of callows grows. Every new-
corner adds to the working force. All are welcomed by
both queen and fellows. To her, all are children; to
them, she is a common parent and sovereign, as loyally
and lovingly recognized as though they had been the
fruit of her own ovaries. At last all the captured brood
have matured, and have joined the working- band, and
the rooms and galleries (the houses and highways) of the
new city are astir with busy life.
Soon Queen Rubicunda begins to function as mother.
The wee white eggs which she drops are cared for by
the black workers and nursed into life. They are young
Ruble undas! Their ruddy skins are in sharp contrast
with the black skins of the Subsericean auxiliaries. It
matters not. There is no distinction. They are citizen?,
all, of one commune; sisters, all, of one family; inheritors,
all, of one nest-odor the real badge of a common citizen-
Meantime Rubicunda has undergone a noteworthy psy-
chical change. The instincts of warrior and sovereign
gradually yield to those of mother and founder. She
screens her own person, since her life is needful to per-
petuate the colony, and for the same reason permits her-
self to be guarded and cared for by the workers. Thus
the commune is founded and the normal activities of an
ant city grow up and go on.
The case here given of the way in which Rubicunda
founds a colony is a typical successful one. But for
every such success there have been a multitude of failures.
And that is well for other tenants of the earth ; for, con-
sideling the vast number of migrants from one nest at
the marriage-flight, were not the losses of life enormous
our world might be transformed into an ant-hill! As it
is, all males perish, and comparatively few r females gain
a foothold upon active communal life.
But we are yet far from accounting for the origin of
that feature in the slave-holding ant's habit which, per-
haps, is the most striking to the ordinary student of
animal behavior viz., the issuing forth in martial bands
Fig. 91--WORKER ANTS DEPORTING CAPTIVES OR THEIR FELLOWS
to sack and despoil neighboring communes of other spe-
cies, and to transport them to their own nest to enter
upon a state of servitude (Fig. 91).
One needs to keep in view the fact that the primary
aim of a Sanguine slave-maker's raid is not to recruit the
tale of laborers, but to supply food. The acquisitive in-
stinct which in seed-eating ants, as Pogonomyrmex and
Pheidole, is expressed in storing grains and oily seeds,
has outlet in Sanguinea-rubicunda and her kind in the
accumulation of the carnivorous food of which all ants
are fond, and which is stored in the compact form of the
immature young of plundered species. Doctor Forel
observed that his Formica sanguined colonies reared but a
small portion of the cocoons given them as a test. One
formicary to which he gave "a fabulous number" of
FOUNDING OF SLAVE-MAKING COMMUNES
Formica pratensis cocoons during the course of a sum-
mer failed to raise a single one. [F. 1, p. 259.]
Yet there is apt to be, and commonly is, a remainder
that gets adopted. And there will always be among
the imported cocoons some that are near maturing, and
actually mature before they are needed for food. These
imported imagines are born into the native nest-odor^
and are thus qualified for acceptable citizenship: a
status acquired, as Miss Fielde has shown, within the
first three days following emergence. [Fd. 2, p. 320.]
These drop naturally into the services of callows, as
though they were at home, and they make up the con-
tingent of consociates in mixed colonies, and of auxilia-
ries or slaves, which in the Sanguine type of commune is
apt to be less than half the whole number of workers.
No sexed forms are tolerated among these abducted and
adopted citizens, and therefore no rival queens with
conflicting claims disturb the communal peace.
These facts are now well established, and they are
substantially those which Darwin predicated as the basis
of his theory of the origin of the dulotic habit in Formica
sanguinea, and which long ago were approved by such
master myrmecologists as Forel and Wheeler, and such
a philosophical naturalist as Lord Avebury.
The acquisition of the habit of raiding in column has
yet to be accounted for. Every individual ant is, by
the primary necessity of feeding itself and others, a
natural forager. The worker is hardly well out of cal-
lowhood ere the strong instinct of communal benefi-
cence, fortified no doubt by personal hunger, impels it
forth from the home gates to pick up whate'vier edible
it may happen upon. The emmet "conscience' 1 knows
no law of meum et luum, and these solitary plunderers
may be seen everywhere afield. A fallen and bruised
apple or peach or a dropped bit of sweet will at once
demonstrate the presence of these universal foraging
scavengers and robbers. Herein nature has planted in
the individual ant the predatory habit upon which to
build up such an expedition in column as the raids of
Sanguinea-rubicunda disclose. This is the first stage;
we proceed to the next.
The wars of ants, as has been shown, usually arise
from the quarrels of a few often, perhaps, stragglers-
over some treasure-trove. One after another joins the
fray; messengers fly to the respective nests; and soon
numbers of recruits, all throbbing with martial fervor,
are thronging from either communal centre to the battle-
field. This tendency to inarch in file and to mass for
defence and attack, and, indeed, for other matters of
common interest, is ingrained with most species, and
seems to strengthen as the colonies grow.
With the honey-ants of the Garden of the Gods, as
the author has shown, such an assemblage occurs before
the evening excursion after honey-dew. [Met 1 , o, p. 24.]
Toward sunset the workers begin to gather around the
single crater-like gate of the home mound. Soon the
summit is covered with the yellow adventurers. At last
the break is made, and away they go, keeping ^vell to-
gether until the column breaks into sections and inte-
gers at the foraging-grounds, a thick clump of scrub-oak
bushes. Quite the same phenomenon attends the even-
ing outbreak of the cutting ants of Texas. [McC. 6,
p. 243; 10, p. 34.] When the chippage used to barricade
the gates has been removed by the smaller workers, the
leaf -cutters push their way out, and pour forth in
squadrons, a great army, and as such march to the chosen
FOUNDING OF SLAVE-MAKING COMMUNES
fo raging-ground. Indeed, something of the same sort
may be seen, though in cruder and less-concerted (if at
all concerted) form, in the daily raids of the mound-
making ants in their excursions after the honey-dew of
aphides upon the trees growing near by their ant cities.
Enough has been said to show that the movements
of ants in column, especially for a hostile and predatory
purpose, is a tendency, not to say a trait, that appears
in many species. To be sure, it is reasoning per sallem-
and doubtless a wide vault, indeed to infer from such
general tendencies the development of a trait so thorough-
ly fixed and admirably ordered as the predatory raids of
our Sanguine slave-makers. But in the present state of
emmet ethology, some such tentative inference may be
justified, until wider and minuter studies shall enable
naturalists to fill up the gaps in our knowledge. One
may have good hopes that this will yet be done, as prying
naturalists go forth to their true aim,
"And take upon 's the mystery of things
As if we were God's spies."
For surely the hidden things of nature must all be
brought to light ere the divine call of man to the uni-
versal inquisition after truth shall be fully answered.
It is significant that while the black auxiliaries are
given a full share of the commune's diverse services,
their red superiors labor with them side by side, and
seemingly with full efficiency and activity. That the
Rubicundas must be sensible of the advantage of strength-
ening their working force may be conceded, and also
that this may have been in some degree, at least, a factor
in determining their kidnapping excursions. But, man-
ifestly, dependence upon their imported labor has in
nowise reacted unfavorably upon themselves. They have
not deteriorated, but retain their full generic equipment
as builders, nurses, foragers, and workers generally in
all the diversified duties of ant citizenship.
This is in marked contrast with what has occurred in
the Polyergine type of slave-makers. There the workers
have lost all characteristic qualities except the martial.
They have developed into mere vital kidnapping ma-
chines, with those soldierly capacities needed to make
them effective. As slave-catchers, robbers, and fighters
they are highly efficient: but they lack the power to
carry on the ordinary and needful affairs of a commune.
They remind one of those human tribes whose males
function as warriors only, and leave to their slaves and
women the entire work and burden of the commune.
Indeed, with the Polyergines, degeneration has gone so
far that they depend upon their slaves not only to pro-
cure food, but to bestow it. So abject is their estate
that they cannot feed themselves, and, lacking the of-
fices of their slaves, die of starvation. However, as
with the Sanguines, no sexed forms are permitted by the
Polyergines other than of their own species.
It is not the writer's purpose to give here a detailed
account of a slave-maker's raid. He has given that
elsewhere. [McC. 7, p. 71.] But it may be ranked justly
among the most interesting incidents in the history
of insects whose ethology presents some remarkable
analogies to our own social manners. Were we to take
a brief view of such an event, what points would fix
our attention? We would note the organization or com-
munal action implied in the impulse that sends the
raiders forth; the scouting that must precede a sortie in
order to locate the quested objects of assault; the com-
1'OUNDING OF SLAVE-MAKING COMMUNES
munication of antennal signals; the drafting of the red
warriors from whom the attacking contingent is drawn,
and the gathering of the black auxiliaries to tarre on the
belligerents, though in sooth they need no such "very
pregnant and potential spurs." We would note the
forward movement; the ordered march; the vanguard
action with skirmishers from the assailed commune; the
fierce scaling of the Subserieean barricades, and the
plunge into the cleared ways. We would see, perhaps
pity, the futile efforts of the besieged to enguard their
commune gates; the flight of the inmates, bearing their
young, from the pillaged nest; the woe-begone groups of
refugees hiding in the vicinage; the little knots of com-
batants scattered here and there around the field, the
melancholy tailings of a lost battle; the maimed, the
dying, the dead scattered here and there. We would
follow the return column of raiders laden with their
booty of larvae and pupa*, and occasionally adult blacks,
as tender-hefted in this office as they had been ruthless
in assail; the heartening of the pillaged Subsericeans
as they see their foes retiring ; the occasional rallies and
rear-guard attacks to recover some of the spoil, and not
always in vain. We might feel, perhaps, a flush of in-
dignation at the welcome of the well-guerdoned spoilers
to their home commune, with every token of satisfac-
tion (except noise!); and, on the other hand, a touch of
sympathy at the gradual return of the refugees to their
desolate city, with the young saved from the common
spoilage, to take up again the role of communal life.
All these incidents unite to form an event unique and of
A faithful description thereof, were it published with
the bare substitution of human names, would need scant
revision to serve as an accurate account of a scene in
the warfare and predatory expeditions of men. One
who has viewed both events the storming of an ant-
hill and the assault upon a fortified town will vouch
for the striking resemblances that appear throughout
the entire series.
Perhaps those whose sympathies have been keenly
enlisted by the author's recital of the cruelties and suf-
ferings attending the wars of slave-making ants will not
deny the plea for peace, universal peace, as the ultimate
end of civilized men; and meanwhile the mitigation, in
every attainable way and measure, of the awful rigors
v */ C_5
of war as now tolerated in a world still so largely in
a state o^ nature, unhallowed by Christian grace and
PROBLEMS OF SANITATION AND PERSONAL
THE sanitation of cities and homes has come to be
one of the most important problems of human gov-
ernment. It is only recently that it has been con-
sidered with scientific
method and thorough-
ness, and that society
has addressed itself to
its solution with ade-
quate vigor. How is it in ant commues?
Their method of preserving the public
health is summed up in one word clean-
Our treatment of the subject falls nat-
urally under the heads of personal clean-
liness, parental cleanliness, and public
cleanliness. Personal cleanliness with
every ant is a passion.
As much time is given Fig. 92 A THIRD LEG OF ANT,
SHOWING HAIRS AND SPINES
to cleansing the person USED IN PERSONAL CLEANING
as is required, and no
work is so urgent as to interfere with that. Nature has
abundantly provided for the support of the habit which
she has implanted, by the gift of certain implements.
The legs (Fig. 92), which are clothed with hairs, bristles,
J 9 279
and spines that are likely to take up dust, are scraped
against one another to remove the coarser grains of dust,
as a man might scratch a shin-bone with the calf of an
opposite leg. Then they are drawn alternately through
the jaws, which, with
the saliva of the mouth,
act as a sort of scrap-
er and sponge for re-
moving finer particles.
Again, they are used
to comb the head and
antennae, for which
they are provided with
pairs of tibial combs,
coarse-toothed and fine-
toothed, of which our
own toilet articles are
a close likeness. 1
In addition to these
is the tongue (Fig. 93),
a rasped organ similar
to that of dogs and cats.
How effective this is
a View of out-thrust tongue of agri- f or cleanliness in these
cultural ant from above, b.c kn-
larsed views of the "bosses or bulbs" domestic animals most
on the tongue, se Serrate edijes.
md - - Mandible. mx - - Maxilla.
sc Scope of antenna?
persons know. It is
with ants. It is this
organ that is used in those parental acts of cleansing
committed to the nurses. The larva?, from the time
they are taken in charge until they pass into the pupa
1 Nature's Craftsmen, p. 67.
SANITATION- -PERSONAL BENEVOLENCE
stage, are so freely sponged with the rasped and moist
tongues of their caretakers that there is little chance
that dirt or parasite or fungoid germs shall remain.
Even after adult life is achieved the friendly offices of
cleansing are exchanged between neighbors, and one
will see a mutual shampooing among the ants in his
The need for personal cleanliness is greatly increased
by the underground life of ants, which subjects them to
attacks of sundry vegetable moulds and parasitic insects.
Some of my experimental colonies have been destroyed
by mites (Fig 94); and it was pitiful to see the little
creatures' struggles to protect themselves from the in-
vasion of the hordes of minute parasites, against whose
attacks they were seriously, even fatally, hampered by
the artificial conditions of their unnatural life. The value
of special armature of legs and jaws and tongue, and the
habits of ceaseless cleanliness engendered by their use,
were mightily emphasized by one's observation of this
unfortunate episode in the career of these imprisoned
colonies. The thought occurred that the habit of feed-
ing upon fungus growths, and the cultivation of fungus
gardens in the AttidaB (cutting ants), may have arisen
from the use of the tongue and jaws in freeing them-
selves and their commune from the attacks of vegetable
The location of the larvse is often changed, a useful
sanitary precaution. The baneful effects of sudden
changes in temperature and humidity are met by shift-
ing the antlings nearer the surface or farther within
the cone. For such manipulation among the mound-
builders their elevated and perforated structures are
well adapted, and for this, in part, may have been de-
veloped. In small nests one may observe this by turn-
ing over a flat stone on a bright spring day or in early
autumn, when the little heaps of white Iarva3 may be
Fig. 94 MITES THAT ATTACK ARTIFICIAL NEST OF HONEY-ANTS
a Dorsal view, b Ventral view, c Suckers on the same in
different degrees of extension, d Mites upon the cheek of a
seen lying in the top galleries next the stone, whither
they have been brought from the underground rooms
for the sake of greater warmth and health.
Ants enjoy these personal ablutions, as one readily
sees who closely observes them either in nature or in
artificial nests. While engaged therein they put them-
SANITATION PERSONAL BENEVOLENCE
selves into sundry odd positions ; at least, so they seem
to an onlooker, although quite similar attitudes may be
seen in cats and dogs when giving themselves a tongue-
brushing. In mature imagohood they are seen fre-
quently sponging and combing themselves. Dust and
impurities of whatever kind they cannot abide, and are
uncomfortable until rid of the defilement. They are
continually in contact with muck and mud and dust,
Fig. 95 ANTS CLEANING STINGING APPARATUS AND BRUSHING
BACK-HAIRS OF THE HEAD
living as they do on and under the ground in earthen
caverns and cells. Yet who ever saw one looking untidy
and unkempt? (Fig. 95.)
"As tidy as an emmet" would be an apt proverb, and
it would apply with equal truth to bees, wasps, hornets,
yellow- jackets, and other insects. If it be true that
cleanliness is next to godliness, our tidy emmets, not
here and there a rare example, but one and all, would
be fair candidates for canonization, and no advocatus
diaboli could challenge their record successfully. Per-
haps it would be impossible for human laborers, in any
conceivable industrial condition, under any form of
government, to approach even afar off the habitual
cleanliness of working ants.
But one who has mingled much with working-men and
closely observed their manners might venture to sug-
gest that a much closer approach to these exemplary
characters is entirely practicable. That it would greatly
enhance comfort, health, good looks, and that sense of
respectability and personal purity that goes so far to
elevate human nature is hardly to be doubted. But
could employers afford to give their workmen the time
needful to effect such personal cleanliness? Would the
increased efficiency coming with the higher quality of
manhood and womanhood thus attained sufficiently
increase the product and the value of the work to justify
the sacrifice? One asks such questions glibly enough,
but how shall he find a practical answer?
An important item in public sanitation is ventilation.
Since ants are apterous, no such mode of agitating the
air and producing a current by rapid wing movements
is possible as practised by bees and hornets. The former
have squads of winged ventilators just within the hive
gate, the latter just outside the door. How ants pro-
duce a like effect is not yet determined. As most of
them domicile in the ground, they are not as likely to
suffer from heat as honey-bees and hornets. But al-
though their consumption of air is not great, one would
suppose that such crowds of creatures living in such
confined quarters as we have described would soon viti-
ate the atmosphere and make necessary some sort of
Perhaps this is secured among the mound-making
ants by placing the city gates most numerously at the
base of the cone. Through these there is doubtless a
constant or sufficient current of air passing out of the
open doors distributed along the sides and upper parts
SANITATION PERSONAL BENEVOLENCE
of the mound. Moreover, there is an extensive system
of underground galleries reaching, in one case at least,
sixty feet from the central mound, and these are prob-
ably ventilated through the basal gates. But it is not
so easy to see how such vast structures as the nests of
the agricultural and Occident ants can be ventilated
through their single gates.
The problem of drainage must be a pressing one in
ant communes, and its practical solution is an interest-
ing study, although one must depend more upon reason-
able inference than deduction from known facts. That
many nests must be inundated during long and hard
rains is inevitable. That their inmates can endure
a goodly period of submerging without drowning is
known. But some method of warding off or carrying
off or absorbing the excess of water in severe rains and
floods seems to be required.
The conical shape of such nests as are built by mound-
making, Occident, and rufous ants, although perhaps pri-
marily due simply to the natural action of gravitation,
must aid in keeping the inmates dry by shedding the
rains as do our own peaked house-roofs. One would
think that placing the bulk of the gates near the base of
the mound would be disadvantageous until he remem-
bers that the large space above, with its numerous series
of interlacing galleries and rooms, gives an admirable
refuge for the commune's infant charges. There they
may be deported in heavy rains and kept in good con-
In the case of such single-gated cones as those of the
Occident ants, the danger of flooding is less; and, more-
over, there, as with the flat disks of the agricultural ant,
gate-closing can be resorted to. At least one example
of such a mode was observed before a storm, the gate
being shut up by pebbles, earth-pellets, and chippage
precisely as at night. One agricultural ant-disk was
watched during a rain. The harvesting workers rushed
for the central gate from all points of the circle, over
which the water was beginning to gather, and in a
moment the gateway was choked up by the crowd of
insects massed on and around it. I did not think at the
time this was done with the intention of closing the gate
and shutting out the water accumulated in the plaza,
but it had that effect. Possibly it may have been in-
It is probable that in such weather conditions the rain
that enters the nest gradually descends through the
storied rooms and galleries and is partly absorbed dur-
ing descent, and at the bottom of the nest is gradually
taken up by the underlying ground. It may even be
that the lowest cavities, both chambers and galleries, are
left uninhabited to receive excess of intrant rains, or
are vacated during wet weather, that they may serve as
a sort of temporary relief reservoirs.
Livingstone [Lv. 1, p. 353] notes that the ants of Dilolo
(South Africa) manage to preserve their communes upon
plains where water stands so long annually as to allow
the lotus and other aqueous plants to mature. When
all the ant horizon is submerged a foot deep they occupy
little houses built on stalks of grass and placed above
the line of inundation. Livingstone argues that this
must have been the result of experience, since, had the
insects waited until the inundation had invaded their
subterranean quarters, the required soil for fashioning
their elevated nests could not have been obtained.
Some of these raised rooms were the size of a bean, others
SANITATION- -PERSONAL BENEVOLENCE
as large as a man's thumb. Could the great missionary
explorer have fallen upon some species of Cremastogaster,
who thus utilized their tent-building habit?
It is certain that ordinary showers do not stay, but
rather quicken, the commune's activity. One often
sees them followed almost immediately by a vigorous
rush of workers from the gates bearing earth-pellets.
Part of this dumpage may have been the inwash of dirt,
but most of it was evidently new earth, which, moistened
and softened, made easier digging, a fact which these
opportunists at once saw and utilized.
Within the numerous galleries and rooms, all under
cover and in darkness, and thronged by myriads of in-
sects in continual movement, together with eggs, larvae,
and cocoons, there is of necessity much litter of various
sorts. This is regularly removed by the workers, who
may be seen carrying it forth and dumping it at points
outside the walls. The inter urban highways, as uncov-
ered by the writer, were never found obstructed by rub-
bish or fouled by filth. As compared with the streets
of many of our own cities and towns, they were models
The mound-making ants, while keeping at times to
certain fixed trails, do not lay out permanent roads by
which to communicate with surrounding fields. But
with such species as do thus unite their large communi-
ties with near environments, the roads are kept in ex-
cellent condition. For example, the disk or plaza or
pavement of the agricultural ants of Texas is kept during
summer hard and smooth, usually denuded of grass and
weeds, and well policed of extraneous matter. The sev-
eral roads that radiate into the harvest-fields of ant-rice
and other seeds are also kept scrupulously clean.
It has been thought that the hospitality extended by
ants to beetles and other myrmecophilous insects is in
part due to their value as general scavengers just as men
in a ruder stage of civilization have kept (and still keep)
dogs, and as certain communities protect by law turkey-
buzzards and gulls. The relations of these alien associates
to the food supply have heretofore been pointed out,
but their usefulness in removing communal garbage may
be a factor in maintaining these strange companion-
Perhaps one might venture to suggest tnat the steady,
ample, and congenial occupation of ants must contribute
to good health, the chief aim of sanitation. And there-
in may be included an evenly poised temperament, free-
dom from anxiety for present and future support, and
absence of the strain and wear of nerves which come
with troubles over property and the want thereof all
of which are characteristic of ants, and which may be
counted for these and other lower orders a boon of
This estate, with its sequent of perfect content and
happiness, ants attain by complete absorption into the
commune, the entire atrophy of the sense of personal
ambition and possessions. Men seek the same in a way
precisely the reverse: by the enlargement of personal
importance, and the acquirement and accumulation of
personal property. Could the experiment be fairly
made in a human commune, it might be found that men
could more readily and completely secure the repose of
happy minds within wholesome bodies by the methods
of the ants than by their own. In all ages men have
sought this repose in organized communes, religious and
secular; but these have been marked, for the most
SANITATION- -PERSONAL BENEVOLENCE
part, by the elimination of that element which is the
central vital impulse in the life of an ant commune the
production and nurture of the young. When the larvae
and pupae are removed from ants they soon degenerate
and decay. Is it different in human communes? Under
natural law, can it ever be so ?
In the various phases of sanitation considered we
observe that ants attack the problem by precisely the
same method of communal labor that prevails in other
departments of public service. Just as every citizen
is a warrior without a board of war, and a policeman
without a police department, and a worker without a
board of public works, so without a board of health every
citizen of these emmet republics is a sanitarian. No
street commissioners are needed to purge the public
highways, for every citizen feels in herself the responsi-
bility of a street commissioner. Literally, every ant
looks out for her own premises; and not only so, but for
her neighbor's premises as well. In fact, there is no
distinction in this regard, for every part of every street
is held to be equally the charge of every ant. What a
paradise of wholesome purity our city streets might be
if citizens would take such an attitude, or if they would
go as far as unanimously to abstain from causing litter
and scattering filth! We shall never reach the ideal
City of Health until the individual conscience has been
educated and elevated to the duty of entire cleanliness
within the dwelling and around it, and upon the public
streets and parks, as personally due to community.
It is perhaps due to their overmastering patriotism
that one fails to discover individual benevolence in ants.
Friendships and personal affection in the limited and
specialized sense familiar among domestic animals are
A XT COMMUNITIES
unknown. There are, indeed, actions that at first give
the impression of such sensations, and it is not easy at
times to mark the difference between the product of
communal instinct and the promptings of real individual
kindness. But the evidence, on the whole, seems to be
against the existence of any sentiment separate from
that Spirit of the Sodality which, with and under all
serisori-motor reflexes, sweeps on the mass and the
individual alike, with whatever variations, diversions,
and seeming contradictions, to the one sovereign end-
the perpetuation of the commune.
A few examples from my studies of the honey-ants
will illustrate this view. It was seen in exploring the
nests in natural sites that the workers showed great in-
terest in the preservation of the rotunds, or honey-bear-
ers. As the honey-rooms were opened, and the rotunds
disturbed from their roosts, the workers of all castes
rushed eagerly to them and dragged them into the un-
broken interior (Fig. 96). Sometimes several would join
Fig. 96 HONEY-ANT WORKERS IN ACTION
a Honey-ant dragging a rotund up a wall, b Workers
moving a rotund, or honey-bearer
SANITATION- -PERSONAL BENEVOLENCE
in removing one rotund, pushing and pulling her along.
A sketch was fortunately caught of a worker-major
dragging a honey-bearer up the perpendicular face of a
cutting made while excavating a nest. The mandibles
of the two insects were interlocked, and the worker
backed up the steep, successfully drawing her protegee.
This interest in these unwieldy rotunds is maintained
in the daily life of the formicary. They are regarded as
dependents, like the queen, the virgin females, and males,
and are fed and tended as such. Here is at least the
semblance of beneficence, but it is doubtful if such action
passes beyond the control of communal instinct; for,
on the other hand, a number of examples fell under
notice which caused doubt as to the existence of a per-
sonal sentiment toward special cases of need, outside
the routine limits of communal service. I observed
several of these.
After the soil was duly prepared in my artificial nests,
the ants were introduced and left to work out their
own habitations. The honey-bearers were thus mingled
upon the surface with the workers, upon whom fell the
entire task of home-making. The latter at once began
digging galleries and rooms. In bringing up and dis-
tributing the pellets, there was much opportunity to
show carefulness and tenderness toward the honey-
bearers scattered over the surface. No such action was
noted, although I was anxious to discover such excel-
lences in my little charges. On the contrary, there were
exhibitions of what seemed cruel neglect and even posi-
tive cruelty. For example, the grains of soil, instead of
being dumped on unoccupied spots, were heaped around
the rotunds until the poor creatures were literally buried
alive. It would have been easy for the carriers to draw
their fellows aside or to go around them ; but this did not
occur to them, or the disposition to such service was
Again, as the galleries were made and honey-rooms
gradually took shape, most of the rotunds managed to
roll into them and secure a place therein. They pain-
full} 7 attained their perches unaided by the workers.
Some of them on their route got fastened in the gang-
way in most uncomfortable positions, with heads down-
ward and bodies variously awry. The workers passed
by and over them continuously for many days without
the least apparent concern, and without one observed
effort to relieve their comrades, who could have been
righted readily and drawn into the chambers.
Once more, the rotunds often dropped or were shaken
from their perch against the roof to the floor. They
remained just as they fell, except when they were able
to clasp some near-by clod, or bit of gravel, or surface of
a wall. In such case they recovered their perch or
put themselves in a comparatively comfortable position.
The greater number, however, lit upon the round ab-
domen in such wise that the body was erect and the
legs were thrust out unsupported. These unfortunates
were faithfully attended, often cleansed and caressed,
after the prevalent hereditary manner toward depen-
dents, but in no single instance did the workers attempt
to right them and restore them to the roof. Yet they
were well able to do so, and the fallen rotunds were in
sore need of help. Some of them lived more than two
months in this awkward position (Fig. 97).
But evidently thev were most uncomfortable, for the
few who were within my reach eagerly accepted aid.
The offered stick or quill was clasped so firmly by the
SANITATION PERSONAL BENEVOLENCE
mandibles, sometimes aided by the feet, as to enable
me to transfer the bulky creatures to any point near by,
and even to lift them out of the nest. Here, again, the
idea, or at least the act, of helpfulness was lacking. If
Fig. 97 HONEY- ANTS AND ROTUNDS
1. Worker honey-ant feeding a rotund. 2. A rotund honey-ant
partly buried by mining workers. 3. A rotund in trouble a
" priest " and a Levite near by.
we suppose the power to communicate their distress and
desires to have been possessed by the honey-bearers, we
must think the workers even yet more lacking in feeling
and intelligence. In any case the power of benevolent
initiative seemed wholly wanting.
One more case. A honey- bearer was partly buried
under her perch, that portion of the roof having fallen.
Her abdomen was quite covered by the sandy particles
at the margin of the little landslide. A rescue would
have been easy, but it was not undertaken. A sketch
made shortly after the accident shows a worker-minor
standing before the rotund with head and body erect,
antenna? attent, with every mark of curious interest in
her pose. She watched the struggles, and (as it seemed
to me) the mute appeals of her unhappy fellow, who by
great exertion had heaved up the clod somewhat, and
then ' ' passed by on the other side." Meanwhile a second
worker was perched atop of the clod, coolly and cozily
combing her back hair and antenna? ! This tableau fitly
characterizes the workers' behavior in such cases.
Such facts, which might be multiplied, incline one
to the view that personal benevolence, as distinguished
from tribal or communal benevolence, does not exist
in ants. I cannot even assent to Lord Avebury's sug-
gestion [Lb. 2, p. 497] that there are " individual dif-
ferences' 3 among them, and that, as with men, there
are priests and Levites as well as good Samaritans. The
apparent cases of beneficence, outside the instinctive
actions that lie within the line of formicary routine, are
so rare and so doubtful as to their cause that, however
loath, I must decide against the existence in honey-ants
of even so much personal benevolence as would make an
emmetonian "good Samaritan."
It is true that many of the above observations were
made upon insects living in artificial conditions, but they
cannot be far away from their habits in a natural site.
Moreover, it is possible, not to say probable, that there
may be species that show greater aptitude for acts of
personal kindness, just as some have special aptitude for
war, while others are deficient in courage. But all the
facts within the author's knowledge confirm the above
conclusion. The beautiful acts of unselfish devotion
to dependents, especially the helpless young, the self-
denial, the vicarious self-sacrifice, the utter elimination
of self so often seen all these spring solely from the
relations of ants to the commune. They are the acts
of the ant-citizen, in whom all the functions of life are
centred and absorbed, not of the ant - individual, an
independent living unit.
Patriotism, communal loyalty, communism, or by
whatever name we may designate a sentiment or im-
pulse for which, in the poverty of our speech and the
limits of our sympathies with a sphere of life so far out-
side of our own, we have no exact word, is the one passion
that moves the typical ant in its varied acts. Personal
sentiment there is next to none. Outside the routine
of communal duty and service, the actors in an ant com-
mune know no law but the instinct to serve and preserve
the commonwealth. Even in satisfying the demands of
appetite the individual is lost (with the workers, at least)
in the public interests, for even the contents of one's
crop are held at the service of the community.
Of course, there is reason for this condition. An im-
perious law of co-operation in the mass does not favor
the development of strong individual characteristics.
The mechanical order that assures the sodality and safety
of society among insects would be disturbed by strong
personal ties, which would thus introduce a divisive and
enfeebling force, especially in times of communal peril
when all sentiment and service should be concentrated
upon public interests.
A NOTE IN REVIEW
WHILE emphasizing, to the utmost that known facts
can justify, the marks of superior instincts in ants,
and while believing that their social organization has
developed powers and incited to behavior that suggest
human conduct in certain communal conditions, the
author must not be understood as in any degree con-
founding emmet instinct with human intellect. There
is an impassable gulf between them.
To reflect upon a rule of conduct; to decide thereupon
and frame one's behavior accordingly; to assemble facts
gleaned from the past and from the present; to reason
upon and deduce therefrom principles of duty and ser-
vice, of social and individual government, of personal
responsibility and worship and immortality all this,
which is impossible to the ant, but is within the ordinary
powers of man, set him in a class by himself, so far apart
from all other animals that, in contrasting or comparing
the one with the other, and in tracing resemblances be-
tween their physical and social actions, we shall fall into
serious error if we forget this fundamental difference.
When, therefore, we use the word "intelligent" to ex-
press a quality of animal behavior, as it seems necessary
to do in the deficiency of our language, we shall escape
confusion in our philosophy if we remember that " in-
telligent" and " intelligence" as so applied are terms of
A NOTE IN REVIEW
convenience and not of accuracy. Under such limita-
tions, a review of these pages compels the conclusion
that among all the lower animals ants must be ranked
for intelligence as most nearly resembling man in the
quality, variety, and complexity of their achievements.
Can we suggest any natural cause for this?
The development of civilization has come largely
through the interaction of life upon life. And this has
had its chief effect in cities and towns, where human
beings more closely and continuously affect one another.
A trace of that fact remains in the current use among us
of the word "pagan." Christian civilization had its first
and widest acceptance among the cities, while the rural
sections held to the old religious forms. Hence the
Roman name for countryman, rustic (paganus), came
to be the equivalent of non-Christian or heathen.
It is doubtless due to the same underlying fact or force
in nature that ants have acquired traits that place them
among the most intelligent members of the insect world.
The vigorous reflex of life and habits which existence in a
commune compels, widens the horizon of activities and
develops and nurtures facilities and conduct that strong-
ly mark the possessors as superior. To ants as well as
to men, the commune is a school. It would seem to
result that the more general the participation in com-
munal activities, the higher and more widely distributed
will be the advancement.
The idea fancy or fact, as one may choose that the
communes of ants and other hymenopterous insects are
organized after the fashion of human government is
not a novelty. Long ago, Shakespeare, whose high en-
dowments of intellect and sympathies brought him with
almost unerring course to the very arcanum of nature's
mysteries, conceived of the bee commune as a well-
ordered kingdom, whose "divers functions "' are bound
together by "obedience 1 ." The passage is well worth
quoting for its bearing upon the matter before us :
"So work the honey-bees,
Creatures that, by a rule in nature, teach
That act of order to a peopled kingdom.
They have a king and officers of sorts;
Where some, like magistrates, correct at home;
Others, like merchants, venture trade abroad;
Others, like soldiers, armed in their stings,
Make boot upon the summer's velvet buds;
Which pillage, they, with merry march, bring home
To the tent-royal of their emperor,
Who, busied in his majesty, surveys
The singing masons building roofs of gold;
The civil citizens kneading up the honey;
The poor mechanic porters crowding in
Their heavy burdens at his narrow gate;
The sad-ey'd justice, with his surly hum,
Delivering o'er to executors pale
The lazy, yawning drone. I this infer,
That many things, having full reference
To one consent, may work contrariously."
King Henry F., Act I., Sc. 2, 1. 187 sqq.
Making due allowance for poetic license, and taken
in its broad intent and general sense, this passage ex-
presses fairly well the varied duties under one common
discipline that occupy the energies of an ant commune.
The reader can analyze it and follow out the analo-
gies for himself. Perhaps he may be able to point the
needle of his thoughts to that "rule in nature' 1 which
brings about and regulates this "act of order." If so,
the author will confess that he has compassed a mystery
that thus far has eluded his own research.
But for the "act" itself we may venture, without
A NOTE IN REVIEW
justifying the charge of anthropomorphism, to borrow
a word from human civics. Plato distinguished four
kinds of character-making " enthusiasm ' among men
-prophecy, prayer, poetry, and love. These give to
human faculties the stimuli which urge man to achieve
his highest destiny. We might enlarge the list by at
least the element of patriotism, and thus mark another
likeness between man and social insects. For patriotism
-that is, a supreme devotion to the commonwealth-
is the prevailing " enthusiasm ;; among ants, bees, and
wasps. With them one finds no discount laid upon
the communal workers and military guards. All, except
needful dependents, are laborers and warriors. All are
alike devoted to the civil welfare and defence.
Many times and in many ways the devotion of ants to
their commune has been tested. The rule is well-nigh
invariable of instant and absolute self-abnegation, and
surrender of personal ease and appetite, life and limb,
to the public welfare. The posting of sentinels at gate-
ways is customary, and they are apt to know first the
approach of danger. With heads and quivering anten-
na^ protruded from the opening, these city watchmen
not only dispatch within all news of threatening peril,
but rush out with utter abandon to face the foe. With
ants patriotism is not " second nature"; it is instinctive,
inborn, seemingly as strong in the callow antling as in
the veteran brave. It has no second place. It is first,
It must be confessed, however, that it is rigidly ex-
clusive. Racial catholicity is not an emmetorian virtue.
Ants are without that elastic hospitality which embraces
and assimilates all species of their family. Even the
slave-makers hold their domestic auxiliaries strictly
distinct in the radical function of propagating and thus
perpetuating the species.
If one should seek some directing or governing per-
sonal power of that system by which the complex and
varied results which these studies note are wrought out
with almost unbroken unison among hundreds of thou-
sands, even millions, of individuals, he would find his
research thwarted at every point. There is a smack of
royal authority in the title " queen 7 given the ant in
whom is concentrated the function of maternity for
renewing the life of the whole community. But in this
sense the word is figurative, for queenhood means sim-
ply motherhood. Beyond the brief initial period of a
formicary, when the solitary female in her circular cell
has unlimited power over her scant domain, and in her
single person represents the norm of all reproductive,
nutritive, constructive and administrative forces, an ant
queen has not a shadow of that function of rulership which
the words "queen" and "king" express in human relations.
From the moment that she has raised around her a
band of workers strong enough to take up the labors of
the growing society, she becomes merely an immense
vital organ for producing eggs. She does not even care
for these, as she did with the first few laid; for they are
whisked away by attendant workers the instant they
leave her body, and thenceforth are the common charge
of the colony. A girdling corps of courtiers attends
her day and night, whose members care for her personal
wants, restrain her movements within safe limits, and
collect and distribute to the nurses the precious atoms
of life upon which her importance depends. But no
breathing of rulership or authority or governing in-
fluence of any sort issues directly from her.
A NOTE IN REVIEW
Where, then, is the headship of the State? Is it vest-
ed in the courtiers nearest the queen's person? Are
they a sort of cabinet or board of governors, among whom
the several functions of governing are distributed? All
observations show that their chief office is to secure to
the community the eggs upon which its future exist-
Is government lodged with the sentinels, who seem to
represent the fighting or defensive elements of society?
There is no war-lord. There is no commander-in-chief.
There is no standing army and no soldier class in the
large majority of species, although in " necessity's sharp
pinch'' every citizen becomes a soldier. Certainly our
mound-making ants, at least, are not a military govern-
ment, though every guardian of the gates is at once
and equally the embodiment of military authority, and
bears in its own person every grade from general to
Perhaps the nurses, who include in their nursery ser-
vice the duties of hospital corps and medical staff, are
the fountains of governing power? No! Among men
it may be true that "the hand that rocks the cradle
rules the world"; but nursing ants in their offices
keep themselves to their own sphere, and cannot be
said even figuratively to sway the policies of the ant-
Shall we, then, fall back upon the great body of build-
ers and foragers and laborers generally ? Are they organ-
ized into "unions," with chapels, delegates, brotherhoods,
chiefs, boards, circles, and assemblies, who have grad-
ually developed an influence that controls the common-
wealth? On the contrary, perhaps the most perfect-
known example of absolute "individualism' 1 ' among so-
cial creatures is a typical ant society. Therein every
ant is, as nearly as is conceivable, a law unto itself.
Here is neither congress nor legislator. Every individual
seems to follow the desires and devices of her own heart.
But here also is such perfect obedience to "law" as only
insect commonwealths show among all known organized
societies. Where is the source and where the potent
centre and motor of this law?
The mysteiy deepens as one's inquiry widens. Here
is effective government; but where are the governors?
"Those that think' -sang Goldsmith "must govern
those that toil." Here are toilers enow; but of that
higher, or supposed higher, class who think out the
problems of State and the means for their solution
the most careful searching finds no trace. The varied
and complicated facts of governments, as men know
them, here meet the observing naturalist and the
speculating philosopher. But neither naturalist nor
philosopher has yet put his probe upon the secret
source of government among ants and other social
Can there be government without visible governors?
Here in our ant city one sees home-making, home-guard-
ing, home-nurturing; the building of roads, store-rooms,
nurseries, vast structures that relatively rival those of
the great centres of human population. Here one sees
the gathering of supplies; the storing of crops; the
waging of war; the utilizing of captives from alien tribes
for co-operative service; the keeping of domestic herds;
the policing and sanitation of habitations and streets;
the nurture and care of the young. All the practical
results of organized society one sees in the emmet State.
Is not this government?
A NOTE IN REVIEW
" For government, though high and low and lower,
Put into parts doth keep in one consent,
Congreeing in a full and natural close
But where is the master musician and his orchestral
leaders? Whose hands arrange the parts, direct the
performers, give unity to the whole, and from their
wondrous "congreeing" of action bring forth this per-
fect piece of social harmony? The Spirit of the Com-
mune! do you say? It is a phrase to drape our igno-
rance. The reasonable answer still evades the student's
grasp. Though the natural philosopher may justly
claim that "in nature's infinite book of secrecy a little
I can read," here is a page for whose interpretation no
Daniel yet has come. Beyond the veil of recorded
science an insoluble, at least an unsolved, mystery lies.
This we perceive: Every ant is a law unto itself;
and in every individual the self-directing faculty is well-
nigh perfect. There is no private property. All citizens
are equals absolutely equals in ownership of the com-
munal property and in the use of, the authority over, and
the service and responsibility for the same. All serve,
save natural dependents; but all apparently are free to
choose the quality, the period, and the amount of service.
There is no visible head, no representative class or body
within which the control of the commonwealth is em-
bodied; and yet, by some occult force hitherto un-
known to men, all the beneficent effects of government
are wrought out with the regularity and precision of
an automatic machine. It is true to-day, as when
Solomon announced it many centuries ago, that this
work goes on without "guide, overseer, or ruler."
Here, in this strange commune, with its absolute law
and impersonal but imperial executive, the self-directing
])o\ver of every individual seems perfect. Is this social-
ism? nature's type of a practical socialism? Learn
this, then : if socialism as a form of human government
would be equally or even approximately successful, it
ni ust first attain that perfect individual discipline and ab-
solute self-control, self -abnegation, self-surrender, and self-
devotion to the good of the whole community that one sees
in a Commonwealth of Ants.
TABLE OF AUTHORS AND
FOR the sake of economy in space, and to save
breaking up the pages with foot-notes, the authors
and works referred to in the text of this book are des-
ignated thus: Every author, as a rule, has been given
as a symbol the first or the first two letters of his
name. His several works used, the titles of which are
printed in full in this table, are indicated by a number
or by successive numerals. For example, [McC. 7, p.
243] shows that the text refers to the seventh title
(Nature's Craftsmen) listed under the name McCook.
Due acknowledgment has thus been made throughout
the text of those to whose writings the author has been
largely indebted. Yet, among these fellow-naturalists,
he here expresses his special indebtedness to the veteran
myrmecologist Prof. August Forel, M.D., of Zurich,
who inspired and guided his earliest studies; and to
Prof. William Morton Wheeler, of Harvard University,
who stands primus inter pares among the later and
younger myrmecologists, by whose devoted and in-
telligent labors the writer and readers of Ant Com-
munities have profited.
Ad. ADLERZ, GOTTFRID.
Myrmecologiska studier. Formicoxenus nitidulus. Of-
versigt af Kongl. Veterenskaps-Akademiens For-
handlingar, 1884. Nr. 8, Stockholm.
Ai. AITKEN, E. H.
Red Ants' Nests. Bombay Nat, Hist. Soc., 1890, vol.
v., No. 4.
An. AXDRE, ERNEST.
1. Les Fourmis. Paris, Libraire Hachette et Ce 7 1885.
2. Species des Formicides d 'Europe. Gray (Haute-
Saone). Francis Bouffant, 1881-1882.
AY. LORD AVEBURY. See Lubbock.
C. CALDERWOOD, PROF. HENRY, LL.D., University of
The Relations of Mind and Brain. London, Macmillan
& Co., 1884.
De. DEAN. G. A.
The Mound-Building Prairie Ant (Pogonomyrmex occi-
dcntalis). Bulletin 154, Kansas State Agricultural
E. EMERY, PROF. CARLO, Bologna.
1. Zur Biologie der Ameisen. Sonderabdruck aus dem
Biologischen Centralblatt. Bd. xi., Nr. 5 u. 6
ausgegeben am 1. April, 1891.
2. Arbeiterstandes bei den Ameisen. Ueber Entstehung
des Soziallebens bei Hymenopteren. The same.
Bd. xiv. January, 1894.
3. Beitrage zur Kenntniss der nordamerikanischen
Ameisenfauna. Zoologischen Jahrbiichern. Abth.
pin. Syst., etc. Siebenter Band, pp. 633-681.
4. Ossewationi ad Esperimenti sulla Formica Amazzone.
Nota letta alia R. Acad. delle. Institute di Bologna.
Fd. FIELDE, Miss ADELE M.
1. Tenacity of Life in Ants.
2. Portable Ant-Nests. Biol. Bull., vol. ii., No. 2, 1900.
3. Artificial Mixed Nests of Ants. Biol. Bull. No. 6,
TABLE OF AUTHORS AND REFERENCES
Fo. FOREL, PROF. AUGUST, M.D., Zurich.
1. Les Fourmis de la Suisse. Geneve. 1874.
2. Etudes Myrmecologiques en 1884, avec une description
des organes sensoriels des antennes. Bull. Soc.
Vaud. Sc. Nat., xx., 19.
3. Die Nester der Ameisen. Neujahrsblatt heraus-
gegeben von der Naturforschenden Gesellschaft
auf das Jahr, 1893.
4. Ants' Nests. The same as No. 3, translated by Pro-
fessor Wheeler. Smithsonian Report for 1894, pp.
5. Ants and Some Oth r Insects. Translated from the
German by Prof. William M. Wheeler. The Open
Court Pub. Co., Chicago.
6. Habits of North American Ants. Revista di Sci.
t Biol., vol. ii. , No. 3, pp. 1-13. (Quoted by Wheeler.)
7. Etudes Myrmecologiques en 1886. Annales de la
Societe Entomologique de Belgique, tome xxx.
8. Hermaphrodite de 1'Azteca instab'lis SMITH. Bull.
Soc. Vaud. Sc. Nat., xxviii., 109. PL xvi.
9. Eine myrmekologische Serienreise nach Tunesien und
Ostalgerien nebst einer Beobachtung des Herrn
Gleadow in Indien iiber Aenictus. "Humboldt"
Band ix., Heft 9.
Go. GOULD, THE REV. WILLIAM, Exeter College, Oxon.
An Account of English Ants. London, 1747.
H. HUBER, M. P.
The Natural History of Ants. Translated by J. R.
Johnson M.D., London, 1820.
Hb. HUBER, JAKOB.
Ueber die Koloniengri'mdung bei Atta Sexdens. Biol.
Centralbl. xxv., 1905, pp. 606-619, 625-635.
Hu. HUBER, FRANCIS.
New Observations on the Natural History of Bees.
3d ed., Edinburgh, 1821.
I - -VoN IHERING, DR. H.
1. Ameisen von Rio Grande do Sul. Berlin Entomog.
Zeits. Bd. xxxix., 1894, Heft 3.
2. Die Anlage neuer Colonien und Pilzgarten bei Atta
sexdens Zool. Anzeig. 21 Jahr 9, 1898.
Ja. JANET, CHARLES, Paris.
1. Etudes sur les Fourmis. Structure des Membranes
Articulaires des Tendons et des Muscles (Myrmica,
Camponotus). Limoges, 1895.
2. Les Fourmis. Address on the Occasion of the Re-
union Generate annuelle de la Societe Zoologique
de France Paris, 1896.
3. Etudes sur Fourmis. Note 17, Systeme glandulaire
tegumentaire de la Myrmica rubra. Note 18,
Aiguillon de la Myrmica rubra. Appareil de ferme-
ture de la glande a venin. Paris, Georges Carre et
C. Naud, 1898.
4. Anatomic du corselet de la Myrmica Rubra Heine.
Note 19, Memoires de la Societe Zoologique de
5. Rapport des Animaux Myrmecophiles avec les Four-
mis, pp. 1-99. Limoges, 1897.
6. Sur les Muscles des Fourmis, des Guepes et des Abeilles.
Extrait des Comptes rendus hebdornadaires des
seances de 1'Academie des Sciences. Paris, 1895.
Lb. LUBBOCK, SIR JOHN, BART. (Lord Avebury).
1. Ants, Bees, and Wasps. London, Kegan Paul, Trench
& Co., 1882.
2. Journal Linnean Soc. Zoology, vol. xii., p. 487, Ob-
servations on Ants, Part iii.
3. Observations on Ants, Bees, and Wasps. Part xi.
Lim. Soc. Journal Zoology, vol. xx., pp. 118-136.
TABLE OF AUTHORS AND REFERENCES
See also the same journal for Lubbock's original
observations on Ants, subsequently published in
book form, substantially.
4. On the Habits of Ants. Popular Science Monthly,
Li. LIVINGSTONE, DAVID, LL.D., D.C.L.
Travels and Researches in South Africa. Harper &
Brothers, N. Y., 1858. Pp. 149, 576, 577.
Ln. LINCECUM, GIDEON, M.D.
On the Agricultural Ant of Texas. Proceed. Acad. Nat.
Sci., Philadelphia,, 1866.
Ma. MARSHALL, PROF. WILLIAM, Universitat Leipzig.
Leben und Treiben der Ameisen. Leipzig, 1889.
McC. McCooK, HENRY C, D.D., Sc.D., LL.D.
1. Notes on Architecture and Habits of Formica (Cam-
ponotus) pcnnsylramca, the Pennsylvania Carpenter
Ant. Trans. Amer. Entomolog. Soc., Philadelphia,
vol. v., 1874.
2. The Mound-Making Ants of the Alleghanies, Formica
exsecto'ides. Trans. Amer. Entomol. Soc., vol. vi.,
3. The Agricultural Ant of Texas. 1880. (Book now
out of print.) Lippincotts, Philadelphia.
4. The Honey -Ants of the Garden of the Gods. Pro-
ceed. Acad. Nat. Sci. of Philadelphia, 1881, pp.
17-77, Plates i.-x.
5. The Honey and Occident Ants. (Book now out of
print.) Lippincotts, Philadelphia, 1882.
6. Tenants of an Old Farm. George W.Jacobs & Co.,
Philadelphia. London, Hodder & Stoughton, 1889.
Chapter xiii., p. 231, on Cutting Ants of Texas.
7. Nature's Craftsmen. Harper & Brothers, N. Y.,
8. Ants as Beneficial Insecticides.' Proceed. Acad. Xat.
Sci., Philadelphia, 1882, p. 263.
9. On Certain Ants Associated with the Cotton Worm.
Professor Comstock's Report, 1870, p. 182. U. S.
Depart . Agriculture.
10. Observations on the Cutting Ants of Texas.
Proceed. Acad. Nat. Sci., Philadelphia, 1879, p.
11. Combats and Nidification of the Pavement Ant
(Tetramorium ccespitum). Proceed. Acad. Nat. Sci.,
Philadelphia, 1879, p. 156.
12. Note on the Adoption of an Ant Queen (Crenmsto-
gaster lineolata). Acad. Nat. Sci., Philadelphia,
1879, p. 137.
13. Mode of Depositing the Ant Eggs. Acad. Nat. Sci.,
Philadelphia, 1879, p. 140.
14. Note on Marriage Flight of Ants. Acad. Nat. Sci.,
Philadelphia, 1879, p. 141.
15. Note on a New Northern Cutting Ant (Atta septen-
trionalis McCook). Acad. Nat. Sci., Philadelphia,
1880, p. 359.
16. The Shining Slave-maker Pohjergus lucidus: Archi-
tecture and Habits. Acad. Nat. Sci., Philadelphia,
1880, p. 376. PI. xix.
17. Mode of Recognition Among Ants. Acad. Nat. Sci.,
Philadelphia, 1878, pp. 15 sq.
18. Modification of Habits in Ants through Fear of
Enemies. Acad. Nat. Sci., Philadelphia, 1887, p.
27. (Slave-Making Ants and Their Slaves.)
19. Prolonged Life of Invertebrates. Acad. Nat. Sci.,
Philadelphia, 1887, pp. 369-386. (Sir John Lub-
bock's "Old Ant Queen.")
20. Toilet Habits of Ants. Acad. Nat. Sci., Philadelphia,
1878, p. 119.
21. Note on Mound-Making Ants, Formica exsccto'ides.
Acad. Nat, Sci., Philadelphia, 1879, p. 154. (Gath-
ering Pine-Bark Pellets, Foraging for White Ants,
etc.; Moss-Grown Mounds, Closing Gates.)
TABLE OF AUTHORS AND REFERENCES
22. The Rufous or Thatching Ant, of Dakota and Colo-
rado. Acad. Nat. ScL, Philadelphia, 1884, pp. 57 sq.
23. The Sense of Direction in a European Ant (Formica
rufa). Acad. Nat. ScL, Philadelphia, 1887, pp. 336
sq. (Mounds, Road-Making, Sentinels.)
24. American Spiders and Their Spinning-Work. Vols.
i., ii., iii., quarto. P. Blakiston & Son, Philadelphia.
Ml. MOLLER, DR. ALFRED.
Die Pilzgiirten einiger siidamerikanischer Ameisen.
Jena, Gustav Fischer, 1893.
MG. MOGGRIDGE, J. TREHERNE, F.L.S., London, 1873.
Harvesting Ants and Trap-Door Spiders.
My. MAYR, DR. GUSTAV.
1. Die Formiciden der Vereinigten Staaten von Nord-
amerika, Wien, 1886.
2. Siidamerikanische Formiciden. Wien, 1887.
Ro. ROMANES, GEORGE J.
1. Animal Intelligence. D. Appleton & Co., N. Y.,
2. Intelligence of Ants. Popular Science Monthly,
Sa. SAUSSURE, HENRI DE.
Les Fourmis Americaines. Archives des sciences phy-
siques et naturelles, t. x., Geneve, 1883.
St. STANLEY, HENRY M.
Through the Dark Continent. Vols. i., ii., Harper &
Brothers, N. Y., 1878.
Tr. TREAT, MRS. MARY.
1. A Chapter in the History of Ants. Harper's Maga-
zine, January, 1879, p. 177.
2. My Garden Pets. D. Lothrop Co., 1887.
W. WHEELER, PROF. WILLIAM MORTON, Harvard Uni-
1. Compound and Mixed Nests of American Ants.
American Naturalist, Nos. 414, 415, 417. 418. Re-
print by Ginn c\: Co., Boston, 1901.
2. On the Founding of Colonies by Queen Ants, with
Special Reference to the Parasitic and Slave-Holding
Species. 1906, Bull. Am. Mus. Nat, Hist,, vol.
3. Notes on a New Guest- Ant. Bull. Wisconsin Nat.
Hist, Soc., vol. v., No. 2, 1907.
4. Ethological Observations on an American Ant.
Journal fur Psychologic und Neurologic, ii., pp.
1-31. Leipzig, 1902.
5. A Crustacean-Eating Ant. Biological Bulletin, vol.
vi., No. 6, May, 1904.
6. The Polymorphism of Ants, with an Account of Some
Singular Abnormalities Due to Parasitism. Bull.
Am. Mus. Nat. Hist,, vol. xxiii., 1907.
7. A New Type of Social Parasitism Among Ants. Bull.
Am. Mus. Nat, Hist., vol. xx., art, xxx., pp. 347-
8. The Habits of the Tent -Building Ant (Cremastogaster
lineolata SAY). Bull. Am. Mus. Nat. Hist., vol.
9. Ants' Nests. Translation of Doctor Forel's paper on
Ants' Nests for Smithsonian Report for 1894.
10. Comparative Ethology of the European and North
American Ants. Journal fiir Psychologic und Neu-
rologie, pp. 400-435, Leipzig, 1908.
11. A European Ant (Myrmica hvinodus) Introduced
into Massachusetts. Journal of Economic Entomol-
ogy, vol. i., No. 6, 1908.
12. Studies on Myrmecophiles. I. Cremastochclus. Jour-
nal N. Y. Entomolog Soc., vol. xvi., No. 2, June, 1908.
13. Ditto. II. Hetjerius. Id., Sept., 1908.
14. Ditto. III. Microdon. Id., Dec.. 1908.
TABLE OF AUTHORS AND REFERENCES
Wa. WASSMAN, THE REV. FATHER EDMUND (S. J.).
Die zusammengesetzten Nester und gemischten Kolonien
der Ameisen. Minister, Aschendorffische Buch-
1. Bombay Nat. Hist. Soc., 1892, pp. 56, 58. Quoted
W. 5, p. 256.
ACACIA, Mexican, nest of ants
Accidents repaired, 50.
Achray, ants in Pass of, 58.
Adaptation of means, 50.
Adlerz, Gottfrid, 108.
.Ecophylla smaragdina, 34.
African ants, 34, 192.
Agricultural ant, 20, 21, 22, 55,
130, 187, 206, 257, 286; open
plaza, 56; roads, 56; transpor-
tation, 57; feeding, 84, 146.
Agriculture, Department of, 114.
Agriculture destroying ants'
Alien associates and affinities in
ant communes, 224 et se<j.
Alleghany mound-makers, 151.
Amazons, ants are, 155.
Ancestral habit, 30.
Andre, M. Ernest, 233.
Anger and rudeness, absence of,
toward dependents, 180.
Ant cows. See Aphides,
Antennae, organs of smell, 81,
123; as hearing organs, 148;
sensibility of, 149; functions
of, 150; cleansing, 50.
Antennal parley, 153.
Ants, factors determining be-
havior, 181; lack of personal
benevolence, 292; communal
loyalty the onlv benevolence,
Aphis herds and ant associates,
243 et seq.
Aphides, 30, 102, 227, 244, 245;
of Formica exsectoides, 5, 13.
Aphenogastcr tennesseensis, 35.
Appetite, pleasures of an ant's,
Architecture, nesting, 17; sim-
plest types, 26; development
of, 26, 27.
Argiope aurantium (spider), 124.
Ashmead, Professor, 233.
Asian ant, 34.
Atta fervens, 72, 86, 87, 92, 146;
big-headed workers, 205.
Atta sexdens, 95.
Attaphila fungicola, 229.
Attidsp, 37, 66, 86.
Auditory organs, 147.
Avebury, Lord (Sir John Lub-
bock, Bart.), 152, 210, 273,
BANKOK, Siam, ant of, 31, 32.
Bees, language of, 125, 127;
spiracular voicing, 137.
Beetles, with ants, 225; Lome-
Behavior of ants, factors deter-
Bell wood, Pennsylvania, ant, 35.
Big-headed workers, 116, 203.
Boll- weevil, destroyed by ants,
Brains of ants, 183.
Brunelli, Signer, 140.
Buckley, Prof. S. B., 86.
Building up mounds, 43, 46.
CAGES for stridulating insects,
Calderwood, Prof. Henry, 150.
Callow antlings, 164.
Camponotus abdominah's, 199.
Camponotus americanus, 107.
Camponotus maccooki, 237.
Camponotus pennsylvanicus, 192.
Cnni])onotus rufipcs, 32.
Cannibals, ants as, 80.
Captives carried, 272.
Carpenter ant queen, 18.
Carpenter ants, 222.
Carton nests of ants, 29, 30,
Catacombs of Rome, an analogy,
Caves, mushroom, of ants, 94.
Cecropia adenopus, 36.
Cedius ziegleri, 231.
Cereal foods, taste developed for,
Ceylon ant, 34.
Chalcid parasite, 233.
Challenge of antenna?, 151.
Christian theory of war, 190.
Cicada, stridulating, 142.
Cleanliness, habitual, 50; tools
Cleansing body and head, 283.
Cockroaches, with ants, 225.
Cold, effects of, 228.
Column of march, 117.
Communal analogies, ants and
men, xv, xvi.
Communal dependents, the prob-
lem of, 171 et seq.
Communal life, influence of, 297.
Communal righteousness, 10, 15.
Communicate, how ants, 136 et
Comstock, Prof. J. H., 114.
Cooper, William S., Detroit,
Courage, 11, 12, 198.
Courtiers of queen, 300.
Cowardice, rare, 195; example of,
Cremastogaster, architecture of,
27, 29; form of, 28, 32.
Cremastogaster (brevispinosa} ,
Cremastogaster lincolata, 29.
Cricket, wing-cover of male, for
Criminal and degenerate ants, 9,
Crustaceans, ants feed on, 120.
Cutting ants, 205; underground
engineering, 64; head of, 88;
night raids, 90; plants pre-
ferred by, 91; forms of, 93;
caves of, 94.
Cutting leaves, mode of Atta, 89.
DARWIN, CHARLES, 273.
Dean, Mr. G. A., 25.
Death's-head moth, 138, 139.
Degeneration, without young,
Dependents, immature tax on
communes, 172; winged, 184.
Development of honey-bearers,
Devotion to commune, 13, 194.
Differentiation of functions of
Digestive tract, 110.
Dipteron parasites, 238.
Discord, absence of, 55.
Distribution, universal, 224.
Dolichoderus bituberculatus, 31,
Dorymyrmex flavus, 196.
Dorymyrmex pyramicus flarus,
Drainage of nests, 285.
Dumping-grounds, or chambers,
in nests, 23, 115.
ECITON, or " driver ant," 28.
Eciton Carolinense, 153.
Ectatomma tuberculatum, 114.
Education by the State, 168.
Eggs, eaten, 78; taken from
queen by workers, 161; of
Emery, Prof. Carle, 36, 233.
Engineering, 37, 40, 43, 49, 63.
Engineering methods in ant
structures, 49 et seq.
FACE of ant, 148.
Fecundation, in nest, 174.
Feeding the commune, 101 et seq.
Female ant, 77; after marriage
Female government in nnt com-
munities, 155 et seq.
Female temperament, 156; ex-
hibited with young, 165.
Females, .spurious pseudogynes,
Fetid ant, 257.
Fielde, Miss Adele, 10S, 273.
Florida harvester, 117, 204.
Florissant basin fossils, 176.
Food, 77; supply, variation in,
107; solid, given, 109; bearers'
arrival in nest, 179; quarrels
Food call, in ants, 146.
Foot, 53, 54.
Foraging habit formed, 78.
Forel, Dr. August. 8, 31, 32, 34,
153, 154, 199, 200, 256, 257, 273.
Formic acid of ants, 38, 101;
marking out roads, 59; fumes
as guide, 63; batteries, 210.
Fort Clark, Texas, ants, 65.
Fossil ants, of Oligocene and
Fossils, ants burrow in, 24.
Founder of commune, 77.
Founding community, 17.
Fountain, Gen. S. W., 65.
Fraternization, 6, 7.
Fruits, as ant food, 82.
Forelius faetidus, 257.
Forelius maccooki, 257.
Formica diffitilis-conscians, 264.
Formica ex secta, 8.
Formica ex sectoides, 2, 3; size of
communities, 8, 37, 61, 225.
Formica fn.sca, 263; of Baltic
Formica gnava, 227.
Formica pratensis, 255.
Formica rufibarbis, 256.
Formica Sanguinea, 265.
Formica sanguinea - rubicund a,
Formica Schaufussi, 153, 230, 258.
Formica Schaufussi-inccrta, 230,
Formica subsericea, 153, 199, 263.
Formica ru/a, 58, 61, 265.
Fraternal confederacies, 1 et seq.
Fury, under assault, 41.
GALLERIES, as lodgings, 41;
Galls, oak, as food, 110.
Gardner on insect music, 131.
Gate closing, Occident ants, 68,
70; opening, 69.
Gate closing, cutting ants, 72-75.
Goeldi, Dr., 95.
Good-temper and harmony, 194.
Grass, use of, in building, 38.
Grasshopper, shrilling, 140.
Gryllus riridissimus, 140.
Guards, 188. See also Sentinals.
Gustatory organs, 80.
HAIRS, specialized, 149; and
Hardening material, 42, 44.
Harvesting ants, 119.
Hearn, Prof. Lafcadio, 141.
Herbert, Edmund, 241.
Hill, Mr. R., 24.
ants of, 2.
Honey-ants, 274, 290; nest and
sentinels, 10; exterior of nest,
19; face, 80; abdomens of, 103;
oak galls with, 110; structure,
111; queen, 173.
Honey-bearers in natural site,
Honey-dew as food, 30, 102.
Horse ants, 58.
House-fly spiracles, 136.
Huber, Francis, pere, 137, 138.
Huber, Jakob, 95, 97.
Human parasites, social, 233.
INDIVIDUALISM in ant commune,
Industry, martyrs of, 160.
Insectivorous, habit utilized, 110.
Instinct, co-operating, 41 ; elastic,
50; to preserve species, 180.
Intelligence, ants', key to, 150,
Internal economy of ant com-
Island Heights ants, 99.
JANET, CHARLES, 256.
Japanese insect cages, 141, 142.
KANSAS ants, Sapa Creek, 24, 25.
Katydid, stridulating, 143.
Kauffman, Col. A. B., U. S. A.,
"Kelep," the, 114.
Keys of insects' notes, 131.
Kohlrabi-fungus of cutting ants,
LABIAL palp, 80.
Labor, division of, 9; influenced
by physical condition, 51.
" Ladybirds" are beetles, 248.
Language of ants and other in-
sects, 121 et seq.
Larvae, as food, 83; care of, 164,
281; kept in common, 165, 182.
Lasius flavus, 174.
Lasius fuliginosus, 183.
Lasius niger, 152, 159.
Leadership of commune, 301.
Leaf-made tents, 35.
LeConte, Dr., Coleopterist, 230.
Legs of ants, 279.
Leidy, Prof. Joseph, 24.
Leptogenys elongata, 120.
Leptothorax emersoni, 25,0.
Leptothorax petiolatus, 199.
Life, waste of, 158, 160.
Light within nests, 185.
Lincecum, Dr. Gideon, 65.
Litter cleaned up, 287.
Living chain, working by, 35.
Livingstone, Dr. David, 9, 148,
Lobopelta chinensis, 119.
Lobopolata distinguenda, 118.
Loch Katrine, 58.
Lycurgus the Spartan, 167.
M AGO WAN, DOCTOR, Wenchow,
Maimed, described, 215.
Males, nonentities in work of
commune, 167; not fighters,
Mandibles, 80, 206, 270.
Massagerie among ants, 228.
Mathematical skill in road mak-
Matriarchate, an ant commune,
Maxillary palp, 80.
Mclanoplifs femur-rubrum, red-
legged locust, 141.
Mendicants killed, 109.
Metopina pachycondylce , 236, 238.
Mimetic language, 122, 123.
Mimetic entreaties, 108.
Mites, parasites, 282.
Moggridge, J. Treherne, 116.
Moller's corpuscles, 37.
Moller, Dr. Alfred, 37, 95, 96.
Monomorium pharaonis, 82.
Morris, Rev. George K., 99.
Mosquito voicing, 137.
Mound-making ants of Alle-
ghenies, 2, 3, 85, 101. See
also Formica exsectoides.
Mounds, sectional views, 42.
Mushroom gardening ants, 95,
Mycetophagous ants, 120.
Myrmecocyst us hort ideoru m, 10;
exterior architecture, 19. See
also Honey ants.
Myrmecophylous insects, 229.
Myrmica brevinodis, 250.
"NATURE'S CRAFTSMEN," ant
studies therein, xv; on slave-
making ants, chap, v, 19, 85,
104, 110, -160, 175, 243, 263,
Nest odor acquired by aliens,
Night' life, 224.
Noiseless toil, 48.
Northern cutting ants, 98.
Numbers of ants, 92.
Nurse ants, duties, 167.
Nursery, emmet, 166.
Nurses, female, 165.
OCCIDENT ant, exterior nest, 24;
closing gates, 67, 68; store-
rooms and dumping-rooms,
116; gathering seed, 117; in
action, 196; in duel, 207;
parasitic nests on, 259; sting
of, in fights, 260.
Odor, specific, in recognition, 14;
of nest as giving recognition,
Oils, as ant food, 81.
Olfactory sense, 148.
Opportunists, ants as, 35.
Orange-trees preserved by ants,
Orasema coloradensis, 233.
Orasema viridis, 233, 234.
Orb weaver's eggs, 2.
Overseers, absence of, 40.
Pachycondyla harpax, 240.
Panics, spread of, 178.
Parasites on ants, 233, 236, 282.
Parasitic formicaries, 259.
Parasol or cutting ants, 86. See
Pasteboard ant nest, 31.
Patriotic feeling, supreme, 194.
Peace among men, 278.
Pear-tree psylla, 246.
Pennsylvania harvester, 118.
Pheidole commutata. 108.
Pheidole instabilis, 116, 203, 233,
Pheidole kingi, 233.
Pheidole pennsylvanica, 27, 118.
Philadelphia Academy of Natural
Philosophy of ants, 51.
Physical condition affected by
Physical expression as language,
Pine leaves in nests, 30.
Piping of bee queens, 137.
Plant culture, 98.
Plant-lice, aphides, described, 243.
Play, ants sporting, 186, 187.
Pleasures of, 185.
Plesiobiosis, double nests, 257.
Pluck in small ants, 197.
Pogonomyrmex crudelis, forms of,
Pogonomyrmex occidentals s, 10,
20. See also Occident ant.
Police administration, 14.
Polyergus ru/escens, 153; lucidus,
Polyrachis jerdonii, 34; spinigera,
34; dires, 34.
Portage of pebbles, 25, 26; im-
plements for, 37.
Post-petiole in stridulation, 147.
Providing work for young, 169.
Pseudomyrma belli, 36.
Pseudomyrma (gracilis), 199.
Psychic contagions, 178.
Pteratic language, loO, 132.
Public sentiment, 15.
QUEEN, 268, 271; cutting ant
founds mushroom garden, 96;
mother, 156; surrounded by
guard, 157; eggs of, 158; among
workers, 159; winged, of honey-
ant, 173; not seen in wars, 201.
RAID, or march, of cutting ants,
88, 89, 119; raiding in column,
Rain, effect on work, 37.
Rations, supplying the com-
munal, 77 et seq.
Recognition, 14, 15, 101, 104; in
wars, 220; by odor, 223.
Red ants, house pests, 82.
Republic, the ideal, 171.
Resemblance to spider's work, 35.
Roads, 119; of Formica rufa, 59.
Romanes, George J., 150.
Roofing habit, 24, 25.
Roziies gongylophora, 98.
Rufous ants of Britain, 58.
SALIVARY secretions, 44.
Sanitation and personal benev-
olence, problem of, 279 et seq.
Santiago de Cuba, cutting ants
of, 90, 91.
Scavengers, ants are, 81, 288.
Schaufuss ant, 264.
Schimpfer, Prof. A. F. W., 37.
Scott, Sir Walter. 58.
Scouts, or pioneers, 83.
Scudder, Prof. Samuel C., 133,
Senses of ants, 134.
Sensibilities in antenmr, 149.
Sensory organs, key to intelli-
Sentinels, 123, 124, 299.
Shakespeare on insect com-
Shampoo dejeune, 252.
Shining slave-maker, 263.
Silence of ant work, 45, 48.
Silk-made nest, 32, 34.
Slave-making ants, 200, 258;
communes, founding of, 261
Slave-making habit, how formed,
Slavery, human, compared with
ant, 261-262; raid described,
Slaves eaten, 83.
Slingerland, Prof. M. V., 245.
Social insects, 1, 2.
Soldier caste, 11, 115.
Solenopsis fugax, 254, 256.
Solenopsis molesta, 257.
South African ants, 9.
South American ant, 32.
Sphinx atropos, 138.
Spiders, American, and Their
Spinning Work, McCook, 177.
Spiders, stridulating, 40; an-
cestral, and their habits, 177;
Spinning habit, 32.
Spiracles, use of, 136.
Spirit of the commune, 40.
Stanley, Henry M., 192.
Steering committee for queen.
Stem-mother of aphis, 246.
Stinging organs, rudimentary,
207-209; of Pogonomvrmex,
Store-rooms for grain, 21, 115.
Stridulation, as language, 140;
organs in ants, 143, 145, 146;
modes of, among locusts,
crickets, and grasshoppers,
141, 143; delicate sounds, 144.
Structure of ants, 111.
Subterranean career, 185.
Swarming of sexes, 174.
Sweet tooth of ants, 79.
Symbiosis, with plants, 37, 81.
Sympathetic excitement, incit-
ing to marriagj-flight, 178.
TENACITY of life, 108.
Termes flavipes, 85, 225.
Termites, 84, 117, 119.
Tetramorium ccespitum, 152, 212,
Tidiness of ants compared with
Tmesiphorus costatis, 230.
Tongue, 80; described and illus-
Tools, natural, of ants, 53.
'Topochemical trail," 154.
Trachymyrmex septentrionalis, 99.
Transportation methods, 57.
Trenchers, communal big-headed
Trossachs, ants of, 58.
Tunnels under streams, 66.
UBIQUITY of ants, 79.
Underground wars, 219.
Vieveck, Mr. H. L., 25.
Voice of ants, 136.
Von Ihering, Dr. H., 95, 98.
WAR, how ants carry on, 211;
origin, 274; brutal but nat-
ural, 190; alarms, 193; war
note of bee, 127; return from,
Warrior ants and their equp-
ment for war, 190 et seq., 258;
all workers are, 202.
Wassman, Father Edmund, 200,
231, 254-255, 257.
Waste products disposed of by
Watchers, queen's guard, 162.
Wheeler, Prof. William M., 17,
29, 86, 99, 145, 146, 175, 176,
199, 227, 236, 257, 258, 264,
White ants, 118. See also Term-
Winged forms of ants, 173.
Winged dependents, arrival of
Winter, influence of, 225, 227.
Workers, complex instincts, 182.
Wounded ants uncared for, 214.
Wroughton, Mr., 117.
Wyoming ants, 24.
XEXODUSA CAVA, 230.
YOUNG cared for, 77; care and
presence of, needful, 289.