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Title: Life among the ants

Author: Vance Randolph

Editor: E. Haldeman-Julius

Illustrator: Peter Quinn

Release date: January 7, 2026 [eBook #77638]

Language: English

Original publication: Girard: Haldeman-Julius Company, 1925

Credits: Carla Foust, Tim Miller and the Online Distributed Proofreading Team at https://www.pgdp.net


*** START OF THE PROJECT GUTENBERG EBOOK LIFE AMONG THE ANTS ***




LITTLE BLUE BOOK NO. 833
Edited by E. Haldeman-Julius

Life Among
the Ants

Vance Randolph

Drawings by Peter Quinn


HALDEMAN-JULIUS COMPANY
GIRARD, KANSAS




Copyright, 1925,
Haldeman-Julius Company


PRINTED IN THE UNITED STATES OF AMERICA




CONTENTS


Chapter                                  Page

   1.  Books About Ants                     4

   2.  The Ant’s Body                       5

   3.  Reproduction and Metamorphosis      12

   4.  The Harvesting Ants                 20

   5.  The Mushroom Growers                25

   6.  The Honey Ants                      30

   7.  The Legionary Ants                  36

   8.  The Red Slave Makers                46

   9.  The Amazons and Their Slaves        51

  10. Dairies and Guests                   54




LIFE AMONG THE ANTS




CHAPTER I

BOOKS ABOUT ANTS


There are many references to ants in the works of the ancients (Aesop,
Plutarch, Horace, Ovid and Pliny), and these were quoted and elaborated
by the mediaeval authors, but modern scientific investigation may be
said to begin with the nineteenth century. Since then an enormous
amount of work has been done by European scientists, but their papers
are scattered through the files of obscure scientific journals in a
great variety of continental languages, and are usually inaccessible or
useless to the American student who wishes to make a serious (but not
_too_ serious) study of ant life and behavior.

The first general treatise in English was doubtless Sir John Lubbock’s
famous work entitled _Ants, Bees and Wasps_, first published in 1881.
This work was for many years a sort of standard textbook on the
subject, and is still well worth looking into.

Another book which may be of use is _Animal Intelligence_, by George
Romanes. The sixth edition, which appeared in 1895, devotes more than
one hundred pages to the habits of ants.

Eric Wasmann has written a great number of books and papers about ants,
one of the best of which has appeared in English as _The Psychology
of Ants and of Higher Animals_, published in 1905. All of Wasmann’s
works are valuable and well worth reading, but they are marred by his
constant references to philosophical and theological matters which are
of no great interest to the general reader. Father Wasmann feels called
upon to demonstrate that ants, as regards their psychical powers,
are much nearer to man than are the anthropoid apes, and is forever
interrupting himself to defend his vitalistic biology and condemn the
theory of organic evolution.

By all odds the best work available on the subject is the large volume
called _Ants_, written by Professor William Morton Wheeler of Harvard
University, and published in 1910. This book is, in fact, not merely
the best but the only book required by the average student. There is,
of course, a great deal of material which is uncomprehensible to one
who has no particular technical background, but the whole thing is so
admirably arranged that the student has only to glance through the
table of contents to locate matter suited to his taste and training. I
have made a very free use of _Ants_ in the preparation of this booklet,
some sections of which are little more than epitomes or abstracts of
Wheeler’s chapters.




CHAPTER II

THE ANT’S BODY


The body of the ant, like those of other insects, is segmented, and
covered with a hard chitinous external skeleton. It is separated by
constrictions into three distinct parts, the head, which bears the eyes
and mouth-parts; the thorax, to which the wings and legs are attached;
and the abdomen, which contains most of the entrails and the sexual
apparatus.

_The Head, Eyes, and Mouth-parts._ The head varies greatly in shape
and size, but always bears a frontal plate or _clypeus_, just above
which the two jointed _antennae_ or feelers are attached. The antennae
contain a great number of minute structures which are supposed to be
connected with the sense of smell. Three small simple eyes or _ocelli_
are set in the top of the head, and two large _compound eyes_ are
located one on either side. The eyes are always very well developed in
the males, and somewhat less so in the females; the eyes of the workers
are relatively small, and the ocelli are sometimes lacking altogether.
The compound eyes are the principal organs of vision, while the ocelli
are supposed to register only very near objects.

Just below the clypeus are the mouth-parts, consisting of the _labrum_
or upper lip, a pair of powerful _mandibles_, another pair of jaws
called _maxillae_, and the _labium_ or lower lip. Both maxillae and
labium bear little _palpi_ or feelers, and are plentifully supplied
with taste-buds containing the gustatory cells. The tongue or _glossa_
with which the ant laps up its food is attached to the upper part of
the labium.

_The Thorax, Legs and Wings._ The ant’s thorax consists of four
segments. The first segment is known as the _prothorax_; it is quite
small, and bears the first pair of legs. The next segment, the
_mesothorax_, carries the second pair of legs and the front wings--when
wings are present. The third segment or _metathorax_ bears the
third pair of legs and the hind wings--if there are any wings. The
fourth segment is really a part of the abdomen, and is known as the
_epinotum_. On each side of the thorax are two breathing-holes or
_stigmata_, which communicate directly with the _tracheae_ or windpipes
which supply air to the interior tissues.

The ant has six legs, one pair attached to each of the three segments
of the thorax proper. Each leg consists of five parts, the _coxa_, the
_trochanter_, the _femur_, the _tibia_, and the _tarsus_ or foot. The
wings are four in number, and the venation is similar to that found
in other members of the order Hymenoptera, but the wings are not much
used in classification because the workers are always wingless, and the
females wear wings only for a part of their lives.

_The Abdomen and Its Appendages._ The ant’s abdomen is divided into two
parts, the slender _pedicel_ which articulates with the last segment
of the thorax, and the larger part of the abdomen called the _gaster_.
The pedicel is provided with a file-like structure, which by rubbing
against a non-striated segment produces a sound of very high pitch.
In some species the females and workers bear stings and poison glands
in the last segment of the gaster. The female has no ovipositor. In
the male the tip of the gaster usually bears three pairs of sexual
appendages; the two outer pairs are used in clasping the female during
copulation, and the inner pair, when held tightly together, form a
tube which functions as a penis.

_The Alimentary Canal._ The mouth is located between the maxillae,
and is provided with a little pouch called the _infrabuccal cavity_,
which is used to hold solid matter while the liquid nutriment is
being sucked out of it. When this has been accomplished the pellet
is thrown out. The liquid food passes back into the _pharynx_, and
then on through a slender tube called the _esophagus_, which is lined
with fine hairs. In the gaster the esophagus expands into the _crop_,
which acts as a reservoir; no food is absorbed through its walls,
but is often regurgitated to feed the young. Just back of the crop
is the _proventriculus_ or gizzard, the movements of which provide
the suction by which liquid is drawn up the esophagus and into the
crop, and the force by which food is regurgitated. The true _stomach_
is rather small, and it is here that the food is both digested and
absorbed. The _small intestine_ communicates with the stomach by a
valve, and is connected with a number of _Malpighian tubes_ which act
as kidneys, absorbing liquid waste from the blood and pouring it into
the intestine. The large intestine or _rectum_ receives the feces and
urine from the small intestine and expels them from the body by way of
the _anal opening_.

_The Circulatory System._ The _blood_ of the ant, like that of other
insects, is colorless, and contains several kinds of corpuscles. Its
function is to carry food from the stomach where it is absorbed to
other parts of the body where it is needed. The blood of insects has
no red corpuscles, and does not carry oxygen about. The blood is not
confined in definite veins and arteries as in the higher animals, but
percolates about through the entire body cavity. There is a simple
_heart_ in the dorsal part of the abdomen which pulsates and forces
blood forward through an _aorta_ into the head, from which it seeps
gradually back into the abdomen, to be pumped forward through the aorta
again. Thus a sluggish circulation is maintained.

[Illustration: Fig. I. Diagram showing internal structure. 1, mouth; 2,
pharynx; 3, infrabuccal cavity; 4, aorta; 5, esophagus; 6, heart; 7,
crop; 8, small intestine; 9, stomach; 10, Malpighian tubes; 11, large
intestines or rectum; 12, anal opening.]

_Respiration._ Ants have neither lungs nor gills, and the blood does
not carry oxygen into the cells and carbon dioxide out as in the higher
animals. As in most other insects, air is taken into the body through
breathing-holes or _stigmata_, and brought into direct contact with the
tissues. There are ten pairs of these stigmata in the ant--two pairs in
the thorax and eight in the abdominal segments. Each opens through a
sort of valve into a _trachea_ or wind-pipe, which branches until its
ramifications extend to all parts of the body. When certain muscles
contract the size of the body increases, and air is drawn in through
the stigmata; when the size of the body is decreased the air is forced
out. The incoming air brings in the necessary oxygen, and the outgoing
current is laden with carbon dioxide waste from the tissues.

_The Nervous System._ The _brain_ proper is a mass of nerve matter in
the head just above the esophagus, but the _subesophageal ganglion_ is
very close to it, and the two are connected by heavy fibers on each
side of the esophagus, so that the whole thing has the appearance of
a brain with the gullet running through the middle of it. The major
part of the upper brain is connected with the compound eyes, but there
are nerves also which supply the ocelli, the antennae, the pharynx,
the labrum, and muscles in the head. The subesophageal ganglion gives
off nerves to the mandibles, maxillae and labium. From the lower back
part of the subesophageal ganglion the _ventral nerve cord_ arises, and
runs through the thorax and far back into the abdomen. This cord bears
three large _thoracic ganglia_ which innervate the muscles of the wings
and legs. In the abdomen are eleven smaller _abdominal ganglia_, with
nerves running out to supply all of the abdominal organs. The so-called
_sympathetic system_ consists of a few very small ganglia and nerves
not directly connected with the ventral nerve cord, which function in
connection with the digestive organs.

_The Reproductive Organs._ The _ovaries_ of the female or queen ant
are located in the upper and front part of the gaster, and each is
connected by a slender _oviduct_ with the _uterus_. The uterus is
continuous with the _vagina_, the external opening of which is located
near the tip of the abdomen. At the top of the uterus is a small pouch
called the _seminal receptacle_, which receives the sperm from the male
in copulation. The spermatozoa live in this pouch for several years,
and meet and fertilize the eggs as they descend into the uterus from
the ovaries.

The organs of the worker are similar to those of the queen, except that
they are very much smaller, and are usually incapable of functioning
normally. Worker ants have never been seen to copulate. The _testes_
of the male ant are located in the front part of the gaster, and are
connected by the _vas deferens_ with the _seminal vesicles_. Tubes from
the vesicles unite to form the _ejaculatory duct_, which is connected
with the _penis_ at the tip of the abdomen.




CHAPTER III

REPRODUCTION AND METAMORPHOSIS


Like their relatives the bees and wasps, ants have developed two
types of females, so that a colony contains three distinct sorts of
individuals, known as males, females, and workers.

_The Male._ The male is less subject to variation than either the queen
or the worker. The body is usually slender and graceful, the eyes and
antennae are well developed, and the mouth parts rather small and weak.
In most species the male is winged. As in the bees, the one great
function of the male in the colony is to copulate with the female or
queen, so as to supply her with sperm to fertilize future eggs. The
male is not killed in the course of the sexual embrace, as the drone
honeybee is, but usually dies soon afterward.

_The Female._ The true female or queen is usually larger than either
the male or the worker; the head, eyes, and mandibles are well
developed, and the abdomen is very large to contain the reproductive
organs. The female is usually winged at the time of mating, but the
wings are loosely attached and she loses them as soon as the nuptial
flight is over. The wings and legs are stouter and shorter than those
of the male, in most cases. In a few species the females have no wings,
and in others it is the males which are wingless. No case is known in
which neither male nor female is provided with wings.

_The Worker._ The worker is an undeveloped, wingless female. The eyes
are small, and the ocelli are often lacking; the antennae, legs, and
mouth parts are strong and well developed. There is a great deal
of variation among workers; one common variant is the _dinergate_,
or soldier--a form with a very large head and mandibles adapted to
fighting. The sex organs of the worker are unquestionably female, but
they do not ordinarily function, and a worker has never been seen to
copulate.

_Mating._ In species in which both the male and female are winged,
mating occurs in the air, as in the nuptial flight of the queen bee. In
the case of the honeybee, however, there is only one queen to a great
number of drones, while with the ants there may be hundreds of queens
and drones in the air, all copulating at once. Another difference is
that the mated females do not often return to the parent colony, as the
queen bee always does. When the mating hour draws near all the ants,
even the nearly blind and wingless workers, rush out of the nest in
great excitement, and the air is soon full of flying ants. Copulation
usually begins high in the air, but the linked pairs often fall to
the ground together. In the mating of bees the male is almost always
instantly killed, the genital organs and entrails being torn out of
his body. This mutilation never happens among ants, but the male’s
life-work is ended with the sexual act, and he usually dies shortly
afterward.

_The New Colony._ As soon as the mated female is upon solid ground
again she tears off her wings, or removes them by rubbing against
some solid object. This done, condemned to a crawling, terrestrial
existence for the rest of her days, she sets out alone to establish
a new colony. She digs a hole in the ground, or in rotten wood, or
under a flat stone, seals up the opening, and sits down in the dark
until the eggs in her abdomen are mature. Sometimes this takes weeks
or even months, and during this time the queen has nothing to eat,
but lives by absorbing the large wing-muscles which she will never
use again. Finally the eggs are deposited, being fertilized by some
of the spermatozoa which were obtained from the male, and which are
stored in the spermatheca, a little pouch just above the uterus. When
the larvae hatch she feeds them with a secretion from her salivary
glands. The resulting ants are normal workers, except that they are
unusually small. Sometimes it takes nearly a year to rear this first
brood, and all this time the queen has eaten absolutely nothing. As
soon as the workers are old enough they dig passages to the open air,
and enlarge the nest by adding galleries and runways. They drag in
food and feed the exhausted female, who from this time forward does
nothing but eat and lay eggs--the brood being cared for entirely by the
workers. From now on the female is a timid, photophobic, rickety old
egg-laying machine. During her long fast the great wing-muscles have
been absorbed, leaving the thorax hollow, so that she floats if placed
in water. Only a very few females can survive the ordeal necessary to
found a new colony--probably only one of many thousands which undertake
it. It is a beautiful example of the Darwinian phenomena of survival.

The procedure described above is the usual one in most species of
ants. It was guessed at by Huber in 1810, but the first man to watch
the actual founding of a new colony was an American named Lincecum,
about 1866. In 1879 Sir John Lubbock observed the whole process in an
artificial nest, and his account of the process has since been verified
by numerous other investigators.

In certain species, however, the queen is unequal to the task of
founding a family in this manner. In this case she must return to the
parent colony, join a queenless colony of her own or an allied species,
or raid a small colony of aliens. In this latter event she kills them
all, and adopts their eggs and brood.

_Complete Metamorphosis._ Like the butterflies and beetles, ants have
a complete metamorphosis, that is, they pass through four distinct
developmental stages. In many other insects--the grasshoppers for
example--the metamorphosis is said to be incomplete, because the newly
hatched young have the same general form as the adult, and their
development is merely a matter of increase in bulk.

_The Egg._ Ant’s eggs are very small, rarely more than one-fiftieth of
an inch in length, and are very seldom seen by the casual observer, who
mistakes the comparatively large cocoons for eggs. The egg is usually
elongated, and consists of the germinal spot, the yolk, and the thin
transparent shell called the chorion. The eggs look very much alike,
and one cannot predict whether a given egg will produce a male, a
worker, or a queen. Some eggs are fertilized by sperm stored in the
female’s spermatheca, others are deposited without fertilization, while
those laid by workers are certainly not fertilized, since workers do
not copulate. In bees and certain other related insects it has been
found that unfertilized eggs always produce males, but whether this is
always true in ants is still an open question.

Very little is known of the embryological development of the ant, but
the unhatched larva certainly has traces not only of antennae and legs,
but remnants of certain abdominal appendages not present in the adult
ant, and evidently harking back to more remote ancestors. The egg
usually hatches about twenty days after it is laid, but the length of
this period varies greatly with the temperature.

_The Larva._ The newly hatched larva is a soft, semi-transparent grub,
with a fat body, slender crooked neck and small head. There are no
eyes, but the mouth-parts are fairly well developed, and ten pairs of
stigmata are usually present. The body is covered with short fine
hairs. The digestive system is well formed, but there is no connection
between the stomach and the intestine, so that the larva has no
movement of the bowels until it is about to transform into the next
stage. The accumulated feces in the lower part of the stomach may often
be seen as a black spot showing through the semi-transparent walls of
the body.

[Illustration: Fig. II. Cross-section of an ant-hill, showing the
arrangement of larvae and pupae according to size. (Adapted from
Andre.)]

The larva is fed by the workers, the food being either regurgitated
liquid food or pieces of fresh vegetable or animal matter. It has been
found in the case of the bees that the kind of food given the larva
determines whether it will develop into a queen or a worker, but we
have no definite information about this matter among the ants.

When the larva is fully grown, usually about a month after hatching, it
is buried in the ground by the workers, and spins a silken cocoon about
itself. All ant larvae have spinning organs in the head, but some do
not spin cocoons, and in this case are not buried, but undergo their
metamorphosis in the open chambers of the nest. The larva now voids its
accumulated feces, sheds the larval skin, and appears as the pupa, the
third stage in the ant’s development.

_The Pupa._ In the pupal stage the ant has most of the appendages and
organs of the adult, but they are small and folded close against the
body. The pupa lies quietly, is not fed, and ordinarily gives no signs
of life at all. Gradually the various parts develop, the darker color
of the adult appears, until finally the mature pupa has very much the
appearance of the imago. Then the cocoon is opened by the attendant
workers, the young ant dragged out and fed, and begins its life as an
adult. The pale, newly emerged ant is known as a _callow_. The pupal
stage usually lasts from fifteen to twenty days, but is sometimes much
longer in cold weather.

_The Adult._ The general appearance and characteristics of the adult
are described elsewhere in this book. The total time of development
from the deposition of the egg to the appearance of the callow varies
from about sixty days to five months, and is considerably longer than
the corresponding period in most other insects. The queen bee, for
example, passes through all three stages in about sixteen days, while
some butterflies are developed in less than twenty-five days. Another
interesting feature is the extreme longevity of the adult ant. The
males are short-lived, but the workers of many species live for four
or five years, and the queens for still longer periods. Janet kept one
for fully ten years, and Sir John Lubbock had a queen in his possession
from December, 1874 to August, 1888, “when she must have been nearly
fifteen years old, and, of course, may have been more,” since he had no
means of knowing her age at the beginning of her captivity.




CHAPTER IV

THE HARVESTING ANTS


The works of Pliny and other ancient writers contain references to ants
which collected great stores of seeds, and these accounts were quoted
by numerous mediaeval authors. Modern students of ants, however, worked
mostly in northern and central Europe, and as they did not find any of
these harvesting ants they were rather inclined to dismiss the classic
stories as fiction pure and simple, and class the seed-gathering ants
with the unicorn and the mermaid.

In 1829, however, one W. H. Sykes, an Englishman located in India,
reported that certain ants near his station not only collected great
quantities of grass seed, but after a heavy rain could always be
seen bringing their cereal out of the underground granaries to dry
it in the sun. These observations went far to vindicate the ancient
naturalists, and the work of J. T. Moggridge, in 1873, completed the
vindication. Moggridge watched the workers bring in the seeds, bite
off the germinating part to prevent the seeds from sprouting, and
store them in the nests, which often contain a pint or so of grain. By
examination of these hoards he identified as many as eighteen different
families of plants represented in a single nest. Despite the efforts to
prevent germination by biting off the radicles (a fact noted by Pliny
some sixteen hundred years before) many of the seeds do sprout, and
thus the harvesting ants play a part in the distribution of plants. Of
this subject Moggridge says: “As the ants often travel some distance
from their nest in search of food, they may certainly be said to be,
in a limited sense, agents in the dispersal of seeds, for they not
infrequently drop seeds by the way, which they fail to find again,
and often also among the refuse matter which forms the kitchen hidden
in front of their entrances, a few sound seeds are often present, and
these in many instances grow up and form a little colony of strange
plants. This presence of seedlings foreign to the wild grounds in which
the nest is usually placed, is quite a feature where there are old
established colonies of _Atta barbara_, where young plants of fumitory,
chickweed, cranesbill, Arabis thaleana, etc., may be seen on or near
the rubbish heap.... One can imagine cases in which the ants during
the lapse of long periods of time might pass the seeds of plants from
colony to colony, until after a journey of many stages, the descendants
of the ant-borne seedlings might find themselves transported to places
far removed from the original home of their immediate ancestors.”

There are many species of harvester ants in America; one of the most
interesting is _Solenopsis geminata_, popularly known as the fire-ant
because of its readiness to use its painful sting. Although the
fire-ant certainly stores up seeds, often to the extent of damaging
crops of soft fruits like strawberries, it will also eat insects, or
almost anything else that it can get. The nests are usually found
beneath flat stones, and in some localities are so common and so
populous that Wheeler refers to the fire-ant as being “in possession
of a large portion of the soil of the American tropics.” In Louisiana
and other southern states these ants nest along the shores of lagoons
and bayous; when the floods come and the nest is submerged the workers
cling together in a ball as much as eight inches in diameter, with the
brood in the center. This ball floats in the water, the ants constantly
shifting about so that very few are drowned, and very little brood
lost, until they are able to effect a landing.

The so-called Texas harvester (_Pogonomyrmex molefaciens_) has become
famous because a man named Lincecum, about 1862, published a paper in
which he claimed that this ant actually _plants_ seeds in the ground,
weeds and cultivates its fields all summer, gathers the crop, dries
it in the sun, and finally stores it away in subterranian granaries.
This story was accepted and promulgated by Charles Darwin, and so was
believed in many quarters. It seems to rest solely upon the fact that
ant-rice (_Aristida_) is usually found growing about the nest, although
it may occur nowhere else in the immediate vicinity. “Four years of
nearly continuous observation,” writes Wheeler, “enable me to suggest
the probable source of Lincecum’s misconception. If the nests of this
ant can be studied during the cool winter months--and this is the only
time to study them leisurely, as the cold subdues the fiery stings of
their inhabitants--the seeds, which the ants have garnered in many
of their chambers will often be found to have sprouted. Sometimes, in
fact, the chambers are literally stuffed with dense wads of seedling
grasses and other plants. On sunny days the ants may often be seen
removing these seeds when they have sprouted too far to be fit for food
and carrying them to the refuse heap, which is always at the periphery
of the crater or cleared earthen disk. Here the seeds, thus rejected as
inedible, often take root and in the spring form an arc or a complete
circle of growing plants around the nest. Since the _Pogonomyrmex_
feeds largely, though by no means exclusively, on grass seeds, and
since, moreover, the seeds of Aristida are a very common and favorite
article of food, it is easy to see why this grass should predominate in
the circle. In reality however, only a small percentage of the nests,
and only those situated in grassy localities, present such circles.
Now to state that _molefaciens_, like a provident farmer, sows this
cereal and guards and weeds it for the sake of garnering its grain, is
as absurd as to say that the family cook is planting and maintaining an
orchard when some of the peach stones, which she has carelessly thrown
into the backyard with the other kitchen refuse, chance to grow into
peach trees.”

Wheeler has also observed the mating flight of the Texas harvester,
and his graphic description is worth setting down in its entirety:
“During three successive years (1901-1903) at Austin, Texas, the
nuptial flight of _molefaciens_ took place on one of the last days
of June (28 and 29) or the first in July. On one of these occasions
(July 4, 1903) the flight was of exceptional magnitude and beauty. A
few days previous the country had been deluged with heavy rains, but
Independence Day was clear and sunny, the mesquite trees were in full
bloom and the air resounded with the hum of insects. For several days
I had seen a few males and winged females stealthily creep out of the
nest entrance as if for an airing, but hurry back at the slightest
alarm. From 1:30 to 3 o’clock, however, on the afternoon of July 4,
all the numerous colonies I could visit during a long walk west of the
town, gave forth their males and females as by a common impulse. The
number issuing from a single large nest was often sufficient to have
filled a half liter measure. Soon every mound and disk was covered with
the bright red females and darker males, intermingled with workers,
many of whom kept on bringing seeds and dead insects into the nest
as unconcernedly as if nothing unusual were happening. The males and
females, quivering with excitement, mounted the stones or pebbles of
the nest or hurriedly climbed onto the surrounding leaves and grass
and rocked to and fro in the breeze. Then, raising themselves on their
feet and spreading their opalescent wings, they mounted obliquely one
by one into the air. I could follow them only for a distance of ten or
twenty meters when their rapidly diminishing bodies melted away against
the brilliant cloudless sky. Many pairs, hesitating to take flight,
chased one another about on the surface of the nest. The amorous males
seized many of the females before they could leave the ground. Lizards
crept forth in great numbers and gulped down quantities of the fat
females, while others were borne off into the air by large robber flies
(_Asilidae_). By a little after three o’clock the males and females had
left the nest and only the workers were seen pursuing the quiet routine
business of bringing in seeds.”




CHAPTER V

THE MUSHROOM GROWERS


In tropical and subtropical America there are about one hundred species
and varieties of ants which have most extraordinary habits, and are
grouped together in the Myrmicine tribe _Attii_. These ants are usually
rather small and dull colored, and, while they are powerful and
industrious diggers, are not given to rapid movements as most ants are,
but walk slowly and sedately about. When picked up they do not struggle
as many other ants do, but feign death after the manner of certain well
known beetles.

It was long noted that the _Attii_ carried great quantities of leaves
into their nests, and there was considerable doubt as to the use to
which these were put, some observers believing that they were used
immediately as food, and others contending that they served as roofing
and carpets in the underground passageways. Belt, a naturalist who
lived in Nicaragua, was probably the first to discover the secret
of the leaves. Digging into one of the nests in his garden, he was
surprised to find no great quantity of leaves in any of the passages,
although ants were continually bringing them in at the entrance. The
chambers were always partly filled with “a speckled, brown, flocculent,
spongy-looking mass of a light and loosely connected substance.... This
mass, which I have called the ant-food, proved on examination to be
composed of minutely subdivided pieces of leaves, weathered to a brown
color, and overgrown and lightly connected together by a minute white
fungus that ramified in every direction throughout it.... When a nest
is disturbed and the masses of ant-food spread about, the ants are in
great concern to carry away every morsel of it under shelter again; and
sometimes, when I dug into the nest, I found the next day all the earth
thrown out filled with little pits that the ants had dug into it to get
out the covered up food.”

Further investigation brought Belt to the conclusion that the _Attii_
do not eat leaves at all, but use them as manure to grow fungus on; and
further, that they feed upon this fungus, and will eat nothing else.
The _Attii_ are, in Belt’s own phrase, “mushroom growers and eaters.”
While leaves are the chief fertilizer, other substances are often
found suitable for growing fungus on; flowers are sometimes used, and
some species are particularly partial to pieces of orange peel. The
temperature and ventilation of the subterranean gardens are matters
of great importance, and there are many small holes which connect
the larger chambers with the surface. These air-shafts are plugged
and reopened at intervals, and by this means the temperature and
ventilation are regulated.

Alfred Moeller was a naturalist who studied the _Attii_ in Brazil, and
published the results of his labors in 1893. He found that the gardens
contain only one kind of fungus, all alien spores being carefully
weeded out. The ants do not allow the fruits to develop, and this has
made the classification of the fungi a very difficult matter. The fungi
found in the _Attii_ nests are different from any others known, but
no one can tell whether they are really distinct species or merely
modified forms of certain common moulds or mushrooms.

Von Ihering, in 1898, discovered that the virgin queen, when leaving
the nest on her nuptial flight, always carries a little pellet of
fungus in her mouth. After being fertilized by the male the queen
shuts herself up in a little burrow and sets about the founding of a
new colony. There are in this case no leaves available, and she starts
the fungus growing upon some of her new-laid eggs, which she crushes
for the purpose, and which seem to work quite as well as the usual
vegetable fertilizer.

J. Huber, in 1905, studied the same problems which interested Von
Ihering, and concluded that the fungus is not grown upon crushed eggs,
but is nourished by the liquid excrement of the queen. He describes
his observations as follows: “After watching the ant for hours she will
be seen suddenly to tear a little piece of the fungus from the garden
with her mandibles and hold it against the tip of her abdomen, which is
bent forward for this purpose. At the same time she emits from her vent
a clear yellowish or brownish droplet which is at once absorbed by the
tuft of hyphae. Hereupon the tuft is again inserted, amid much feeling
about with the antennae, in the garden, but usually not in the same
spot from which it was taken, and is then patted into place by means
of the fore feet.... According to my observations, this performance
is repeated usually once or twice an hour, and sometimes, to be sure,
even more frequently.” Although, according to Huber, the eggs are not
used directly as fertilizer for the fungus, the same result is brought
about indirectly, as the female is accustomed to feed upon her own
new-laid eggs. Huber estimates that nine out of every ten eggs laid are
eaten at once by the mother. The young larvae, too, are fed with eggs
thrust directly into their mouths by the queen. When the adult workers
appear, however, they live exclusively on the fungus which has been
growing during their larval life, and feed the queen upon fungus also,
while continuing to supply the larvae with their mother’s eggs. After
a week or so the workers dig their way out of the chamber, bring in
leaf-manure for the garden, and the fungus is no longer cared for by
the queen, who now gives all her attention to the serious business of
egg-laying. As the fungus becomes more abundant under this cultivation
it is fed to the larvae also, and eggs are no longer used as food by
any of the individuals in the hive.

The extraordinary habits of the Attine ants have fascinated many
students, and a number of theories about their development have
been advanced. Forel suggested that the ancestors of the present
mushroom-growers must have lived in rotten wood, and fed upon the
fungus which grew upon the moist walls of their nests, or upon insect
excrement. Von Ihering thinks that they may have developed from the
harvesting ants, which gradually acquired such an appetite for the
fungus which happened to grow in their granaries that the original
stores came to be used only as fertilizer. Wheeler points out that,
besides the Attine ants, there are several kinds of beetles and
termites which cultivate fungus upon their own excrement, and suggests
that originally this was the method employed by the ants. Later on they
came to use the excrement of other insects, and finally passed to the
addition of leaves and other non-fecal vegetable matter.

As has been said above, the _Attii_ are primarily tropical and
subtropical insects, but a few species have come north into the United
States. They are found chiefly in peninsular Florida, in southern
Texas, and in Arizona, although one species has been reported as far
north as southern New Jersey.




CHAPTER VI

THE HONEY ANTS


Many species of ants are in the habit of collecting nectar from
flowers, and the sweet juices excreted by plant-lice, until the crop
is greatly swollen. When they arrive at the nest, however, the sweets
are soon regurgitated and fed to the larvae. Any worker ant is able to
expand its crop to a certain extent, but in some species this power is
developed to an enormous degree. In still other tribes this peculiar
capacity seems to be limited to certain individuals. In the true honey
ants only a comparatively small number of workers are capable of this
honey-carrying, and these individuals are known as honey-bearing or
_repletes_. The repletes never accompany the other workers on their
foraging expeditions, but remain always in the nest, and are used as
living bottles in which to store the nectar brought in from the fields.

In some North American species of _Myrmecocystus_ the abdomen is
distended to such an extent that the repletes are unable to move about
without serious danger of bursting open, and spend their lives hanging
in clusters from the ceilings of certain chambers in the nest. These
honey ants are found in desert regions from central Mexico as far
north as Denver, Colorado, and have since ancient times been highly
prized as sweetmeats by the aborigenes of this region. Honey ants were
described in Mexican publications as long ago as 1832, but the first
important study was made by McCook, whose investigations were carried
out in the so-called Garden of the Gods, near Manitou, Colorado, about
1882. He found several very large nests, covering an area of more than
six feet in diameter, and extending three feet below the surface of
the ground. One of these nests contained some three hundred replete
honey-vessels hanging by their claws from the ceiling, and so distended
with honey that, once fallen from their positions, they were quite
unable to get back up again. McCook saw the ordinary workers bringing
in great quantities of nectar and honeydew, which was immediately
regurgitated and fed to the repletes or _rotunds_, as he called them,
and thus stored up in a living reservoir until needed.

[Illustration: Fig. III. Repletes of a common honey-ant. (From a
drawing by Wheeler.)]

It was formerly supposed that the sweet liquid was kept in the stomach
of the replete, but Forel, in 1880, showed that it is in reality
the enormously distended crop which functions. McCook made careful
dissections which bore out Forel’s views, and demonstrated that the
replete has all the abdominal organs of the ordinary worker, although
these are flattened against the body wall and rendered inconspicuous by
the distension of the crop.

McCook rejected the view that the replete belongs to a separate
caste, saying that “a comparison of the workers with the honey-bearer
shows that there is absolutely no difference between them except in
the distended condition of the abdomen.... The process by which the
rotundity of the honey-bearer has probably been produced, has its
exact counterpart in the ordinary distension of the crop in overfed
ants; the condition of the alimentary canal, in all the castes, is
the same, differing only in degree, and therefore the probability is
very great that _the honey-bearer is simply a worker with an overgrown
abdomen_.... Thus workers are transformed by the gradual distension
of the crop and expansion of the abdomen into honey-bearers, and the
latter do not compose a distinct caste.”

[Illustration: Fig. IV. Repletes of a honey-ant (_Myrmecocystus
hortideorum_) hanging from the roof of a honey chamber. (After McCook.)]

Just why these repletes should be developed in some species and not
in others is a mooted question. The fact that they are found only
in desert regions in North America, Australia, and South Africa may
mean that a dry climate is one of the important conditions of the
phenomena. Forel said: “The extraordinary distension of the crop seems
to be frequent in the Australian species of the general Melophorus,
Gamponotus and Leptomyrmex. I suppose that this is due to the extremely
dry climate of the country, which must compel the ants to remain,
often for long periods, in their subterranean abodes. At such times
a store of provisions in living bags must be very useful to them.”
Wheeler, in commenting on the above statement by Forel, writes: “There
can be little doubt of the truth of this statement, but I believe that
it should be expressed in a different manner. The impulse to develop
repletes is probably due to the brief and temporary abundance of liquid
food (honeydew, gall secretions, etc.) in arid regions and the long
period during which not only these substances, but also insect food
are unobtainable. The honey is stored in the living reservoirs for
the purpose of tiding over such periods of scarcity, and the ants
remain in their nests because they do not need to forage. Hence the
confinement mentioned by Forel is not the immediate but one of the
ulterior effects of drought. I am convinced from my observations on
desert ants that no amount of drought will keep these insects in their
nest when they are in need of food.

“While excavating the nests of _M. hortideorum_ I was impressed with
certain peculiarities in their structure and situation, which seem to
be explainable only as adaptations to the development of repletes.
One of these peculiarities is the great hardness of the soil that is
preferred by the ants. This is the more astonishing because the workers
are very slender and delicate organisms. It is evident that such soil
is well adapted to the construction of vaulted chambers like those in
which the repletes hang, whereas soft or friable soil would be most
unsuitable. The development of repletes also makes it necessary for the
ants to seek very dry situations for their nests. Hence we always find
them, in the environs of Manitou at least, on the summits of ridges
which shed the rain very rapidly. The honey chambers must be kept dry,
both to prevent the disastrous results of crumbling and slipping walls
and to obviate the growth of mould on the repletes, which are, of
course, imprisoned for life in dark cavities and filled with substances
that are favorable to the development of fungi. I believe also that
the size of the nest openings and galleries, which are so much larger
than would seem to be required by such small, slender ants, may be
an adaptation to securing plenty of fresh air in the honey chambers.
If these suppositions are correct, there is obviously a reciprocal
relation between the replete habit and an arid environment: the ants
store honey because they are living in an arid region where moisture
and food are precious, and the storing of honey in replete workers, in
turn, is possible only in very dry soil.”




CHAPTER VII

THE LEGIONARY ANTS


These insects, which Wheeler describes as “the Huns and Tartars of the
insect world,” are found in tropical Africa and Asia, and in the warmer
parts of America. There is a great variation in size and appearance
between the different castes, the females and workers being blind and
wingless, while the males have well developed wings and large compound
eyes. Some of these ants have no fixed habitation, but wander from
place to place, traveling mostly at night, and camping during the day
in any shallow hole that affords a temporary shelter. They cannot
endure the direct rays of the sun, and Savage, in 1845, observed that
“if they should be detained abroad till late in the morning of a
sunny day by the quantity of their prey, they will construct arches
over their path, of dirt agglutinated by a fluid excreted from the
mouth,” except when they can remain concealed by thick grass or leaves.
Sometimes the soldier ants form a sort of network arch with their own
bodies, and Savage says that “whenever an alarm is given the arch is
instantly broken, and the ants, joining others of the same class on
the outside of the line, who seem to be acting as commanders, guards
and scouts, run about in a furious manner in pursuit of the enemy.
If the alarm should prove without foundation, the victory won or the
danger passed, the arch is quickly renewed, and the main column marches
forward as before in all the order of a military discipline.”

In these marches the ants carry their eggs, larvae and pupae with them,
these being borne in the mandibles of the _minima_ or small workers,
and the whole column lives by foraging. Savage’s description of their
predatory habits is well worth quoting here: “They will soon kill the
largest animal if confined. They attack lizards, guanas, snakes, etc.,
with complete success. We have lost several animals by them--monkeys,
pigs, fowl, etc. The severity of their bite is increased to great
intensity by vast numbers, to a degree impossible to conceive. We may
easily believe that it would prove fatal to any animal in confinement.
They have been known to destroy the _Python natalensis_, our largest
serpent. When gorged with prey it lies motionless for days; then,
monster as it is, it easily becomes their victim.... Their entrance
into a house is soon known by the simultaneous and universal movement
of rats, mice, lizards, Blapsidae, Blattidae, and of the numerous other
vermin that infest our dwellings. Not being agreed, they cannot dwell
together, which modifies in a good measure the severity of the driver’s
habits, and renders their visits sometimes (though very seldom in my
view) desirable. Their ascent into our beds we sometimes prevent by
placing the feet of the bedsteads into a basin of vinegar, or some
other uncongenial fluid; this will generally be successful if the
rooms are ceiled, or the floors overhead tight; otherwise they will
drop down upon us, bringing along with them their noxious prey in the
very act of contending for victory. They move over the house with a
good degree of order, ransacking one point after another, till, either
having found something desirable, they collect upon it, when they may
be destroyed _en masse_ by hot water; or, disappointed, they abandon
the premises as a barren spot, and seek some other more promising
locality for exploration. When they are fairly in we give up the house,
and try to await with patience their pleasure, thankful, indeed, if
permitted to remain within the narrow limits of our beds or chairs.
They are decidedly carnivorous in their propensities. Fresh meat of all
kinds is their favorite food; fresh oils also they love, especially
that of _Elais guiniensis_, either in the fruit or expressed. Under my
observation they pass by milk, sugar and pastry of all kinds, also salt
meat; the latter, when boiled, they have eaten, but not with the zest
of fresh. It is an incorrect statement, often made, that _they devour
everything eatable_ by us in our houses; there are many articles which
form an exception. If a heap of rubbish comes within their route, they
invariably explore it, when larvae and insects of all orders are borne
off in triumph--especially the former.”

Sometimes, instead of camping in shelters on the ground, these ants
climb up into a tree and hang together in a cluster like a swarm of
bees. Savage reports a colony suspended from a low tree: “From the
lower limbs (four feet high) were festoons or lines of the size of
a man’s thumb, reaching to the plants and ground below, consisting
entirely of these insects; others were ascending and descending upon
them, thus holding free and ready communication with the lower and
upper portions of this dense mass. One of these festoons I saw in the
act of formation; it was a good way advanced when first observed:
ant after ant coming down from above, extending their long limbs and
opening wide their jaws, gradually lengthened out the living chain till
it touched the broad leaf of a _Canna coccinea_ below. It now swung to
and fro in the wind, the terminal ant meanwhile endeavoring to attach
it by his jaws and legs to the leaf; not succeeding, another ant of the
same class (the very largest) was seen to ascend the plant, and, fixing
his hind legs with the apex of the abdomen firmly to the leaf under the
vibrating column, then reaching with his fore-legs and opening wide his
jaws, closed in with his companion above, and thus completed the most
curious ladder in the world.”

Similar chains are used in bridging little rills or even small brooks,
but when a real flood occurs a different procedure is adopted. In this
case they cling together so as to form a large ball, with the eggs and
young in the center, and float away upon the water until a safe landing
can be effected.

There are several kinds of legionary and driver ants in America; some
species have been found as far north as Texas and even Colorado, but
most of them are confined to the tropics. These ants usually do not
spend all of their time on the march, but have permanent nests, from
which they sally out at intervals on foraging expeditions. Belt offers
a graphic description of the sortie of a colony in Brazil: “One of the
smaller species (_Eciton praedator_) used occasionally to visit our
house, swarm over floors and walls, searching every cranny, and driving
out the cockroaches and spiders, many of which were caught, pulled or
bitten to pieces, and carried off.... I saw many large armies of this,
or a closely allied species, in the forest. My attention was generally
first called to them by the twittering of some small birds, belonging
to several different species, that followed the ants in the woods. On
approaching to ascertain the cause of the disturbance, a dense body of
the ants, three or four yards wide, and so numerous as to blacken the
ground, would be seen moving rapidly in one direction, examining every
cranny, and underneath every fallen leaf. On the flanks, and in advance
of the main body, smaller columns would be pushed out. These smaller
columns would generally first flush the cockroaches, grasshoppers and
spiders. The pursued insects would rapidly make off, but many, in
their confusion and terror, would bound right into the midst of the
main body of ants.... The greatest catch of the ants was, however, when
they got amongst some fallen brushwood. The cockroaches, spiders and
other insects, instead of running right away, would ascend the fallen
branches and remain there, whilst the host of ants were occupying
all of the ground below. By and by up would come some of the ants,
following every branch, and driving their prey before them to the ends
of the small twigs, when nothing remained for them but to leap, and
they would alight in the very midst of their foes, with the result of
being certainly caught and pulled to pieces. Many of the spiders would
escape by hanging suspended by a thread of silk from the branches, safe
from the foes that swarmed both above and below.”

[Illustration: Fig. V. Legionary ants attacking a snake.]

Some of the Brazilian species are more nomadic in their habits.
Belt says: “I think _Eciton hamata_ does not stay more than four or
five days in one place. I have sometimes come across the migratory
columns. They may easily be known by all the common workers moving in
one direction, many of them carrying the larvae and pupae carefully
in their jaws. Here and there one of the light-colored officers
moves backwards and forwards directing the columns. Such a column is
of enormous length, and contains many thousands, if not millions,
of individuals. I have sometimes followed them up for two or three
hundred yards without getting to the end.... They make their temporary
habitation in hollow trees, and sometimes underneath large fallen
trunks that offer suitable hollows. A nest I came across in the latter
situation was open at one side, and the ants were clustered together
in a dense mass, like a great swarm of bees, hanging from the roof
but reaching to the ground below. Their innumerable long legs looked
like brown threads binding together the mass, which must have been at
least a cubic yard in bulk, and contained hundreds of thousands of
individuals, although many columns were outside, some bringing in the
pupae of ants, others the legs and dissected bodies of insects. I was
surprised to see in this living nest tubular passages leading down
into the center of the mass, kept open just as if it had been formed
of inorganic material. Down these holes the ants who were bringing the
booty passed with their prey. I thrust a long stick down to the center
of the cluster and brought out clinging to it many ants holding larvae
and pupae, which were probably kept warm by the crowding together of
the ants. Besides the common dark-colored workers and light-colored
officers, I saw there many still larger individuals with enormous jaws.
These they go about holding wide open in a threatening manner, and I
found, contrary to my expectation, that they could give a severe bite
with them, and that it was difficult to withdraw the jaws from the
skin.”

Sumichrast, who studied some of the Mexican legionaries in 1863,
noted many seemingly aimless migrations, “which they undertake at
undetermined epochs, but in relation, it appears to me, with the
atmospheric changes. What traveler, passing over the _tierra caliente_,
has not encountered the phalanxes of _tepeguas_ upon the paths of the
primitive forests? What inhabitant of these countries has not, at least
once, been unpleasantly torn from the arms of sleep by the invasion
of his domicile by a black army of _soldados_?... Besides the changes
of domicile which are so generally in relation with the atmospheric
variation as to serve as a rule to the inhabitants of the country,
the _Eciton_ devotes itself every season to excursions for pillage,
destined to supply the larvae with nourishment. Nothing is more
curious than these _battues_ executed by an entire population. Over an
extent of many square meters, the soil literally disappears under the
agglomeration of their little black bodies. No apparent order reigns in
the mass of the army, but behind this many lines or columns of laggards
press on to rejoin it. The insects concealed under the dry leaves and
the trunks of fallen trees fly on all sides before this phalanx of
pitiless hunters, but, blinded by fright, they fall back among their
persecutors and are seized and dispatched in the twinkling of an eye.
Grasshoppers, in spite of the advantage given them by their power of
leaping, hardly escape more easily. As soon as they are taken, the
_Eciton_ tears off the hinder feet and all resistance becomes useless.”

The same author describes with some feeling their habit of invading
houses. “These visits ordinarily take place at the beginning of the
rainy season, and almost always during the night. The expeditionary
army penetrates the habitation which it proposes to visit at many
points at once, and for this purpose divides itself into many columns
of attack. One is apprised very soon of their arrival by the household
commotion among the parasitic animals. The rats, the spiders, the
cockroaches, abandon their retreats and seek to escape from the attacks
of the ants by flight. Alimentary substances the _soldados_ hold in
no esteem, and they disdain even sugary things, to which the ants in
general are so partial. Dead insects even do not seem to invite their
covetousness. It has often happened to me to be obliged to abandon
my abode, without having time to carry away my collection, to which
they have never done the least injury. The trouble occasioned by these
insects in entering houses is more than compensated by the expeditious
manner in which they purge them of vermin, and in this view their visit
is an actual benefit.”

As these ants are usually quite blind and their movements are directed
(so far as we can tell) by the sense of smell and contact alone, it
is quite remarkable that they are able to move about so readily, and
become familiar with their surroundings in less time than their seeing
relatives. Forel wrote in 1899: “Throw a handful of _Ecitons_ with
their larvae on a spot with which they are absolutely unacquainted. In
such circumstances other ants scatter about in disorder and require an
hour or more to assemble and bring their brood together and especially
to become acquainted with their environment, but the _Ecitons_ do this
at once. In five minutes they have formed distinct files which no
longer disintegrate. They carry their larvae and pupae, marching in a
straight path, palpating the ground with their antennae and exploring
all the holes and crevices till they find a suitable retreat and enter
it with surprising order and promptitude. The workers follow one
another as if at a word of command, and in a very short time all are in
safety.”




CHAPTER VIII

THE RED SLAVE MAKERS


The European ant known as _Formica sanguinea_ is blood-red in color,
and is one of the most industrious, versatile, and belligerent insects
known to man. This species, according to Wheeler, “assails any intruder
with its mandibles, simultaneously turning the tip of its gaster
forward and injecting formic acid into the wound.”

Although _sanguinea_ is widely known as a slave-holding species, it
is by no means wholly dependent upon its slaves, but is quite able
to dig its own nest, gather food and rear young without the aid of
any slaves at all. “There is,” said Wheeler, “nothing to show that
the slaves contribute anything more to the communal activities than
would be contributed by an equal number of small _sanguinea_ workers.”
Many observers have reported slaveless colonies of _sanguinea_ which
seemed to be flourishing, and Wasmann found that the youngest colonies
contain, as a rule, more slaves than the older nests. He also reported
an inverse ratio between the number of slaves and the size of the
colony, some of the very largest being practically slaveless.

The slave-hunting expeditions of the _sanguinea_ are said to occur
only two or three times a year, and the general procedure is described
by Wheeler as follows: “The army of workers usually starts out in the
morning and returns in the afternoon, but this depends on the distance
of the _sanguinea_ nest from the nest to be plundered. Sometimes the
slavemakers postpone their sorties till three or four o’clock in the
afternoon. On rare occasions they may pillage two different colonies in
succession before going home. The _sanguinea_ army leaves its nest in
a straggling, open phalanx sometimes a few meters broad and often in
several companies or detachments. These move to the nest to be pillaged
over the directest route permitted by the often numerous obstacles in
their path. As the forefront of the army is not headed by one or a few
workers that might serve as guides, but is continually changing, some
dropping back while others move forward to take their places, it is
not easy to understand how the whole body is able to go so directly to
the nest of the slave species, especially when this nest is situated,
as is often the case, at a distance of fifty or a hundred meters. We
must suppose that the colony has acquired a knowledge of the precise
location of the various nests of the slave species within an area of
a hundred meters or more of its own nest. This knowledge is probably
acquired by scouts leaving the nest singly and from time to time for a
period of several weeks, and these scouts must be sufficiently numerous
to determine the movements of the whole worker body when it leaves the
nest. This presupposes not only a high development of memory, but some
form of communication, for the nest attacked is usually one of many
lying in different directions from the _sanguinea_ nest.

“When the first workers arrive at the nest to be pillaged, they do
not enter at once, but surround it and wait for the other detachments
to arrive. In the meantime the _fusca_ or _rufibarbis_ scent their
approaching foes and either prepare to defend their nest or seize their
young and try to break through the cordon of _sanguinea_ and escape.
They scramble up the grass-blades with their larvae and pupae in their
jaws or make off on the ground. The sanguinary ants, however, intercept
them, snatch away their charges, and begin to pour into the entrance of
the nest. Soon they issue forth one by one with the remaining larvae
and pupae and start for home. They turn and kill the workers of the
slave-species only when these offer hostile resistance. The troop of
cocoon-laden _sanguinea_ straggle back to their nest, while the bereft
ants slowly enter their pillaged formicary and take up the nurture of
the few remaining young or await the appearance of future broods.

“Forel is of the opinion that many of the young brought home by the
sanguinea are eaten, for the number of those which eventually hatch and
become auxiliaries is very small compared with the number pillaged
during the course of the summer. Wasmann believes, however, that the
forays take place for the specific purpose of obtaining young to rear.
This seems to be disproved by the fact that even small _sanguinea_
colonies are quite able to get along without slaves and by the
insignificant number of these individuals in many nests. Darwin has
interpreted the surviving and adopted workers as a kind of by-product,
or as representing food which the ants failed to eat at the proper
time, and such they would appear to be in the adult colony, though, as
we shall see, they have an additional significance as the result of an
instinct inherited by the _sanguinea_ workers from their queen. That
the foray is, to some extent at least, due to the promptings of hunger,
seems to be shown by the fact that _sanguinea_ sometimes plunders the
nests of ants which it could not adopt as slaves.”

Wasmann describes the military expeditions of the so-called sanguine
slavemakers (_F. sanguinea_), which generally hunt in companies of
from twenty to fifty workers, “with the purpose not only of stealing
the neuter pupae of the slave species, but often also of pillaging
the nests of smaller ants belonging to the genus _Lasius_, the
larvae, pupae and winged individuals of which are carried off to be
devoured. During the time of the nuptial flight of _Lasius niger_, many
_sanguinea_ colonies are hunting in the vicinity of their nest for the
heavy _Lasius_ females which drop to the ground. Then either singly
or with united forces these robbers pull their victims into their
strongholds, where they are mercilessly slaughtered. On the afternoon
of August 24, 1888, I witnessed such a typical hunting expedition of
several _sanguinea_ colonies near Exaten, Holland, on the outskirts of
a fir plantation. The road passing the nests was covered far and wide
with _sanguineas_ rushing upon every _Lasius_ female that dropped from
the air, as upon a welcome booty. Within the space of an hour I counted
more than one hundred females of _Lasius niger_ that fell victims to
the hunters.”

There are several species and sub-species of _sanguinea_ in the United
States, and the habits of these differ in several particulars from
those of their European relatives. Wheeler reports that although he
has found plenty of slaveless colonies, most nests contain slaves in
much greater number than do similar colonies in Europe. He thinks this
due in part to the fact that the American species make more frequent
raids, and partly also because the species chosen as slaves are “much
more cowardly and docile” than the victims of the slave-hunters of
the Old World. The actual tactics employed in the raids do not differ
essentially from those of the European species.

It was long supposed that new colonies of the _sanguinea_ were founded
in this wise: When the queen descends from her nuptial flight she
either brings up a brood of her own like many common ants, or she is
adopted into a nest of one of the slave species. On either of these
suppositions it is difficult to explain how the slave-making instincts
could be transmitted to the workers, because the latter have no
offspring and the queen was supposed to lack the slaving instincts. In
1906, Wheeler cleared the matter up by introducing a _sanguinea_ queen
into a nest containing workers, larvae, and cocoons of one of the slave
species. She was immediately attacked, but beat off her assailants,
killed a number of them, and captured a large number of cocoons, which
she carried into a separate chamber and defended against all comers.
Here she waited until the workers emerged from the captured cocoons;
these workers, of course, attached themselves to her and soon gained
possession of the whole nest. This experiment shows clearly that the
_sanguinea_ queen really possesses all the slave-making tendencies
exhibited by the workers in their raiding, and solves the problem of
the inheritance of these instincts.




CHAPTER IX

THE AMAZONS AND THEIR SLAVES


Another type of slave-owning ants is represented by the genus
_Polyergus_, found in both Europe and North America, and known as
amazons. Slavery among the amazons is a very different thing from
the casual master-servant relationship found in the various species
of sanguinary ants. The _sanguinea_ are quite able to build nests,
gather food, and rear their young unaided by slave labor, and slaveless
colonies are not at all uncommon, but the amazons are absolutely
dependent upon their slaves, and no amazon colony could exist without
them. As Wheeler says, the amazons “are even incapable of obtaining
their own food, although they may lap up water or liquid food when
this happens to come in contact with their short tongues. For the
essentials of food, lodging and education they are wholly dependent on
the slaves hatched from worker cocoons that they have pillaged from
alien colonies. Apart from these slaves they are quite unable to live,
and hence are always found in mixed colonies inhabiting nests whose
architecture throughout is that of the slave species. Thus the amazons
display two contrasting sets of instincts. While in the home they sit
about in stolid idleness or pass the long hours begging the slaves for
food or cleaning themselves and burnishing their ruddy armor, but when
outside the nest on one of their predatory expeditions they display
a dazzling courage and capacity for concerted action compared with
which the raids of _sanguinea_ resemble the clumsy efforts of a lot of
untrained militia. The amazons may, therefore, be said to represent
a more specialized and perfected stage of _dulosis_ than that of the
sanguinary ants. In attaining to this stage, however, they have become
irrevocably dependent and parasitic.”

The same author describes a slave-hunting foray of the European
species. “The ants leave the nest very suddenly and assemble about
the entrance if they are not, as sometimes happens, pulled back and
restrained by their slaves. Then they move out in a compact column
with feverish haste, sometimes, according to Forel, at the rate of a
meter in 33 seconds, or 3 cm. per second. On reaching the nest to be
pillaged, they do not hesitate like _sanguinea_ but pour into it at
once in a body, seize the brood, rush out again and make for home.
When attacked by the slave species they pierce the heads or thoraces of
their opponents and often kill them in considerable numbers. The return
to the nest with the booty is usually made more leisurely and in less
serried ranks. The observer of one of these forays cannot fail to be
impressed with the marvelous precision of its execution. Although the
ants may occasionally lose their way and have to retrace their steps or
start off in a different direction, they usually make straight for the
nest to be plundered. They must, therefore, like _sanguinea_, possess a
keen sense and memory of locality. There can be little doubt that they
often leave the nest singly and make a careful reconnoissance of the
slave colonies in the vicinity.”

One can hardly believe that as soon as the fighting is over these
warriors relapse into their accustomed lethargy, and are fed and cared
for by their slaves, who often prevent them from leaving the nest,
and sometimes, when they have wandered away, pick them up bodily and
carry them home by main strength. When a colony moves to a new home
the whole enterprise is left to the slaves, who choose and prepare the
new nesting site, and carry the warriors to it. In the case of the
_sanguinea_ it will be remembered that it is the masters who carry the
slaves on these occasions.

An American amazon which has been the subject of considerable study is
_Polyergus breviceps_, found in the mountainous regions of Colorado
and New Mexico. This species is very striking in appearance, the
worker and queen being of a rich purplish-red color, while the male
is jet-black with white wings. A peculiar feature of the _breviceps’_
raiding parties is that there are no casualties on either side. The
slave species offer no real resistance, and the amazons simply put them
gently to one side, take their larvae and pupae, and go their way.

We do not know exactly how new amazon colonies are established. Forel,
Wasmann and Viehmeyer have agreed that the queen lacks the domestic
instinct, and therefore the new colony must be founded by the slave
species, which cares for the amazon females. It has been shown that the
adoption occurs readily enough in artificial nests. Some experiments by
Wheeler gave rather conflicting results, and he closes his discussion
of the matter by saying: “It will be necessary, therefore, to study
this question further before making definite statements in regard to
the method employed by our American amazons in establishing colonies.”




CHAPTER X

DAIRIES AND GUESTS


The peculiar symbiotic relations between ants and aphids is worth a
brief description. The aphids or plant-lice live in colonies upon
certain plants, and feed upon juices which they suck from the foliage.
The liquid excrement of these insects is sweet, and a surprisingly
large amount is voided--Bŭsgen found that the maple aphid produces
as many as forty-eight drops in twenty-four hours. This substance is
sometimes so abundant that it covers the leaves and even drips down to
the ground; it is known as honeydew, and some rustics still believe
that it somehow falls from heaven. The ants are very fond of this
honeydew, and some species live upon it almost exclusively at certain
seasons, and locate their nests always near good aphid-pastures. The
ants never kill and eat aphids as they do other insects, but protect
them against their enemies. They even carry them about from one pasture
to another, and some species build little sheds and corrals in which
their aphids are confined just as we confine cattle. Sometimes the ants
simply lap up the honeydew as it falls upon the leaves, but in most
cases they _milk_ the aphids by gently stroking them with the antennae,
which causes the emission of a drop of the sweet liquid. Some kinds of
aphids have developed a circle of stiff hairs around the anal opening,
which thus retains the honeydew till the ant comes for it. Not only do
the ants care for and milk the adult aphids, but they rear them from
the eggs. Huber, Lubbock and others have seen ants collecting aphid
eggs in the Autumn, and it has been found that these eggs are stored in
the nest until they hatch, when the young plant-lice are carried out
and placed on a suitable food-plant. On cold or rainy days they are
taken back into the nest; when the weather moderates the ants carry
them out to pasture again.

The scale-insects and mealy-bugs (_Coccidae_) also produce honeydew,
and are visited by the ants precisely as the aphids are. The _manna_
of the Biblical story, according to Wheeler, “is now known to be the
honeydew of one of these insects (_Gossyparia mannifera_) which lives
on the tamarisk. This excretion is still called _man_ by the Arabs who
use it as food.” Forel, Cockerell and Wheeler have seen ants tending
great herds of coccids, and a few of these insects are found in many
nests.

Several kinds of tree-hoppers bear a similar relation to ants. Bare,
who studied these matters in Argentina, “watched the larvae of various
species of _Centrotus_ being assiduously attended by ants. The larvae
are gregarious, frequenting the succulent shoots of plants, and have an
extensile organ at the extremity of the body, from which the coveted
fluid is emitted.” Wheeler observed whole colonies of ants herding
leaf-hoppers in Colorado, and reports that these novel milk-cows
“responded to the antennal caresses of the ants in precisely the same
manner as the plant-lice and scale-insects.” Some ants confine their
tree-hoppers in sheds and shelters similar to those used for the aphids.

The relationship of ants to certain small caterpillars (the larvae
of some of the _Lycaenid_ butterflies) has been known for a long
time. These little caterpillars, when caressed on the posterior
end by the antennae of the ants, give up a drop of sweet liquid,
doubtless very similar to that produced by the aphids and coccids.
These larvae are often found in the ants’ nests, and some of the newly
emerged butterflies have been seen to come out of the ant-hills.
It is said that the ants protect the caterpillars from the attacks
of hymenopterous parasites, and De Niceville is authority for the
statement that the butterfly will not lay her eggs when there are no
ants about: “If the right plant has no ants, or the ants on that plant
are not the right species, the butterfly will lay no eggs on that
plant. Some larvae will certainly not live without the ants, and many
larvae are extremely uncomfortable when brought up away from their
hosts or masters.”

[Illustration: Fig. VI. A small myrmecophilous cricket (_Myrmecophila
nebrascensis_) gnawing at the tibia of the Texan harvester-ant. (After
Wheeler.)]

Besides the ants’ relationship with the insects which produce sweet
substances, there are symbiotic relations of a very different type with
a group of insects known as _myrmecophiles_--ant-guests. These insects,
at one stage or another, live in the ant-hills. At least fifteen
hundred species of ant-guests are known, and Escherich estimates that
there must be at least three thousand altogether. Wheeler thinks that
even this estimate is probably too low. At least a thousand of the
known species are beetles, and most of the rest are insects of one kind
or another, but there are about sixty arachnids and a few crustaceans.

Some of the myrmecophiles are not _friends of ants_ as the name
implies, but mere interlopers--scavengers, robbers and assassins.
There are a number of small beetles which live in the less frequented
galleries of the nest, eat dead ants and brood, kill ailing or crippled
ants, and even attack healthy adults when they catch them alone or
at some disadvantage. Some of these beetles resemble ants in general
appearance, a mimicry which is doubtless of considerable value to
them. The ants kill these pests whenever they can, but many are
protected by their ability to emit an evil-smelling substance which
puts the ants to flight. Others will be killed at once if confined in a
small chamber with a few ants, but in a large nest are able to escape
by reason of their agility.

Another class of myrmecophiles, known as _synoeketes_, or tolerated
guests, live in the ant-hills without attracting any great attention,
being treated with contemptuous indifference by their hosts. The larvae
of certain moths and flies, a large number of beetles, and numerous
other insects are of this class, and feed largely upon the refuse of
the kitchen-middens. Wasmann has studied a group of beetles which
live with the nomadic Doryline ants. These camp-followers mimic the
legionaries, and march along in their columns apparently unnoticed,
being allowed to share the prey taken by the blind warriors. Other
beetles live in the nests of the _sanguinea_, and feed largely upon the
tiny parasites from the bodies of their hosts. Certain minute wingless
crickets are very abundant in many nests; they are seen to lick the
bodies of the ants, and it is supposed that they live upon some
cutaneous secretion.

The insect called _Attaphila_ is a sort of miniature cockroach, which
lives with the fungus growing _Attii_, and is, according to Wheeler,
the only insect known to be on intimate terms with these ants. A
peculiar thing about the _Attaphila_ is that the last joint of the
antennae is nearly always bitten off. This insect was formerly
supposed to feed on fungus, but has since been found to lick the
surface secretions from the ants’ bodies. A little beetle called
_Oxysoma oberthueri_ is very like _Attaphila_ in its habits, “mounting
the bodies of its host and licking or shampooing them with great
eagerness.”

Very different from the furtive, barely tolerated myrmecophiles
described above are the three or four hundred species known as true
guests, which, to quote Wheeler again, “are no longer content to be
treated with animosity or indifference, but have acquired more intimate
and even friendly relations with the ants. These true guests are not,
therefore, to be found skulking in the unfrequented galleries of the
nest, or suspiciously dodging about among the ants, but live in their
very midst with an air of calm assurance, if not of proprietorship.”
Among these are many beetles bearing tufts of hair which produce some
aromatic secretion very pleasing to the ants. The ants rush to lick the
odorous tufts, are caressed by the peculiar antennae of the beetle,
and feed the latter with regurgitated food. Many of these beetles are
cleaned and shampooed by the ants, are often carried about, and favored
in other ways, despite the fact that they sometimes devour the ant
brood. Some of the smaller species are totally blind, and are permitted
to ride about on the ants’ backs for hours at a time.

Another sort of guest is the little mite called _Antennophorus_,
which Janet has found in the nests of several European ants. These
mites attach themselves firmly to the body of their host, and it is
interesting to note that no matter how many are present on a single
ant, they are always so placed that the weight is properly distributed,
and the host’s progress not interfered with. These creatures remind
one of the ticks found on higher animals like dogs, but they are not
parasites in the sense that ticks are--they do not suck the ant’s
blood, but reach out and snatch their nutriment from the drops of
regurgitated food as they pass from one ant to another.

[Illustration: Fig. VII. Showing two minute myrmecophilous beetles
(_Oxysoma oberthueri_) feeding on the surface secretions of an ant.
(Adapted from Escherich).]

The ants do not bother _Antennophorus_ much, but there is another mite
called _Cillibano_ which is a true blood-sucker, and which they seize
and tear to pieces whenever they can. A little blue fly (_Orasema
viridis_) is common in the nests of several Texan and Mexican ants; its
larvae attach themselves to the ant larvae and live as parasites. Both
the larvae and the adult, however, are fed and fondled by the ants.

Besides these external parasites there are many grubs and worms which
live inside the body of the ant, and are comparable to the pin-worms
and tapeworms which dwell in the human intestine. These creatures have
not been studied extensively, however, and very little is known of
their habits and metamorphosis.

       *       *       *       *       *




Transcriber’s note


Minor punctuation errors have been changed without notice. Italization
was standardized.

Illustrations tags have been moved so they do not break up the
paragraphs.

Spelling was retained as in the original except for the following
changes:

  Page 7: “female during copulalation”      “female during copulation”
  Page 17: “the arangement of larvae”       “the arrangement of larvae”
  Page 18: “the ant’s tevelopment”          “the ant’s development”
  Page 29: “habits of the Attiine”          “habits of the Attine”
  Page 29: “besides the Attiien ants”       “besides the Attine ants”
  Page 44: “itself every reason to”         “itself every season to”
  Page 50: “of several _sanquinea_”         “of several _sanguinea_”
  Page 51: “the _sanquinea_ queen”          “the _sanguinea_ queen”
  Page 55: “is known as honey dew”          “is known as honeydew”
  Page 55: “honey dew, and some species”    “honeydew, and some species”




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