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IMOMLR AAMMOMS

REV. E. WASN ANN “ee

SYCHOLOG GYorANTS
oO} ae |

THE LIBRARY
OF
THE UNIVERSITY
OF CALIFORNIA
LOS ANGELES

John M. Mehutens

COMPARATIVE STUDIES

IN THE

Psychology of Ants and of
Higher Animals.

ERIC WASMANN, S. J.

Plus enim formicularum et apicularum opera
stupemus quam immensa corpora balaenarum,
(S. Augustine, De Civit, Dei, |. 22, c. 24, n. 5.)

Authorized English Version of the second German Edition.
Enlarged and revised by the Author.

St. Louis, Mo., AND FREIBURG, (BADEN),
Published by B. HERDER.
1905.
LONDON AND EDINBURGH: SANDS & CO.

— BECKTOLD—
PRINTING AND BOOK MFG. CO.
ST. LOUIS, MO.

PREFACE TO THE FIRST EDITION.

OME time ago we published an essay entitled

“Instinct and Intelligence in the Animal King-
dom,” examining in detail the concepts of instinct
and intelligence, with their application to animals.
The discussion showed that intelligence is the spiritual
power of abstraction, and not the mere faculty of
forming complex sense-representations; for the laws
of association in sense-perceptions belong to the sphere
of instinctive sensitive life and not to spiritual intelli-
gence. Now, what modern animal psychology terms
“intelligence of animals,” is nothing but inborn
instinct, raised to a higher level of perfection by the
individual’s sensuous experience. This, in its turn,
is based on the very same laws of association of sense-
representations. Hence, there is no reason for ascrib-
ing to animals intelligence in the strict sense. Indeed,
our reasoning led us to take a further step, and we
proved that animals have no intelligence at all. If
they were gifted with a spiritual power of abstraction,
it would necessarily be manifested in their outward
actions, especially by the formation of an arbitrary
phonetic or graphic language. Animals, however,
have no language; hence, they have no intelligence.

Besides, we have shown in the same essay that
the manifestations of the psychic life, both of higher
and of lower animals, are to be judged according to
one and the same critical standard. The anatomical

iii

Preface to the First Edition.

difference, that exists between the sense organs and
the nervous system of Arthropods on the one hand and
of Vertebrates on the other, is not a sufficient a priori
reason for ascribing intelligence to the latter only and
denying it to the former. The proof of these asser-
tions forms the groundwork of the present essay. We
shall compare more at length the psychic life of the
most “intelligent” Arthropods, namely the ants, with
that of the higher Vertebrates and of man. From this
discussion we shall learn, whether the “missing link,”’
with which modern evolutionists hope to bridge over
the chasm between the instinct of animals and the
spiritual soul of man, is to be looked for in ants or
in the higher Vertebrates, or whether, in fact, it exists
at all. Biologists will be pleased to find that the
present essay contains many new observations on the
habits of ants and their guests.

PREFACE TO THE SECOND EDITION.

HE numerous observations of modern scientists,

illustrating the relations between the psychic life
of ants and of higher animals, have been extensively
utilized in preparing this second edition. We have
paid due regard to the observations and experiments,
published since 1897, on the differentiation of castes
in bee-hives. Besides, we have turned to account the
results of a statistical chart now completed, extending
over five years and comprising all the colonies of
Formica sanguinea in the neighborhood of Exaten,
Holland. This ant is the most interesting of all
European species. Thus we are able to publish many
new facts of interest in scientific biology regarding
the slave-making habits of this ant, its methods of
nest construction, its relationship to its guest
Lomechusa, and the influence of the latter in the
differentiation of castes in ant communities. Finally,
two additional illustrations of Lomechusa strumosa
and of its larva have been added in the text.

TRANSLATOR’S PREFACE.

TASMANN’S “Instinct and Intelligence in the
Animal Kingdom,” which appeared recently in
an English dress (Herder, St. Louis, Mo.), was so
favorably received, that it has been thought advisable
to follow it up with this translation of another essay
by the same author. These two books supplement
each other, as may be gathered from the frequent
cross references they contain, and, more especially,
from the close relationship of the subjects of which
they treat.

The best recommendation of Wasmann’s biological
and psychological essays is given in the following
lines of W. M. Wheeler, Prof. of the University of
Texas:1 “Wasmann in his numerous writings has
undoubtedly done much, at least in Germafiy, towards
the exposure of this pseudo-psychology (of Brehm,
Buechner and others) and a more rational conception
of ant behavior. His long familiarity with these
animals and their guests has given him a singularly
lucid insight into their activities. My own more
limited observations on our North American species
lead me to agree with him so far as the facts are con-
cerned, and many of the inferences which he has
drawn from them.” As to his additional remark:
“T am constrained to say, however, that I cannot adopt

1) “The Compound and Mixed Nests of American Ants,” in
“American Naturalist,” Vol. XXV, 1901, p. 808.

Translator’s Preface.

either his psychological definitions or his psychogenetic
reservations,”—we call the critic’s attention to the end
of the fourth chapter of “Instinct and Intelligence in
the Animal Kingdom,” where Prof. Wheeler’s objec-
tions have been answered.

In order to make the English translation more
valuable for North America, the author has kindly
added a series of notes and observations on the ant
fauna of the United States. He has added, moreover,
the figures representing the North American form of
Formica sanguinea and that of its guest Xenodusa
cava. The present work, therefore, is more than a
translation; it may be called a new edition, revised
and enlarged by Father Wasmann.

Canisius College, Buffalo, N. Y.

vii

CONTENTS.

PAGE.

Pretace to the first: edition... 22032) See er
Preface to the second edition .........cccceccccccces
WERORERIOES “BERTR CE eS eg se al Sans a eee tas

INTRODUCTION.

Some ancient and modern views of the psychic life of
ants and of higher animals......3...:.-.ccceccses

CHAPTER I.
COMMUNITY LIFE IN THE ANIMAL KINGDOM.

1. A general view of the forms of animal communities.
Different degrees of community life in the animal kingdom.
Communities of bees and ants compared. Ant states, the
most perfect animal societies, both simple and complex.
2. The social foundations of ant states............. eiese'
Polymorphism, the organic foundation of ant societies.
Psychic ties in ant colonies. It is untenable to identify the
human “state”? with that of ants. ‘‘Automatic” instincts
cannot sufficiently account for the latter.
3. The communities of higher animals compared with
SMOBE OF ANUS vc ok Fn PES i oo a
Mutual warning of danger. Sentinels. Mutual “charity.”
Nursing of the sick. Co-operation and division of labor.
Common defense. ‘‘Fidelity and obedience” in animal so-
cieties. Brief summary.

CHAPTER II.
WARFARE AND SLAVERY IN THE ANIMAL KINGDOM.

1. Wars among higher animals...............ceceees

Fables and stories of wars between troops of apes. The

latter use no weapons or tools,
viii

iii
a
vi

Contents.

PAGE.

2. The war-like expeditions of the Amazon ant and the

FOCmiea SOUQUINEE Gi A eins oh i a h

Their tactics, the “humanity” of the victors. The military
skill of the Amazon ants; their inability to feed themselves.
The military tactics of the sanguine slavemaker (Formica
sanguinea) discussed on psychological grounds.

3. The pretended “automatism” in the psychic life of ants

The individuals of one and the same colony act differently

in combat. Acquired individual habits. Persecution of toler-

ated or even of true guests. Ants tamable. Psychic influence

of the numbers of a colony on their courage in combat.
Marked ‘‘heroism” of individuals. Martial sports?

4. The slave-making instincts of Formica sanguinea......
“Tradition and instruction” in insect communities. Experi-
ments made with “self-taught” ants and bees. Seemingly
intelligent plasticity of the instinct of slavery in Formica
sanguinea, both with regard to the number and to the species
of its slaves. On the nature of slavery in ants. The correct
explanation is neither anthropomorphism nor mechanic automa-
tism, but something between the two.

* Other wats atid alliances of 'Shts oc cio. ccs dae ce oe vanes
Fights between neighboring colonies. Alliances between
hostile colonies, and their psychological explanation. Summary.

CHAPTER III.
ARCHITECTURE IN THE ANIMAL KINGDOM.

1. A general survey of the architecture of animals........
2. The nests of ants. ve ota elek ealee age eae arse
Various and wbincaee: forms of their nests. Nests of dif-
ferent ant species. Psychological explanation of this difference.
3. The nests of Formica sanguined........ cece ceeceeee ;
Variety and plasticity in their forms. Variable panes of
the nests of a colony. Periodic changes of nests; different
places of residence in different seasons. These phenomena
psychologically explained. Power of adapting the construc-
tion of their nests to the surroundings, in order to secure
them against hostile inroads.
_ 4. How do ants build their nests?................. Reni.
No rigid system of co-operation; seemingly intelligent self-
determination. Suitable consideration of the different con-
ditions of- temperature and moisture. Architecture of birds
compared with that of ants; with that of mammals, in par-

ticular of beavers.
ix

103

Contents.

PAGE.

3. Other purposes, for which ants employ their architec-
fibals: Ska ce ese RAN ere een Paes eect

Stations, roads, galleries, stables for aphides, granaries,
ramparts, etc. The grave-yards of ants in fiction and in
reality.

6. Is the building activity of ants guided by intelligence?..

Ants using their larvae as spinning-wheels. The “bridge
building” of ants according to W. Marshall and Buechner.
Experiments of Lubbock and Bethe. My own experiments.
Results summed up.

CHAPTER IV.
BREEDING AND NURSING IN THE ANIMAL KINGDOM.

1. A general view of the breeding instincts of animals ...

Breeding and family life in animals with reference tc the
preservation of species. This relationship psychologically
explained. Breeding in social insects. The breeding instinct
of ants dependent on the laws of organic development.

2. Care of the young ‘among ants. 0.0.0.0 0s ce Ae

Psychic characteristics of their breeding instinct and of its
different manifestations. The differentiation of castes in
bees and ants influenced by education. Intermediate forms
between females and workers in ants, and their probable
explanation. Pseudogynes and their relation to Lomechusa.
Attachment of ants to their brood. Fond “mothers” and
“aunts.”

3. Adoption instincts in the animal kingdom...... canes

Their frequent occurrence. Psychological explanation. Ants
nursing Lomechusa-larvae. Different treatment of these
larvae, both in normal sanguinea colonies and in such as
contain pseudogynes. W. Marshall and the nursing of eggs
of aphides by ants; “intelligent’? foresight of the future;
comparison with the nursing of Lomechusa-larvae. Adoption
instincts in birds and mammals. Results summed up; con-
clusions,

Conclusion. 30334 42.3. SD Gee TY & asa eo oe ee ares ;

The automatism and the plasticity of instinct both in ants
and higher animals reviewed. Man the only being in the
visible creation gifted with reason. Humanization of animals
is unscientific, and injurious to the moral order of human
society,

153

173

192

INTRODUCTION.

VEN in ancient times, observers of animal life

noticed that bodily size and psychic perfection
are not always in direct proportion, but that the
reverse is not unfrequently the case. Thus Aristotle’
declared that keenness of perception (rqv Tijs Siavoias
dkpi(Beav) was often more manifest in smaller
than in larger animals. Nor did it escape the great
Stagirite, who was not only a logical thinker, but also
a skilful observer, that many animals of low rank
in the zoological scale were endowed, in some way,
with a higher psychic life than the highest mammals,
so much so, that its manifestations could be com-
pared with human institutions only. He mentions,
especially, ants and bees among those “bloodless”
animals which possess a more intellectual soul than
_ many animals of the other kind.?, The same thought
was expressed by St. Augustine, one of the loftiest
Christian minds, in the following terms: “We admire
the works of the tiny ants and bees more than the
bulky forms of whales.”’* And a distinguished modern
naturalist, Emil Dubois-Reymond, has acknowledged

1) “Hist. animal.,” 1. 9, c..7 (Becker I, 612).

2) “De partib. animal.,” 1. 2, c. 4 (Becker I, 650). Aristotle’s
division of animals into those with red blood and those with colorless
blood in reality coincides with that of Vertebrates and Non-Vertebrates. |
He uses the term “Bloodless Animals” for those which have no red
blood.

8) “De civ. Dei,” 1. 22, c. 24, n. 5 (Migne XLI, 792).

2 Introduction.

in the name of his colleagues:t “With reverential awe
does he (the naturalist) gaze at the microscopic speck
of nervous substance, which harbors the soul of the
ant with its industry, its instincts of architecture,
order, fidelity and courage.”

Surely, it was not without great reason that
scientific observers of recent times applied themselves
to the most careful and detailed examination of the
life of ants, especially since the publication of Pierre
Huber’s classical “Recherches sur les Moeurs des
Fourmis indigénes’ (1810). Very many interesting
facts of great value for psychological research have
thus been furnished. However, dabblers in popular
science, who viewed things from the standpoint of
“vulgar psychology,” as Wundt termed it, misinter-
preted these facts in a very unscientific manner; for
they tried to draw conclusions from them which led
to the humanization of animals, and denied the
existence of any essential difference between the
psychic faculties of man and brute. It is not so very
long since Ludwig Buechner endeavored to pro-
mote these ideas in his “Geistesleben der Tiere”
(Berlin, 1876). As is generally the case with such
shallow elaborations, Buechner has found not a few
imitators and plagiarists. Therefore, it may not be
out of place to examine these deductions from the
standpoint of critical psychology.

Sir John Lubbock, who, devoted himself to the
study of ant life with the accuracy of a professional
scientist, and who carefully refrained from the

1) “Ueber die Grenzen des Naturerkennens.” Lectures by E.
Dubois-Reymond, 1st issue (Leipzig, 1886), p. 127.

Introduction. 3

humanizing tendencies of modern times, states in the
introduction to his book, “Ants, Wasps and Bees,”
that ants rank next to man in the scale of intelligence,
and that in psychic faculties they approach nearer to
man than the Anthropoid apes even. George Romanes
in the sixth edition of his book, “Animal Intelligence”
(1895), devotes more than one hundred pages to ants,
and thus indicates the great importance he ascribes
to their psychic qualities.

Prior to Lubbock’s work on ants, another prom-
inent investigator of ant life, Dr. Augustus Forel, in
his “Fourmis de li Suisse” (1874) had expressed
the opinion that the principal factor in the psychic
activity of ants was not individual intelligence, but
social instincts (p. 444). Although he pretends to
find even among ants remarkable proofs of intellect,
he maintains that it cannot compare with the individual
intelligence of the higher Vertebrates (as apes, seals,
elephants, etc.). Most of my critics, likewise, espe-
cially Forel and Smalian, in discussing my book
“The Compound Nests and Mixed Colonies of Ants,”
conceded that ants were guided in their life and doings
almost exclusively by their social instincts. With the
higher Vertebrates, however, intelligence is said to
preponderate gradually over instinct. This is postu-
lated by Darwin’s theory of evolution, which other-
wise would be unable to explain the mental evolution
of man from the animal kingdom. The only possible
explanation according to this theory is to assume .
that, to the individual mammal-intelligence of the
hypothetical ancestors of man, there was added,
through the development of community life, a higher

4 “Introduction.

degree of perfection in their social instincts, and that
thereby the higher animal was gradually transformed
into man. ;

The tenability of this assumption will be discussed
in the following chapters. It is understood, that in our
comparative investigation, we shall be guided, not by
the postulates of evolutionist theories, but by the prin-
ciples of critical psychology, set forth at length in our
former essay, “Instinct and Intelligence in the Animal
Kingdom” (Herder, St. Louis, Mo., 1903).

Lately there has been invented a theory on the
psychic life of ants, which is dtametrically opposed
to the popular attempts at humanization. Alb. Bethe
has tried to set down ants and bees as mere “reflex
machines,” devoid even of the simplest sensitive per-
ception and cognition, whilst he considers the intelli-
gence of higher animals to be beyond all doubt. Thus
he hoped to succeed in destroying the parallelism
established by us between the psychic faculties of ants
and those of higher animals, from which we had
drawn the conclusion: we do not need ant intelligence,
therefore neither animal intelligence. Bethe’s work is
of undoubted value on account of its attack on the still
wide-spread popular views regarding ants as intelli-
gent, human beings in miniature.? His theory has

1) “Duerfen wir den Ameisen und Bienen psychische Qualitaeten
zuschreiben?” Bonn, 1898. (“Archiv fuer die gesamte Physiologie,”
LXX, 15-100.)

?) In this regard the “‘Betrachtungen ueber die staatlich lebenden
Immen,” published against Bethe’s essay by Charles Sajé in
“Prometheus” (10 Jahrg., 1899, Nr. 486 and 487), go far beyond what
is admissible. Similarly the essay by Kienitz-Gerloff, “Besitzen die
Ameisen Intelligenz?” in ‘Naturwissenschaftl. Wochenschrift” (XIV,
1899, n. 20 and 21),

Introduction. 5

already, on another occasion,’ been subjected by us to
a thorough discussion. Besides, in a longer essay,
especially intended for professional zoologists,? we have
since then perfected our former argument, proving
that ants are no more mere reflex machines than dogs
and apes are intelligent beings. The theoretical side of
Bethe’s psychological views was also noticed in the
second edition of our essay “Instinct and Intelligence
in the Animal Kingdom” (chapters 7 and 8).* In this
work, therefore, we shall return to them but occa-
sionally, to show the fatal results of attempting to
vindicate the intelligence of higher animals by com-
pletely denying psychic activities in ants.

1) “A new ‘reflex theory of ant life’? (Biolog. Centralbl., XVIII,
1898, n. 15, p. 577-588). a

2) “Die psychischen Faehigkeiten der Ameisen.’”’ Stuttgart, 1899.

(“Zoologica,” Heft 26) p. 134 and foll. with 3 plates.
3) p. 144 and following.

CHAPTER I.
COMMUNITY LIFE IN THE ANIMAL KINGDOM.

1. A General Survey of the Forms of Animal Com-
munities.

HE multiplicity of bodily shapes apparent in ani-

mals is not more remarkable than the variety
found in their biological relations. The majority of
animals, lower as well as higher, live singly, and only
temporarily join other individuals of the same ‘species
for breeding purposes; no lasting psychic tie unites
them with others of their species. Other animals live
in pairs during the breeding season, and remain united
until the young are old enough to shift for themselves ;
this is the case with most birds and mammals. If the
offspring continue to remain with their parents, fani-
ilies develop into herds, embracing the members of
different, allied families. Thus, v. g., wild oxen and
horses, the chamois, antelopes and many apes are
gregarious animals. While real gregariousness is
based on family ties in a wider sense and is mostly
permanent, other animals flock together to form troops
or hordes to undertake journeys in common, e. g., our
migratory birds in autumn, the Scandinavian lem-
mings, etc. Insects, too, gather into similar temporary
masses of individuals of the same or closely allied
species, assuming the form of migrating swarms.
Migrating locusts are known to everybody; but also

8 Chapter I.

butterflies, dragon-flies and other insects have been
observed to form similar swarms.

But few animal species are so perfect in their social
organization, that the members of the family construct
their habitations, rear their offspring and provide for
their food in common. These are what Aristotle calls
Zéa rodtixa, animals leading a well regulated social
life, comparable, in a way, to the social life of man.
These animals are chiefly the so-called state-forming
insects, the social wasps, bees, ants and termites. With
the two latter social life is carried to the highest degree
of perfection found in the whole animal kingdom.
True, also among birds, the social weavers (Ploceus)
construct habitations in common, inasmuch as they
build their nests close together, and beavers unite in
colonies to build their dams, when different pairs are
interested in raising the water level at the same spot.
But what is wanting in the associations of higher ani-
mals is co-operation, including some suitable division
of labor for the rearing and nourishing of their off-
spring. The combination of all these elements of
social life is found only among the social insects, and
in a prominent degree among ants.

Viewed from the standpoint of comparative psy-
chology, social is preferable by far to single life. In this
connection, of course, we mean a social life based on
social instincts, on the laws of sensitive cognition, and
not merely a union caused by the laws of vegetative
life, as is the case with certain animal conglomerates,
as sponges, corals, polyps and many species of Tuni-
cates. The bond, which unites the different individuals
of these species to a colony, is entirely material. They

Community Life in the Animal Kingdom. 9

live together from. immediate, vegetative necessity ;
for they literally grow as branches from a common
trunk. As it is an immediate vegetative necessity for
plants to bring forth twigs, leaves and blossoms, so
mere vegetative necessity forces a colony of Siphono-
phores to separate into different loosely connected
individuals, some serving the purpose of nutrition
(nutrient polyps), others of propagation (sexual
polyps), of perception (perception polyps), of loco-
motion (swimming polyps), and of protection (pro-
tective polyps). To apply to the members of such
colonies the term “persons” (eating persons, swim-
ming persons, etc.), as Haeckel and several other
zoologists have done, is evidently out of place, because
this term implies a psychic independence which these
animals do not possess. It would be more justifiable
to conceive the whole growth of .Siphonophores as
one individual of imperfect unity, consisting of various
members, which, on account of their different func-
tions can more fitly be termed “organs” than “persons.”

The similarity of social life in the colonies of
polyps and of ants is very slight and superficial. The
latter, in opposition to the former, consists of indi-
viduals organically separated and independent in their
psychic activities. The members of an ant colony are
complete individuals united to each other, not by the
laws of vegetative growth, but by tmstinctive sym-
pathy. This kind of co-habitation must indeed be
- regarded as a higher manifestation of psychic life
unknown among solitary animals.

It is true, with the state-forming insects also, the
instinctive association of the individuals of a colony

10 Chapter I.

is based on an organic, i. e., vegetative fact, namely
on the eeicd oe from one and the same ecb
called a “queen.”

Honey-bees have never more than one queen in the
hive, ants may have several of them. The instinctive
dependence of bees on their queen is not so great as
was formerly believed. Moreover, in the bee-hive the
queen has essentially no other function than that of
laying eggs; for the rest, her attitude towards the
social activities of the colony is entirely passive; even
when the bees are swarming the old “sovereign” is
generally hurried along by the crowd of her “faithful
subjects ;” she does not lead the expedition, neither
does she determine its direction.t However, a swarm
of bees deprived of their queen will disperse, because
they have no common center of attraction, no point
of crystallization, so to say, around which to form a
new colony. In bee-hives the instinctive bond uniting
queen and workers is closer than among ants, because
the odor emitted by the queen exercises a far more
powerful attraction? on the workers than in the case

1) Abbé J. J. Kieffer communicated the following observations:
“An old queen must often be actually forced out of the hive by the bees
already swarming; sometimes the bees are gone, the queen being left
behind in the hive. In other cases I observed that the old queen had
dropped to the grotind; in spite of this, the bees settled at quite a
different place on some tree, and suffered themselves to be put in a
new. hive which, however, they soon left again, because the queen was
missing.”

2) How powerful is this attraction, can be gathered from an obser-
vation made by Fr. Spillmann, S. J., in June, 1896. On catching a
cluster of swarming bees, a few hundred workers had remained in the
catching apparatus and could not find their way to the new hive. Led
by their sense of smell, however, they clustered around a queen that
had been lying dead on the ground for eight days, although it belonged
to a different hive,

Community Life in the Animal Kingdom. 11

of ants. Thus the queen of the bee-hive becomes in a
higher degree the principle of union for the workers
of her colony and for the regulated exercise of their
instincts. For this very reason but one full-grown
queen is tolerated in a bee-hive, while in ant nests
several may be found. What follows may ultimately
account for the fact.

The workers among ants live much longer than
among bees. According to my observations, our
Formica species, as a rule, attain an age of two, some-
times of three years, whilst the workers among bees
die after a few weeks or months. For this reason a
colony of ants can continue to exist for several years
without a queen, and even produce males through
parthenogenesis. This longer duration of life with
workers among ants may perhaps explain, why they
are less dependent on their queen than bees, and fairly
accounts for the fact, that, with ants, queens are
needed as the unitive principle of the colony in a far
inferior degree. Hence in a community of ants the
number of generating, impregnated females may be
almost unlimited. In a populous nest of the hill-ant
(Formica rufa) near Exaten (Holland), I once found
more than sixty full-grown queens. A similar num-
ber I met with in a nest of the small, red stinging
ant (Myrmica scabrinodis). In fact, by far the
greater part of European ant species have, as a rule,
several queens in every colony of long standing.
With foreign ants the case is pretty much the same.

A community of bees, therefore, having only one
queen, may aptly be compared with a monarchy. But
on account of the great number of oviparous mem-

12 Chapter I.

bers and the consequent greater independence of
instincts in the single worker, an ant colony bears the
stamp rather of democratic, republican, even socialistic
institutions. Viewed from the standpoint of compara-
tive psychology, the community life of ants is more
perfect than that of bees, on account of the greater
psychic independence of each individual. It is this
quality of individual independence that lends to ant-
states, among all associations of animals, the greatest
resemblance to the political societies of man based on
individual intelligence and free will. This resemblance
is of course never more than mere analogy; but it is the
highest degree of analogy known to exist between the
social institutions of man and of the brute. Nor is the
term “state” applicable to the social organizations of
ants or, in fact, to any animal community, in any other
than a metaphorical’ meaning; yet it applies more
perfectly to ant states than to any other family of
insects, and to insect states rather than to those of any
other animals.
_ Another important reason, why with ant colonies
the use of the term “state” is comparatively more
appropriate than with the social organizations of other
animals is, because colonies of ants are often not
merely “enlarged families,’ but contain also members
of entirely different species which are hospitably shel-
tered in the colony. Thus a simple ant colony comes
to be a compound animal society. The above-men-
tioned strangers are partly ants belonging to other
1) On this point vide A. Espinas, “Des sociétés animales” (2e éd.)
p. 372. Also Karl E. v. Baer (in Stoelzle, ““K. E. v. Baer und seine

Weltanschauung” [1896], p. 300); W. Wundt, “Vorlesungen ueher die
Menschen- und Tierseele,” 2d ed., p. 451.

Community Life in the Animal Kingdom. 13

species, living in the colony as “auxiliaries” or
“slaves”; partly they are members of altogether dif-
ferent orders of insects, especially of certain beetles,
as the genera Atemeles and Lomechusa, which are
accorded a friendly reception by the ants, are licked
and fed, their larvae being reared by the ants as if
they were the latters’ own.? This is a special form
of community life (symbiosis), found nowhere else
throughout the animal kingdom. Symbiosis is only
equal to real community life, when the members
engage in mutual psychic intercourse. Between a
hermit crab and a sea anemone that settles on the
former’s back, between a small fish (Trachichthys tuni-
catus) and a large sea nettle harboring it within the
circle of its tentacles,* there is a mutual relation (mutu-
alism) useful to both of them, without, however,
approaching any psychic intercourse, although the one
instinctively looks for the other. There is a similar
relation between ants and many of their tolerated
guests, whilst their relation to their slaves and to their
genuine guests attains a higher degree of psycho-
logical intercourse and becomes real community life.
Moreover, parasites, hostile intruders and indifferently

1) See Wasmann, “Die zusammengesetzten Nester und gemischten
Kolonien der Ameisen,” part II.

2) See the “Autobiography of a Lomechusa,’ in “Stimmen aus
Maria Laach,’”’ LII .(1897), 69, where the literature of the subject is
enumerated. The number of the regular nestmates of ants and
termites is rather considerable. Our ‘‘Kritisches Verzeichnis der
*myrmekophilen und termitophilen Arthropoden,” published in 1894,
already contains 1,246 ant guests and 109 termite guests, having the
most various biological relations to their hosts. Since then many new
species from all quarters of the world have been discovered and
described.

3) See “Zool. Anzeiger,”? Vol, XI (1888), n. 278, p. 240.

14 Chapter I.

tolerated cohabitants are found in the society of many
higher and lower animals. They are present like-
wise in the nests of social wasps, hornets and bumble-
bees; but genuine guests (Symphiles), which, in spite
of their morphological difference, are treated by their
hosts as enjoying equal rights, as members of the
family, are met with only among ants and termites.
That stray chamois or steinbocks should join a herd
of goats, is evidently something quite different from
the fact that ants keep aphides and scale-insects as
their milk cows, and tend even their eggs; or that
they feed from their own mouths certain species of
beetles, which on being licked afford the ants a special
pleasurable sensation, herein treating them the same
as they do their own comrades and larvae. The
mutual social relationship which is here seen to exist
between the animals of different species, and which
we term Symphily (ow-¢iia) is by far more per-
fect. Although, as we shall show later on, it is inti-
mately connected with the instinct of adoption which
occurs also among higher animals, the relation exist-
ing between ants on the one hand and their slaves
and genuine guests on the other, is nevertheless a
form of perfect Symbiosis unparalleled among the
Vertebrates.

2. The Social Basis of Ant States.

As was already indicated, the ultimate foundation
of ant states is organic. It is organic, not only be-
cause it is due to the descent from a common ovip-
arous female, but more especially because it is con-
ditioned, in its essential outlines, by polymorphism,

Community Life in the Animal Kingdom. 15

in other words, by bodily difference in the individuals
of a colony. Ant states are organically divided into
fixed groups of different “castes,” possessing different
corporal and psychic qualities. These castes take their
origin from the peculiar organic development of ants;
they depend on laws of vegative growth, not on the
intelligence and free will of individuals, as do the
classes of human society. By far the majority of
members of ant colonies consist, of course, of wing-
less neuters, which go by the name of “workers” or
simply “ants.” These workers are a secondary form
of the female, the ovaries being stunted, while brain
and instincts are all the more highly developed.t
With many ants, especially with the genera Pheidole,
Pheidologeton, Eciton, Colobopsis, etc., the workers
are again divided into two more or less strictly sep-
arated castes differing in bodily structure, namely:
workers proper and soldiers, the latter possessing a
comparatively huge head and formidable jaws. The
wingless workers and soldiers are entrusted with the
colony’s social welfare; it is their duty to build the
nest, to tend the young, to gather provisions and to
defend the community against hostile invaders, whilst
the winged males and females attend to the propaga-
tion of the species. After having been fertilized,
which is generally done in the air during their nuptial
flight, the females lose their wings and become
“queens,” either founding new colonies or being taken .
back by workers into their old nest for oviposition.
The basis, therefore, of the so-called political con-

1) Hence they cannot be simply called “stunted females,’ no more
than the workers among bees.

16 Chapter I.

stitution of ants’ is in fact organic; it consists in the
descent from one fertile female, and in the differen-
tiation of the descendants into castes differing in
bodily and psychic qualities, as a result of the very
same specific fertility. The social bond, however,
which unites the members of an ant colony and sep-
arates them from other colonies of the same species,
is psychic and instinctive. It is the feeling of fellow-
ship, the instinct of sociality, resulting from common
descent; it is, moreover, the instinct of imitation
which urges the workers of the same colony to act
in concert. This unity and co-operation is effected
‘by means of a certain sensile feeler language: by a
touch of their feelers thousands of members of a
colony immediately recognize one another as belong-
ing to the same community and effectually discover
the intruder; by taps of their antennae they exchange
their feelings and perceptions and thus draw the
attention of other workers of their colony to the same
work. The same feeler language is also the means
of communication of ants in mixed colonies with their
auxiliaries of other species, and of genuine ant guests
with their hosts.

This distinction between members of their own
colony and those of others is effected by very delicate
organs of smell? situated in the antennae. Members

1) We mean here in the first place the simple ant societies which
embrace no members of different species.

2) We have already proven in our work, “Die psychischen Faehig-
keiten der Ameisen” (“‘Zoologica,” 26th issue, p. 10-16), that there is
not merely question of a “chemical reflex” (as Bethe calls it), but of
a real sensitive perception. On the other hand, Lubbock’s experiments
(“On the senses, instincts and intelligence of animals’? [London, 1889],
p. 233 and foll.) have shown that an arbitrarily chosen sign or pass-

word is equally out of place, as is evident from the fact that an ant
which has lost its feelers is nevertheless recognized by her nest mates.

Community Life in the Animal Kingdom. 17

of the same colony have the same delicate “nest
odor,” and by licking strangers they are able to trans-
fer it to other insects. A beetle of the genus Atemeles
having been licked in a friendly manner by but one
ant of a Formica colony, will be acknowledged as a
friend by the other ants of the same colony, whilst
otherwise they would attack it The “nest odor” can
be communicated to members of other colonies not
only by licking but also by feeding. The smell of the
salivary gland secretions* thus seems to serve ants as
well as bees as a means of recognizing the “citizens
of the same state.”

It is, no doubt, downright nonsense for Buechner*
to put ant states on the same level with human
republics, much more so to consider them more perfect
than the latter. And when modern sociologists* try
to establish their reforms of human society on such
foundations, we are justified in styling their endeavors
utopian schemes. The promoters of such ideas for-

1) More on the significance of the salivary gland secretions as a
means of recognition among ants will be found in the essay mentioned
above “Die psych. Faehigkeiten der Ameisen,” p. 16 and 97 ff. On the
latter pages we have also shown that it is not merely the smell of the
salivary gland secretions adhering to a beetle, that induces the ants to
receive it after it has been licked by a single ant of that colony, but
that, besides, other psychic elements are in play and must be considered
in explaining the fact. See also “Instinct and Intelligence in the
Animal Kingdom,” p. 158.

2) See the interesting little essay by N. Ludwig, ‘‘Futtersaft oder
thierische Veranlagung als der Beherrscher und Ordner geheimniss-
voller Vorgaenge im Bienenvolke,” published by the “Leipziger Bienen-
_zeitung,”’ 1896. Likewise N. Ludwig, “Ueber Geruchempfindung und
Riechorgan der Honigbiene” (‘“‘Natur und Offenbarung,” 1899, 9th
issue, p. 554 ff.).

3) “Geistesleben der Tiere,” p. 52.

4) See e. g. Cognetto de Martiis, “Le forme primitive nella
Mate vs economica.” Torino, 1881.

18 Chapter I.

get, that with man class differences rest on far dii-
ferent bases than differences of castes among ants.
With man they are the outcome of changeable, out-
ward conditions of life, or perhaps the result of the
intelligent free choice of the individuals concerned;
with ants, however, they spring directly from the
hereditary organic laws of polymorphism. Besides,
those socialistic theorists forget that among ants there
exists perfect equality and fraternity between all the
members of a colony, for the very reason that these
animals are guided by their social instincts only, not
by independent reasoning, and that they therefore
are never liable, as men unhappily often are, egotisti-
cally to prefer their individual welfare to the common
weal. If those socialist enthusiasts could transform
men into ants, then they might be justified in pro-
posing ant republics as the ideal political condition.
H. E. Ziegler’ is right, therefore, in saying:
“With ants the social differentiation is conditioned
by organization and instincts, and is thus accurately
fixed and regulated, whilst with man the social differ-
entiation is due to education, exercise and custom;
only the foundation of man’s social life is determined
by certain social instincts, its further development,
however, is regulated by the intellect, by education
and custom . . . . To argue about man’s social
institutions from the relations existing among insects
would be committing a gross error, all the more so,
if one should consider the communistic insect ‘states’
1) “Die Naturwissenschaft und die socialdemokratische Theorie,”
p. 186. See also R. Leuckart, “Ueber den Polymorphismus der

Individuen oder die Erscheinungen der Arbeitsteilung in der Natur,”
Giessen, 1851.

Community Life in the Animal Kingdom. 19

as models of human communism.” Smalian' agrees
with Ziegler on this point, and I hardly believe that
any intelligent naturalist will dispute their position.

But now let us examine the other conclusions these
statements imply. The social life of ants in spite of
its differing essentially from the human state, is
nevertheless the highest degree of community life in
the whole animal kingdom; even the social relations
among the highest apes are far from reaching the
perfection of ant states. The foundation indeed of
social life and division of labor in ant states is organic,
and is to a certain degree predetermined by nature
with aprioristic necessity through bodily polymor-
phism. Nevertheless, also with them the actuation
of the social instincts is guided and determined in its
details by the sensile cognition and experience of the
individuals. Whoever falsely styled this individual
sensuous experience of higher animals, such as dogs,
apes, etc., intelligence,” should not be so inconsistent
as to deny to ants a high degree of the same “individ-
ual intelligence.’ Whoever without previous critical
analysis of his psychological notions maintains down-
right, that associations resulting from the sensile ex-
periences of the individual are intelligent, must credit
ants not only with the highest development of the
social instincts, but also with the highest development
of intélligence found in the animal kingdom. This
we wish to prove more in detail.

1) “Altes und Neues aus dem Leben der Ameisen,” in ‘“‘Zeit-
schrift fuer Naturwissenschaft,” LXVII (Halle, 1894), 39.

2) Which is done by Ziegler and nearly all modern zoologists, as
we have shown in “Instinct and Intelligence,” chapt. 2.

20 Chapter I.

In what does the pretended psychological superi-
ority of the associations of higher animals over ant
states consist? Let us try to clear up this question.

3. The Communities of the Higher Animals Com-
pared with those of Ants.

Both Ziegler and Darwin’ point to the fact, that
the higher mammals, especially apes, “sometimes form
societies for the purpose of receiving notice of danger,
for providing mutual protection and defense, for ob-
taining nourishment, sometimes even for united at-
tacks on their prey.’* Societies of ants have the
very same end in view. Although their main pur-
pose is to rear their young in common, yet those
other secondary purposes are not only not excluded,
but their pursuit and attainment by ants reach a de-
gree of perfection unequaled by the above mentioned
higher animals. However, neither Darwin, nor Es-
pinas, nor Ziegler, nor, in fact, any modern student
of animal psychology has ever succeeded in proving
that apes are conscious of their purpose, and therefore
act with intelligence, and that ants are without con-
sciousness of purpose, and therefore acting merely
from instinct.

Let us consider more closely the different points
of comparison. The higher animals living in hordes
aid their comrades by certain calls, giving warning
of danger. Some of them, e. g., the chamois, post
regular “sentinels” for this purpose. However, ants
do the same and in a manner much more indicative

1) “Descent of Man,” I. Chap. 4,
2) Ziegler 1. c., p. 189.

Community Life in the Animal Kingdom. 21

of intelligence. The whole difference lies in the fact
that instead of calls, the ants use another means of
sensile communication, namely, their feelers. If a
troop of “sanguine slavemakers,” as McCook calls
them, (Formica sanguinea), approaches a nest of the
negro ant (Formica fusca), then the first black ant
which has noticed the foe hurries back into the nest,
communicates her own fright to the other workers by
rapidly tapping them with her feelers and thus gives
a general alarm. The larvae and pupae are hurried
down from the higher parts of the nest into the deeper
galleries and chambers, and if the foe advances as far
as these apartments, the black ants run head over
heels through the secret openings at the opposite side,
and with their precious burden climb up stalks and
bushes to save it from the foe. Sometimes they re-
sort to this final means of escape at the first news
of danger and take to their heels before the van:
guard of the foe has reached the interior of the nest.
In a similar way, but adopting different tactics, the
yellow and the brownish-black meadow ants (Lasius
flavius and niger) struggle for safety, when their
nests are attacked by some Formica species. As soon
as the approach of the foe is discovered, the fact is
announced with lightning-like rapidity throughout
the colony by rapid strokes of the feelers. The
larvae and pupae, the winged males and the queens
are carried to the lowest recesses of the nest, and the
‘avenues to it are hastily blocked up with earth to
prevent the enemy’s advance. Whilst the small
Lasius is constantly closing up the approaches to the
interior of the nest with bits of earth, such of the foe

22 Chapter I.

as have ventured too far, are seized and killed by
_ crowds of the assailed.

If with higher animals it is a mark of intelligence
to “utilize the senses of all for the protection of the
commonwealth,” the same must be said of ants, and
in a more perfect degree. The posting of sentinels
for the protection of the community may be observed
with these social insects just as well, and even better
than with the social apes. In a nest of Formica san-
guinea comprising four species of slaves (or auxil-
iaries), namely, F. fusca, rufibarbis, rufa and pra-
tensis, which is under my observation for many years,
I can verify this fact every day. We _ subjoin a
diagram of this observation nest, as it will often be
referred to in the sequel.

The main nest and its annex are made of glass
plates in wooden frames. The space between the two
plates in each nest is partly filled with earth, their
vertical distance being from Io to 12 millimeters, so
that the ants have freedom of motion to perform their
work without being able to screen themselves from
observation. The upper glass plate is generally cov-
ered with a black cloth; for if light were permitted
to enter, the ants would coat the lower surface of the
glass with earth in order to darken the interior of
the nest. By means of glass tubes the main nest and
its annex are put in communication with each other
and with the other parts of the nest, which are like-
wise of glass. (See diagram.)

In the main nest, which corresponds to the interior
of an ordinary ant nest, the majority of the ants are
to be found with their queens, their larvae, pupae and

Community. Life in the Animal Kingdom.

feeding tube.

aunex.

—

main nest. /

Fig. 1.

24 Chapter I.

guests. In the front nest we see generally a number of
ants basking in the sun or engaging in different labors.
In the top nest a small number of sanguinea, rufa and
pratensis are usually found either on guard, or wait-
ing for the flies or other food which I occasionally
throw in. In the glass bulb of the feeding tube, even
if it happens to contain no sugar or honey, there are
always one or two ants, mostly fusca or rufi-
barbis, which have a special liking for this depart-
ment. Even on the dumping grounds, whither the
ants carry their dead, there are, as a rule, a few ants
to be found, remaining immovable and watching for
any suspicious circumstance in the nest or in its
vicinity. On March 26, 1896, from morning till eve-
ning one F. fusca and one F. pratensis were posted in
the refuse nest; on March 27, at 7 a. m. two F. pra-
tensis; at 10 a. m. two F. sanguinea took their place.
On March 28, in the forenoon, one sanguinea
was on guard, which having been taken out by
me and confined was soon after replaced by another
sanguinea for the rest of the day. On March 29 the
whole day one sanguinea; on March, 30 at 7:30a. m.,
two fusca; when at 8 a. m. I had taken out and con-
fined one of the two, I found that within the space of
half an hour another fusca had taken its place, where-
upon both remained there during the whole of the
forenoon, etc. Only during winter, after I had com-
pletely emptied the refuse nest and left it in the
same condition for a long time, no ants were seen
there for several weeks, because this part of their
nest was no longer $f any vital interesi to them, and
because the cool temperature kept them in the main

Community Life in the Animal Kingdom. 25

nest and in its immediate vicinity. It can hardly be
maintained that this posting of sentinels in the dif-
ferent parts of the nest was merely due to poly-
morphism; for the cognitive and appetitive powers
of the single individual ants of those five species in
various ways take a prominent part in it. As we
shall show hereafter, the same obtains in other forms
of the division of labor in ant states.

“Social animals perform many little services for
each other; horses nibble, and cows lick each other
wherever they feel an itching; monkeys hunt for
each other’s external parasites,” etc. Thus Ziegler
reproduces the statements of Ch. Darwin. But ants
of the same colony are quite as serviceable to each
other. Whoever has kept ants in suitably arranged
nests of observation, where they feel comfortable and
at home, can observe such “acts of charity” a hundred
times a day. Every time I gently lift the black cloth,
which protects the upper glass plate of the main nest
from the rays of the sun, I witness one or more of
these lovely scenes. Just now a worker of F. san-
guinea is lying immovable, stretched on her side, whilst
some of the companions are washing her; a sanguinea,
a fusca and a rufibarbis perform this work, and lick
her carefully, whilst she continues immovable; then
they turn her around and lick her just as carefully on
the other side. After half a minute the light which
floods the nest interrupts the performance, and they
flee to some darker spot, the patient soon following
their example. All the workers of each of the five
ant species living in my mixed colony without distinc-
tion render these services of cleanliness to one another.

26 Chapter I.

Sometimes one of the dominant, sometimes one of the
enslaved species is the recipient, no distinction being
made between masters and slaves in performing these
offices. Just as with cows in licking each other, so with
ants, the performance of this service generally causes
no less satisfaction to the active than to the passive
partner, and, when apes look for each other’s parasites,
we must, in order to arrive at a correct psychological
appreciation of such “kind offices,’ not overlook the
fact that apes devour with great relish the parasites
discovered in the fur of their comrade.

As regards these mutual cleaning services, ants
and the higher social animals are pretty much on a
par. The only difference is, that with ants they occur
much oftener than with the latter. In both they pro-
ceed, in the first place, from the desire for cleanliness,
which is no doubt of an instinctive nature.t In the
second place, they are due to the instinctive, mutual
attachment between the members of animal associa-
‘tions. The fact that ants clean a dust-covered com-
panion by carefully “brushing” her down with their
mandibles and licking her with their tongues, when
viewed from the point of comparative psychology,
finds its explanation in the same psychic motives as
when “apes, after having rushed through a thorny
brake, will examine each other’s fur and extract every
thorn or burr.” To lick off the dust is, by itself, not
more agreeable for ants; than it is for monkeys to
extract the thorns.

With ants the mutual attachment of nest mates

1) See Ballion, “De Vinstinct de la propreté chez les animaux,”
2d ed., Bazas, 1895,

Community Life in the Animal Kingdom. 27

goes so far as to make them carefully tend their
wounded and sick: companions, which is not the case
with gregarious mammals. The above-mentioned
observation nest of F. sanguinea, on which I bestowed
special care and attention, allowed me to observe
several times, so as to leave no doubt of the fact, that,
without distinction of masters or slaves, sick com-
panions, or such as were paralyzed by the formic acid
of hostile ants, were carefully nursed and licked for
whole days, until they recovered. Forel, too, considers
it a general rule, that ants nutse their sick or maimed
companions.! | |

Ants as “sick nurses” seemed so strange to me,
that I was unwilling to admit the fact, until I observed
it myself. The first time was on March 16, 1895. I
had replaced in the main part of the aforementioned
nest a sanguinea which had been paralyzed in one of
the narrow glass tubes by an ejection of formic acid,
and was scarcely able to move in spite of her con-
vulsive efforts. At first her companions, on approach-
ing, appeared to take no notice of her distress. Yet,
after a short time, they began to examine her with
their feelers, and then carried her to another part of
the nest where the greater number were assembled.
In this place the sick ant was lying for the whole day,
surrounded by a number of masters and slaves (fusca)
which, mostly in groups, busied themselves about her.
They licked her carefully, turned her over and licked
her again, examined her with their feelers and licked
her once more. This method of medical treatment was
attended with complete success. The patient had fully

1) See Lubbock, “Ants, Bees and Wasps,” Chap. V, p. 88 fi.

28 Chapter I.

recovered by the next day, whilst without nursing she
would probably have perished, as is generally the case
with ants paralyzed by poison.

If, therefore, on account of this “nursing,” Lub-
bock and Romanes ascribe to ants a certain degree of
“care and tenderness” lavished on their sick and
wounded companions, they are right in so far as those
actions are due to instinctive impulses, and not to the
conscious affections of rational beings. For comparing
the associations of ants with those of higher animals
it may, at any rate, be of particular interest to notice,
that such acts occur also among ants, notwithstanding
their highly choleric temperament.

“Yet, social animals also render more important
services to one another; thus wolves and some other
beasts of prey hunt in packs and aid one another in
attacking their victims. The Hamadryas baboons turn
over stones to find insects, etc., and when they come
to a large one, as many as can stand round, turn it
over together and share the booty. Social animals
mutually defend each other.” This quotation from
Darwin’s “Descent of Man” cannot justify Ziegler
any more than his former arguments in concluding,
that the community life of wild cattle, baboons and
other mammals is more closely related to the social
organisms of man, than that of ants. On the contrary,
the manifestations of social life recorded above occur
with ants even in far greater perfection.

Ants, too, hunt in company, especially the so-called
sanguine slavemakers (F. sanguinea and rubicunda),
the red Amazon ants (Polyergus rufescens, lucidus
and breviceps), and all the species belonging to the

Community Life in the Animal Kingdom. 29

Dorylide genera Eciton and Anomma. These are the
dreaded legionary ants and driver ants of tropical
America and Africa. The military expeditions of
F, sanguinea are generally undertaken in small divi-
sions of from twenty to fifty workers, with the purpose
not only of robbing the neuter pupae of the slave
species (Ff. fusca and rufibarbis), 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 san-
guinea 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 after-
noon of August 24, 1888, I witnessed such a typical
hunting expedition of several sanginea colonies near
Exaten (Holland), on the outskirts of a fir planta-
tion. 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.

The individual initiative of ants is manifested or
such occasions in the same degree as with the higher
mammals; whilst concerted action and _ suitable
co-operation reach even greater perfection than with
the latter. At any time a troop of our common red-
backed hill ants (F. rufa) may be seen on some forest
path, with combined forces dragging to their home a

‘30 Chapier I.

large, heavy dung-beetle (Geotrupes typhoeus) ; or a
number of them are hauling to the nest a large beam—
to our eyes it is but a broken twig—which is more
than fifty times the weight of any single ant! Some
pull in front, others push from behind, and even if the
latter for a few seconds pull in the wrong direction,
they soon notice it, and off it goes in the right direction
to the nest. On April 25, 1897, I observed in the
neighborhood of a pratensis nest near Exaten, two
- workers dragging together a beetle of the genus
Calathus towards their hill; they went at a double-
quick, without hindrance on either side, both ants run-
ning backwards with equal speed.
The mode of acting in concert is different with

different species of ants. Among our Formica species.

it reaches its highest degree of development with the
hill ants (F. rufa and pratensis), the initiative of the
single ants bearing rather a secondary part. The
sanguine slavemaker (F. sanguinea), however, which
is able to proceed unitis viribus wherever it seems suit-
able, combines with this power a remarkable degree
of individual initiative, similar to that noticed in dogs,
apes, and other higher animals.

It is of special interest to watch the co- -opetation

and division of labor of different species in mixed |

colonies of ants. In my above mentioned observation-
nest, which contains besides F. sanguinea four other
Formica species as auxiliaries, these five species have
divided the work necessary for the welfare of the
community, so as to give each species exactly the share
corresponding most to its instinctive preferences. This
division of labor, however, is neither mechanically

Community Life in the Animal Kingdom. 31

defined, nor confined within the cast-iron rules estab-
lished by the specific character of each ant, but the
workers of one species will at least to some degree
take part in the work of any other species. Thus e. g.
the rearing of the young in the main nest (see p. 23)
is chiefly attended to by the sanguineas themselves,
but all the four auxiliary species join in the same
work. In the glass bulb of the feeding tube containing
the sugar, the greater number of visitors consists. of
fusca or rufibarbis filling their crops by licking up
sugar or honey, with which they hurry to feed their
companions in the other parts of the nest. Sanguinea,
rufa and pratensis often prefer to carry the grains of
sugar “in the lump” from the feeding tubes to the top
nest, manifesting again various individual differences
in their mode of action, quite independent of their
specific character. Sometimes the lump of sugar is
carried only as far as y (see diagram on p. 23) to the
top of the tube, whence it is transported by other ants ;
generally, however, it is immediately brought (beyond
x) to the front nest, where it is carried to the bottom
in their mandibles, or else, but rather seldom, it is
simply dropped from a considerable height. This i
observed quite often with sanguinea, more rarely with
pratensis. When I introduce a large fly or some other
live victim into the front nest, it is mostly sanguinea
and rufibarbis that dart upon it furiously, whilst rufa
and pratensis manifest on such occasions remarkable
‘ skill and perseverance in holding down the struggling
and fluttering victim. The sanguinea with their
powerful mandibles attend chiefly to the dismembering
of their prey, whilst the conveyance of the larger pieces

32 Chapter I.

into the inner parts of the nest is generally done by
rufa or sanguinea.

It was of special interest to watch the behavior of
my ants, when a new individual of the beetle Lom-
echusa strumosa which lives with F. sanguinea as a
genuine guest, was introduced into the top nest. At
first, as long as there were no rufa and pratensis in the
nest as auxiliaries, he was, if not descending into the
main nest himself, taken at last by a sanguinea and
carried down, in spite of his obstinate, passive resist-
ance. Later on it was generally rufa and pratensis
that transported the guest who attracted their attention
in a remarkable degree. Once a rufa happened to be
alone in the top nest and for a long time was trying
in vain to get hold of one of the two Lomechusas
happening to be there, when all at once she ran down
to the front nest. Scarcely more than three seconds
had elapsed before she returned with four other rufas
which she had called to her assistance. Now the five
rufas immediately set to work with united efforts
to raise the Lomechusas, each of which was then car-
ried down to the main nest by one of the ants.*

If similar scenes had been witnessed in a society
formed of different species of higher animals, we could
not help admiring the harmonious co-operation and the
suitable, but by no means mechanical, division of labor.
However, it is not the higher animals, but ants that
act in this way, and in order to save the pretended
intelligence of the former, ants are classed as in-

1) A more accurate description of the last-mentioned observation
will be found in our essay, “Die psychischen Faehigkeiten der

Ameisen” (‘‘Zoologica,”’. 26th issue, Stuttgart, 1899), p. 63 ff. in the
chapter on the power of communication in ants,

Community Life in the Animal Kingdom. 33

stinctive automatons, or even as unperceptive “reflex
machines” !

Yet, neither in ants nor in any other animals, are
co-operation and division of labor such as to become
mutual, individual assistance, as is the case with man.
The same object attracts the attention of several indi-
viduals and leads them to busy themselves about it, |
each in its own way. Working in company is due
partly to the similarity of instinctive dispositions in the
single ants, partly to the instinct of imitation. H. v.
Ihering refers to this in the case of the Brazilian leaf-
cutting ants (Atta), and has pointed out the psycho-
logical importance of this difference as it exists
between societies of animals and man.*

Everybody knows that not only the higher mam-
mals but also the social insects unite in defending their
community and especially their young. To be thor-
oughly convinced of this fact you need but step on
a wasps’ nest or sit down on an ant hill. Indeed, the
perfect unity and heroic “self-sacrifice” which social
insects and in particular most ants display in defend-
ing their nests and their offspring are simply unsur-
passed by any other animal. This “unselfishness,”
this “spirit of sacrifice’ and “motherly love” in ani-
mals will be referred to in particular, when we come
to speak of the breeding and nursing instincts.

-Higher gregarious animals, e. g., bisons or baboons,
do not in defending the community against a common
‘foe defend also the individuals as such. When a
hunter lying in ambush has killed one of the herd, the

1) “Die Ameisen von Rio Grande do Sul,” in ‘Berliner Entomo-
logische Zeitschrift,” 1894, 3d issue, p. 346.
3

34 Chapter I.

other bisons generally take an inquisitive sniff at the
corpse, but they make no assault on the foe for the
sake of a wounded or dead companion. Wolves are
far more unceremonious. Instead of devising plans
for revenge, they devour their dead or wounded
“brother.” Ants, when engaged in common defense,
aim at defending the individual of the colony just as
little as do the higher animals. An assailed ant is
never defended by her companions for her own sake.
They rush upon the foe, only because they see in him
a common danger, and because their warlike spirit has
been aroused. This was noticed by Forel and Lub-
bock, and I can only confirm it. Therefore, neither
higher animals, nor ants, when either at work or in
battle, manifest anything like individual assistance in
the human sense of the word.*

“All animals living in a body, which defend them-
selves or attack their enemies in concert, must indeed
be in some degree faithful to one another ; and those
that follow a leader must be in some degree obedient.
When the baboons in Abyssinia plunder a garden, they
silently follow their leader ; and if an imprudent young
animal makes a noise, he receives a slap from the
others to teach him silence and obedience.”

Examples perfectly similar to the one just men-

1) The pretended instances of individual assistance in the legionary
ant Eciton hamatum recorded by Belt (The Naturalist in Nicaragua,
2d ed., 1888, p. 26), are easily explained by the fact that these migrat-
ing ants tried to take along their straggling companions, as is often
the case in migrations of European Formica species. Therefore, there
is no reason why we should credit the Ecitons with a higher “‘sympathy
for their companions” than other ants, as Romanes does = (‘Animal
Intelligence,” 6th ed., p. 48). This case is no proof of sympathy, but
merely a manifestation of the instinct of sociableness.

Community Life in the Animal Kingdom. 85

tioned, and which Ziegler has borrowed from Darwin’s
“Descent of Man,” may, when purged of arbitrary
anthropomorphic interpretations, be recorded also of
ants. He who takes the terms “fidelity” and “obedi-
ence” as they are applied to human beings, namely as
reasonable, voluntary subjection to the demands of
duty and authority, can ascribe “fidelity” and “obedi-
ence” to baboons as little as to ants. From the point
of view of critical psychology it is ridiculous to inter-
pret the “slap” given to the young baboon by its senior
to be, as among men, an admonition to fidelity and
obedience. The imprudent cry uttered by the young
baboon, if the story is to be credited at all, excited the
instinctive anger of the old apes as they were silently
advancing. The instinctive association of certain sen-
sile perceptions with certain sensile impulses affords
a much simpler and more natural explanation of this
fact. If, therefore, Darwin and Ziegler on this account
ascribe to baboons fidelity and obedience in the human
sense,’ they are but arbitrarily humanizing the brute,
as indeed Darwin has done time and again in the book
quoted above.

A slight analogy of what we call fidelity and obedi-
ence may indeed be observed in many animals, not
only in the higher species, but also in state forming
insects. Wherever a certain individual is the center
of operation for the instincts of the rest of the com-
munity, the latter will show it fidelity and obedience.
‘The swarming bees cluster around their queen
“faithful and obedient.” This allegiance is, of

1) See my former essay, “Instinct and Intelligence in the Animal
Kingdom.” (Herder, St. Louis, Mo.)

36 Chapter 1.

course, not so prominent with ants, whose queen
is much less of a center for the instinctive activi-
ties of the workers. With ants it is just the
workers that by their restless activity and the remark-
able display of individual initiative, are most power-
fully stimulating the instincts of their companions to
imitation, and thereby to actual co-operation in a given
work. The only difference between the baboons
described by Darwin and our ants is, that with the
former the instinctive communication between the
single individuals of a troop is effected mostly through
calls, with the latter, however, through taps of the
feelers. But both sometimes resort to more drastic
gestures to supplement their means of “communica-
tion.” If an excited F. sanguinea or fusca can not
‘succeed by taps of her feelers in inducing a companion
to join her work, she sometimes seizes her by the man-
dibles or by a leg and simply drags her to the object
which had first attracted her own attention. In the
same way an ant often protects her comrades from
a threatening danger first noticed by her. In my
observation nests I repeatedly noticed some F. san-
guinea or fusca, by taps of her feelefs or some other
more drastic measures warning their companions to_
be “on their guard.”” When, e. g., I took away the glass
tube connecting the feeding bulb with the top nest
(see diagram p. 23), and caught a few of the “senti-
nels” that instantly sallied forth from the opening of
the top nest ready to fight, I often remarked some ants,
that were posted near the opening of the top nest,
approaching the others, tapping them with their feelers
as a danger signal, and even getting hold of one, that

Community Life in the Animal Kingdom. 37

was about to run out, and pulling her back from the
dangerous spot. To interpret such psychic manifesta-
tions in higher animals as “intelligent actions” is evi-
dently inconsistent with denying to ants an equal or
even higher degree of “individual intelligence.” Criti-
cal psychology will regard such occurrences in ants
as well as in higher animals merely as associations of
sensile representations and impulses, which must be
classed as instinctive sensation, and not as intelligent
thought". The social instincts of animals, which in
their actual use are variously influenced and ruled by
individual sense experience, perfectly explain all the
appearances of “fidelity,” “obedience,” “caution,” etc.,
which occur with state-forming insects not in a lower,
but rather in a higher degree than with apes and other
mammals. To credit higher animals with quasi-human
intelligence is, therefore, to humanize animals in a
manner equally arbitrary and inconsistent.

To sum up the results of our comparative study on
the social life of ants and of higher animals. The
associations of apes and of higher Vertebrates are
based on social instincts, which lead them to co-operate
for mutual protection and defense, and partly, too,
for the procuring of food. This co-operation is more
or less powerfully influenced and varied in its manifes-
tations according to the sensile experiences and affec-
tions of different individuals. Exactly the same mode
of co-operation, but of a still more perfect, suitable,
‘and variable nature, we observe also in ant states.
With these animals, too, it is founded on _ social

99 66

1) See “Instinct and Intelligence in the Animal Kingdom’
(Herder, St. Louis, Mo.), especially Chap, III.

38 Chapter I.

instincts, which, corresponding to their organic poly-
morphism, are different in different classes (castes)
of the state. As regards the application of the sensile
experiences and affections of individuals, there exists
within the range of these classes a very great inde-
pendence and variableness of individual action, which
with several ant species, e. g., the sanguine slavemaker
(F. sanguinea), is scarcely inferior to that observed
amongst higher Vertebrates. Besides, the perfection
of social co-operation of higher mammals is far from
equaling that of ants; for with the latter it extends
not only to protection, defense and hunting, but also
to construction of their dwellings, to the rearing of the
young, and to the support of all the “members of the
state” by comparatively few individuals, going by turns
in quest of food and supplying the community with
provisions. Nothing of the kind is known of apes or
other higher animals. The providing of food_in par-
ticular varies greatly with the different kinds of ants:
it embraces “cattle herding” (the keeping of aphides),
hunting (robbing of insects, in particular, robbing of
the pupae of other ants), agriculture (grain gathering
ants), horticulture (ants raising fungi), etc. Nor are
the military expeditions of several ant species under-
taken merely from want of food, but also for the sake
of making slaves, the ravished pupae of workers of
other ant species being reared as members of their
own state. Through this suitable incorporation of
outsiders into their own colony the community life of
ants in the “mixed colonies” reaches a quasi-intelligent
universality, which is vainly sought for among higher
animals, The same universality is manifested also by

Community Life in the Animal Kingdom. 39

the fact, that many ant species treat like members of
their own family even different orders of insects,
namely the beetles of the genera Atemeles, Lomechusa,
Xenodusa, etc., which are known as “genuine ant
guests,” and that they even tend and rear their young
as if they were their own.

It must, therefore, be conceded that the community
life among ants is more developed and: more perfect
than that among apes and other higher animals ; hence,
from the point of view of comparative psychology,
the communities of ants represent the most perfect of
animal societies.

CHAPTER II.
WARS AND SLAVERY IN THE ANIMAL KINGDOM.
1. Wars Among Higher Animals.

ODERN evolutionists represent the social life
among higher Vertebrates with the aim of mak-
ing it the main support of the bridge spanning the
chasm between man and the brute. Thus, Ziegler
concludes his description by a psychological parallel,
in which, just as Darwin did, he tries to establish the
greatest possible similarity between the social life of
animals and of man. Let us examine this evolution-

istic attempt in the light of scientific psychology.
Says Ziegler : “There exists, therefore, among ani-
mals a social community life similar to what we meet
among the hordes and tribes of uncivilized nations.
Even wars, which have taken place among the hordes
and tribes of the human race since prehistoric times,
have their counterparts in the animal world, as is
proved by the following example recorded by Darwin.
“Brehm states on authority of the well-known
traveler Schimper, that in Abyssinia, when the baboons
belonging to one species descend in troops from the
mountains to plunder the fields, they sometimes
encounter troops of another species and then a fight
ensues ; the geladas roll down great stones, which the
hamadryas try to avoid and then both species, making

a great uproar, rush furiously against each other.’ ”
How far the “therefore,” which should connect

Wars and Slavery in the Animal Kingdom. . 41

the social life of animals and of man, is supported by
facts, has been shown in the preceding chapter. Even
in higher mammals individuals of social communities
co-operate merely as far as their social instincts guided
by individual sensile experience will allow. In man,
however, community life is due to social instincts
as to its foundation only, but in its perfect develop-
ment to the intelligent, free self-determination of indi-
viduals. Ziegler and Darwin are far from having
furnished the proof, that the latter element occurs also
in higher animals. Or do they perhaps think that
the wars which hordes of apes wage against each other
contain this proof? Let us see.

In the above description it is stated that the baboons
roll down stones-at their enemies intentionally, and
thus, as it were, use the stones as weapons, as e. g.,
in 1809 the Tyrolese occasionally did in their struggle
for liberty against the French and Bavarians. But
regarding apes, the statement is a myth. Pechuel-
Loesche corrected the passage in the third edition of
Brehm’s “Tierleben.” “We are told,” he writes, “that
apes defend themselves with broken branches, and it
is pretty generally assumed that they hurl down on
their opponents stones, fruits, pieces of wood and other
objects. This belief is probably due altogether to
inaccurate observation... Its originators and abettors
have perhaps seen only, what they from various

1) We sincerely regret that Mr. Pechuel-Loesche was not allowed
to subject the 3d ed. of Brehm’s ‘“‘Tierleben,’”? which he revised, to a
thorough psychological revision. Although several of the most offensive
passages were corrected or omitted, yet Brehm’s peculiar style has not
changed; he cannot possibly refrain from intentionally humanizing the

brute. See a criticism of this work in “Natur und: Offenbarung,”
XXXVII, 570.

42 ‘Chapter II.

accounts supposed to be the fact, not what took place
in reality. Apes living in trees, in wanton playfulness,
break off withered branches by jumping on them, by
snapping and shaking them ; but they do not throw
them at a person who stands below. Neither do they
throw fruits or other objects which they hold in their
hands; they rather drop them quite naturally on being
-frightened or put to flight. Moreover, baboons, among
which I was able to observe especially the tschakmas,
often watching hundreds of them very carefully, never
think of throwing down stones from their rocky eleva-
tions at their pursuers. It is true, from the place where
they happen to be, stones sometimes roll or fall down,
but merely by chance and also at times, when no enemy
is in sight. . . . Together with my wife, who
derived great pleasure from watching the behavior of
the baboons,—they were often the only living beings,
and very noisy at that, in the rocky deserts of South-
western Africa,—I have minutely studied their doings
precisely on this head to convince myself whether they
actually throw. They assuredly do not.”

What light is thrown by these critical observations
of Pechuel-Loesche on the “individual intelligence”
of apes so highly prized by modern evolution? Light
enough, indeed, but extremely compromising for that
theory. In spite of their highly developed brain, which
in anatomical structure bears the closest resemblance
to the human brain, apes are nevertheless unable to
draw even the simplest conclusions, which might lead
them to the use of branches and stones as weapons.
The spider weaving its ingenious web to ensnare its
prey, or casting out silky threads to entangle its vic-

Wars and Slavery in the Animal Kingdom. 43

tim; or the “ant-lion’” (Myrmeleon formicarius), an
insect belonging to the Neuroptera, the larva of which
from the center of its funnel-shaped sand-pit with its
long and flexible mandibles hurls particles of sand at
an ant, to bring her to the bottom of the pit ; or the
archer fish (To.rotes iaculator), directing jets of water
upon small insects resting on aquatic plants, thereby
bringing them down into the water as his prey : these
animals, so low in the zoological scale, are far nearer
to man, as regards the suitable use of weapons, than
the highest apes, although on evolutionistic principles
the latter only could possibly form the transition
between man and the brute. That apes through their
imitative instincts and by human training are able
to “learn” the use of a few simple tools, only proves
the intelligence of man and the power of sensitive per-
ception in apes. Had apes themselves but a trace of
intelligence, they would have invented long ago even
in their free state of nature the use of a few simple
means of defense, such as branches and stones. But
why did they not? The only possible, scientific answer
is : because they evidently have no intelligence. Not
the brain alone makes man an intelligent being, but
his spiritual soul, and this spiritual soul is wanting in
the highest apes as well as in insects. True, modern
evolution is fond of ignoring all facts, which will not
fit into its tissue of hypotheses. But we need not say
that such a proceeding is highly unscientific.
Darwin’s and Ziegler’s attempt at putting the wars
of apes on a level with those of uncivilized tribes, has

1) See “Instinct and Intelligence,” etc. (Herder, St. Louis, Mo.),
p. 60 and 160.

44 Chapter II.

proved unsuccessful. Even the most savage nations
employ tools and weapons of various kinds in order
to catch their prey or to wage war against their foes.
The parallel drawn by Darwin and Ziegler between
the wars of apes and of savages proves to an unpreju-
diced observer the very reverse of what Darwin and
Ziegler intended to prove : it proves the essential dif-
ference between the merely sensitive, psychic faculties
of the highest vertebrates, and the spiritual, mental
faculties of man.

2. The M ilitary Expeditions of the Amazon Ant and
of the Sanguine Slavemaker.

The wars of ants bear far greater resemblance to
human wars than those of the apes. Indeed, ants no
more than other animals use any other weapons than
those furnished by nature, namely their swordlike
mandibles, their poison stings and poison syringes,
but they use them in a manner which of all animal
combats most resembles human strategy. Whoever
watched a military expedition of the red Amazon ants
(Polyergus rufescens) or of the sanguine slavemakers'
will no longer entertain any doubts on the subject.
The Amazon ants, the European Polyergus rufescens
as well as the North American P. lucidus,? advance

1) Since the issue of the book, “Die zusammengesetzten Nester
und gemischten Kolonien der Ameisen” (1891), I have had occasion in
Lainz near Vienna to observe a number of other Polyergus expeditions,
and besides, several sanguinea expeditions near Vienna and in Lim-
burg (Holland), etc.

2) Called by McCook the “shining slavemaker,’”? whose habits he
observed near the Allegheny mountains. There are still three other
subspecies (races) of P. rufescens found in N. America, P. breviceps
Em., bicolor Wasm. and mexicanus For.

Wars and Slavery in the Animal Kingdom. 45

on the war-path in large serried columns, the sanguine
slavemakers, however, the European as well as the
North American,’ in smaller, less serried detachments ;
both, but especially the Amazons, try to storm the
hostile nest by a fierce attack, and to stun the numeri-
cally superior foe and to put him to flight by the
suddenness of the onslaught. Great success generally
attends these tactics. Forel, in his ‘““Fourmis de la
Suisse” (1874), p. 306, has several similar instances,
some of which we wish to bring to the notice of the
reader. When Forel brought a bag containing a
whole colony of meadow ants (F. pratensis), which
in size and strength surpass the Amazons, into the
neighborhood of an Amazon nest, several of the
Amazons at first dashed fiercely into the midst of their
numberless enemies; twenty of them were as a rule
sufficient to rout fifty times that number of pratensis.
Another time an army of Amazons just returning
from the pillage of a slave nest were depositing their
spoils of ant pupae in their nest; previous to setting
out on a new expedition, when Forel at a distance of
one meter from their nest and in the path of their
expedition emptied a large bag of F. pratensis. In
three minutes the whole army of the Amazons had
encircled the hostile camp appearing quite unex-
pectedly. They stormed it in an-instant, drove out
the pratensis and ransacked the nest for its cocoons.—I
would like to hear of apes ever displaying similar
‘military skill.

- It is characteristic of the military tactics of those

1) Formica rubicunda and integra Em, are the principal N. Ameri-
can races of the European Formica sanguinea.

46 Chapter II.

ant species which undertake slave hunting expeditions,
to kill the hostile ants only when resistance is offered.
Fleeing F. fusca or rufibarbis are pursued merely to
obtain the larvae and pupae which they are carrying
off; booty, not slaughter is the object of the victors.
If apes or other higher animals were to act similarly
in their wars, then our modern advocates of evolution
would not fail to make the following reflections:
“Here we find the first traces of genuine humanity,
which shrinks from unnecessary bloodshed; what
these animals consciously aim at is, not to fight, but
to gather the fruits of victory,” etc. In ants such
reflections are readily granted to be ridiculous human-
izations of the brute; but never would it be conceded
in the case of apes, not because the psychic manifesta-
tions are really different, but rather to safeguard the
evolutionistic theories.

The military skill of the Amazons (Polyergus) is
no doubt unexcelled amongst ants, but also amongst
other animals. It is even far superior to the military
tactics of the sanguine slavemaking ants, although the
latter manifest in their whole character a more perfect
development of what is called “individual intelligence,”
i. e., the suitable application of their sensitive experi-
ences. But the Amazons in private life are the dullest
and most awkward “instinct beings’ you can imagine.
Although they are able to take liquid food by licking
just as other ants, they have nevertheless almost totally
lost the instinct of feeding themselves, and would
starve, unless they be fed from the mouth of their
slaves. This fact makes it quite evident, that even in.
the grandest military exploits of the Amazons there

Wars and Slavery in the Animal Kingdom. 47

enters not the slightest trace of genuine intelligence,
but only instinctive sensitive faculties; for, an animal,
that even in a state of utmost destitution is unable to
combine his feeling of hunger with the perception of
nourishment and the impulse to eat, can surely not
be credited with even the lowest degree of delibera-
tion. “A being that is physically able to eat, but has
lost the habit of tt, is the greatest libel on animal intel-
ligence.’’*

Against this conclusion Dr. Smalian? has raised an
objection which we are now going to examine. He
believes our argumentation unsound; and asks “How |
does Wasmann know that the Polyergus are at all able
to feed? The basis of his argument is in concluding
from the nature of the eating organs the ability to
eat. And he states, that he has once seen Polyergus
taking food independently; however, the matter is
doubtful; for in the case of animals which otherwise
never feed themselves but are always fed by others,
it is impossible to know, whether the food they
touched was actually consumed.”

That Smalian should make such an objection may
be explained only by assuming that he does not know
the mode of life of Polyergus from actual observa-
tion; otherwise he would hardly have been led to attack
our argumentation. Besides, he has not reproduced
in full the proofs which he controverts. Indeed, it
was also from the anatomical structure of the mouth-
parts of this ant that we drew the conclusion, that no
organic impossibility prevented the independent feed-

1) “Die zusammengesetzten Nester und gemischten Kolonien der

Ameisen,”’ p. 204. :
2) “Altes und Neues aus dem Leben der Ameisen,” p. 42,

48 Chapter II.

ing of Polyergus. Our chief argument, however, was
the biological fact, that the Amazons do really some-
times lap up liquid food, if by chance it comes in con-
tact with the lower parts of their mouth. Dr. Smal-
ian has undervalued this fact. Not only once, but
repeatedly I saw and followed it up with a lens, how
some Amazon which had pierced an ant pupa with its
mandibles, licked up with her tongue the fluid flowing
from the wound, and sometimes spent a considerable
time in this occupation. Now, since the reception of
food in ants generally takes place by licking, it is hard
to understand, why it should be impossible to know
in this case, whether the food has “actually been con-
sumed.”

-Besides Dr. Smalian has failed to notice Adlerz’
observations mentioned in the very passage quoted by
him. Like myself, Adlerz has witnessed that the
Amazons frequently lick up the moisture condensed on
the glass walls of their artificial nests.. That Amazons
are able to feed independently is, therefore, an estab-
lished fact which can not be done away with. Why,
therefore, do they starve, when they are confined in
a test tube together with some honey or some appetiz-
ing ant pupae, but separated from their slaves by which
they are wont to be fed? The only possible and psy-
chologically correct answer is : because their hunger
does not compel them, like other animals, to seek for
food themselves, but only to beg food of other ants
by taps of their feelers. The sensitive perception of
the food placed immediately before them, in spite of
their feeling of hunger does no longer excite in them
the natural imffulse of tasting it. With these ants

Wars and Slavery in the Animal Kingdom. 49

the instinct of independent quest of food and of its
independent reception has perfectly degenerated. They
have become utterly dependent on their slaves. Once
more we ask Dr. Smalian and other friends of animal
intelligence: Is it possible that a being, which pos-
sesses but a trace of intelligence, should no longer be
able to combine the sensitive perception of its proper
food with the feeling of hunger? Therefore we still
maintain: A being that ts physically able to eat, but
has “unlearned” the habit of it, is the greatest libel
on animal intelligence.

The brilliant military talent of the Amazons is,
therefore, a merely instinctive power, which is assisted
by no individual intelligence. Just the most wonderful
manifestations of the psychic life of animals, such as
to a superficial observer exhibit the most striking
resemblance to intelligence, upon closer inspection turn
out to be evident proofs of the want of individual
intelligence in animals. The brighter the light, the
darker the shadows.

The sanguine slavemakers afford us far better
ground than the Amazons for assuming, that in their
military expeditions individual intelligence comes in
for a considerable share. Some scouts of F. sanguinea
happening upon a nest of some slave species return
with the news. As soon as the favorable moment
for an expedition has arrived, they go ahead showing
the way. Upon arriving at the hostile nest they gener-
‘ally do not rush blindly to the attack, but institute a
formal blockade; then, whilst one detachment impet-
uously forces its way to the interior, others keep a
sarcit watch on the outside and relieve the flying

50 Chapter II.

inhabitants of their larvae and pupae, which are the
only objects the robbers have in view. On the part
of the sanguineas this shows great cunning and looks
very much like intelligence. If a troop of. apes at
war with others were to surround the forest home of
their foe and if a select squadron of the assailants were
to penetrate into the woods, whilst the other part lying
in ambush tried to capture the fugitives, how our
modern evolutionists would be delighted with these
apes! Such an argument for animal intelligence they
would deem absolutely irrefutable, and they would
no doubt allow this to be an “intelligent stratagem.”
But sad to say, not apes but merely ants are skilled in
such stratagems; yes, ants whose brain “can by no
means compare with the brain of the higher animals!”
If the development of the brain is the real cause of
intelligence, then, of course, apes ought to be at least
as intelligent as ants, or rather far more intelligent.
In reality the reverse is the case, and thus, things
look rather queer for modern evolutionism.

Let us return to the military tactics of the sanguine
slavemaking ants. One characteristic feature, that of
reconnoitering the nest they wish to plunder, they have
in common with the Amazons. With these latter ants,
according to Forel’s observations, and my own, single
individuals are wont to set out to investigate the site
of a slave nest, and thus frequently enable the whole
army of Amazons to advance in serried columns over
a distance of thirty yards or more almost in a straight
line to the place they had marked out. This surprising
fact repeatedly observed by Forel and by myself can
not be explained in any other than the above-men-

Wars and Slavery in the Animal Kingdom. 51

tioned way. With the Amazons, however, this system
of reconnoitering cannot possibly have anything to do
with individual intelligence, but only with instinctive
sense faculties. This we have sufficiently proved -
before. Hence, the necessary conclusion is, that also
in the case of the sanguine slavemakers, instinct aided
by sensitive experience will suffice to explain the same
fact. Therefore, to postulate intelligence proper for
such an explanation would be arbitrary humaniza-
tion of the brute.

The other seemingly very intelligent feature in the
military tactics of F. sanguinea is their habit of setting
out in smaller detachments more or less independent,
and uniting only when one of the bands has some-
where met with stronger resistance. Since most
colonies of the negro ant (F. fusca), which is the
ordinary object of their slave hunting expeditions, are
not very populous, and as the inmates generally take
to their heels at the first attack of the sanguineas, the
latter’s tactics of dividing their forces is evidently
appropriate to the usual conditions. Yet, if the assault
is directed against an unusually populous and well
defended nest of F. fusca, or against a large nest of
the far more warlike F. rufibarbis, the same tactics
frequently prove very disastrous to a considerable part
of the assailing sanguineas. The first troop of the
marauders venturing too close to the hostile nest is
attacked and overpowered by the defenders, and sus-
tains great losses, before any of the robbers are able
to hurry back to call for assistance. If in the military
tactics of F. sanguinea there were any question of intel-
ligence or rational deliberation on the part of the single

52 Chapter II.

individuals, they would surely show the prudence and
precaution of previously exploring more accurately
the forces of the foe they want to attack. Thus, they
would not dare an assault upon stronger slave nests,
until a greater number of forces were collected; then
they would, like the Amazons, fall upon the hostile
nest in compact masses of many hundreds or thousands
at a time, and would take the hostile position by storm
without any considerable loss. Why does such a
change never occur in the tactics of the sanguine slave-
makers? A colony of these robbers, which for many
successive years has pillaged the slave nests of the
neighborhood and has experienced the different resist-
ance offered by different hostile colonies, could easily
remember their respective strength and could regulate
the manner of future attacks according to this knowl-
edge. It would be all the easier for them to make an
intelligent use of their former achievements and
reverses, because the worker ants generally live for
the space of at least two or even three years. And yet
not a trace of all this can actually be found. F.
sanguinea will forever cling to her wonted tactics of
setting out in small, scattered bands, even if bloody
failure should ever so often be the result. To an
unprejudiced psychologist such facts bear sufficient
evidence of the fact that the warfare of F. sanguinea
as well as of Polyergus is guided merely by hereditary
instincts, not by individual intelligence. Those tactics
were not invented by the intelligence of the ants;
otherwise the same intelligence of the ants would be
able to perfect and to develop them. Yea more: the
assumption of ant intelligence is contradictory to the

Wars and Slavery in the Animal Kingdom. 53

fact that those tactics are specifically constant, and are
specifically the same throughout the entire territory
inhabited by F. sanguinea.

Dr. Smalian has tried to invalidate this conclusion
also. Here is his objection.’ “It was totally wrong of
Wasmann to demand,? that the sanguineas should
change their tactics, instead of continually attacking
in small troops and thus being easily overpowered by
large troops of fusca or rufibarbis. This mode of war-
fare is inborn, and therefore instinctive, no less than
the pillaging habit itself.”

Dr. Smalian is wrong in ‘believing that we had
in reality demanded of F. sanguinea to change her
hereditary, instinctive stratagems. Our demand was
merely the well known method of argumentation ex
absurdo, which the critic seems to have misunderstood.
In the supposition assumed by Smalian, but rejected
by us, that ants besides their instinct possess also a
certain degree of genuine intelligence, it is perfectly
justifiable to demand that this intelligence should also
be manifested and displayed. If their tactics are inborn
only as to their outlines, this manifestation ought
necessarily to consist in changing them intelligently
according to circumstances, and consequently in their
gradual perfection. But there is no trace of any such
advancement towards perfection, and therefore we are
right in concluding: These red marauding ants have
only instinct, not intelligence. This mode of argu-
‘mentation cannot seriously be styled “totally wrong.”

Wherever the sanguine slavemakers live, they will

1)°L.¢.,. p. 41.
2) “Die zusammengesetzten Nester,” etc., p. 203.

54 Chapter Il.

follow the habit of invading nests of certain smaller
species of Formica, and of rearing the robbed worker
pupae, partly at least, as auxiliary ants for their own
colony. It is, moreover, a constant characteristic of
F, sanguinea to have rather a small number of slaves,
if compared with those of the Amazons. . With these
latter the slaves are far more numerous than the
masters, with the former it is the reverse. Likewise,
the specific military tactics are everywhere equally
constant with both ant species. From the Alps to
England and Scandinavia, from Holland to the
Caucasus, F. sanguinea nowhere changes her habits
and customs. Even her North American sub-species
(rubicunda Em.) shows the same instinct of slave-
making, and this in the same specific form. The only
difference is, that one of the two European slave
species, F. fusca, is represented in the North American
rubicunda colonies by a closely allied variety, namely
by F. subsericea.*_ Since the separation of North
America from Europe was completed in the Tertiary
age, the enslaving habits of the sanguineas and their
military tactics must have been essentially the same in
the Tertiary as they are today. This is the most natural
explanation for the specific uniformity of that instinct
in the different parts of the globe. One thing, how-
ever, is certain: if the impulse of slavemaking and the
specific military tactics of F. sanguinea were due to
the intelligence of the ants, or if they were even in the
slightest degree dependent on it, such a specific uni-
formity existing for thousands of years would be
utterly inconceivable.

1) See Wasmann, ‘“‘Kritisches Verzeichniss der myrmekophilen und
‘termitophilen Arthropoden” (1894), p. 163 ff.

Wars and Slavery in the Animal Kingdom's 55

3. The Pretended “Automatism” in the Psychic
Life of Ants.

Animal intelligence, therefore, has no part either
in the slavemaking expeditions of ants, or in their
military tactics. Yet, the application of these instincts
is not mathematically uniform, They are influenced
and governed by the changeable sensitive perceptions
and individual conditions of the single ants, and thus
great variability exists within specified limits. Those
animal psychologists who, in contradistinction to the
higher animals, call ants mere “instinct automatons,”
or even mere “reflex machines,” are asked to consider
that the instincts of ants are neither more nor less
“automatic” than those of dogs, apes and other verte-
brates. Instances of ‘intelligence in the true sense of
the term can be discovered with the latter as little as,
and even much less than, with ants. Various differ-
ences, however, of individual character, and of. indi-
vidual action, determined by different sense perceptions
and sense experiences, occur with ants as well as with
the higher mammals.

On turning over the stone or the piece of sod
covering a middlesized nest of F. sanguinea, and thus
suddenly exposing the interior to the light, we perceive
all the inhabitants in tumultuous excitement. Part of
the ants furiously biting and ejecting poison attack
the invader; others take care of their imperilled off-
‘spring and in haste carry down the eggs, larvae and
pupae to the lower chambers of the nest; other indi-
viduals of the same colony seem destitute of the
chivalrous spirit of their race for the defense of

56 Chapter II.

country and escape under sheltering grass tufts or
clods of earth; sometimes, even, in the midst of her
fighting, rescuing or fleeing comrades, a sanguinea
presses herself to the soil motionless and, though
mostly for a short time, has recourse to the instinctive
trick of “feigning death ;” in opposition to these, other
sanguineas, finally, seem to be seized by a strange
mixture of courage and fear, by a sort of impotent
rage: not venturing to attack the real foe, they vent
their spite against other objects; with sprawling feet
they crawl along the ground, and with their heads bent
down they furiously bite the sand or stalks of heather,
attacking everything, in fact, but the finger of the
great human monster that robs their nest of Lome-
chusast and other favorite guests. Such scenes as the
one just described I have observed hundreds of times,
and am so accustomed to them, that I find them quite
natural; nevertheless, they are of the utmost import-
ance for comparing the psychic faculties of ants and
those of the higher animals. Packs of wolves or hordes
of apes on similar occasions could display no greater
variability of individual character and of individual
action, than such a colony of sanguineas. Yet, ants,
we are told, are “instinct automatons,’ and apes or
wolves are not!

1) In the colonies of the North American subspecies of sanguinea,
‘F. rubicunda, the European Lomechusa strumosa is represented by an
allied species, Xenodusa cava. Rev. H. Muckermann, S. J., of Prairie
du Chien (Wisconsin) has recently found also the curious pseudogyne
ant form, which is due to the education of the larvae of Lomechusini
by the ants, in the colonies of F. rubicunda. See Wasmann, ‘Neue
Bestaetigungen der Lomechusa-Pseudogynentheorie” (Verhandl. der
Beutsch. Zool. Gesellsch. 1902, p. 98-108 and PI. II.). We shall give
the figure of Xenodusa later on, opposite p. 181.

Wars and Slavery in the Animal Kingdom. 57

A dog biting the stone thrown at him, in his blind
rage acts just as ‘‘automatically” as a sanguinea which
vents her fury on the edges of a glass tube, so that you
can hear the grating noise made by her jaws. And if
certain individuals of an ant colony acquire through
their sense-experience special dispositsons and char-
acteristics, which distinguish them from other individ-
uals of the same colony, then they act “automatically”
as little as dogs or apes, or other higher mammals do.
Some remarkable instances of this may find a place
here.

In the observation nest of F. sanguinea described
on page 23, some beetles called Dinarda dentata, which
I introduced, had at first been received as usual without
difficulty as indifferently tolerated guests, and had
even propagated in the nest. But several times I put in
a little larger Dinarda species (D. Maerkelii), whose
usual host is F. rufa, and when finally some small
sanguineas and their slaves had succeeded in seizing
and killing this beetle, which, as a rule, is unassailable
owing to its wedge-shaped body offering scarcely any
point of attack,t then a number of ants of this colony
gradually took a liking to catching Dinardas, which
liking proved disastrous also to the smaller Dinarda
dentata. Not all the individuals of the different ant
species of that colony have acquired this strange
passion. Among twelve workers of F. sanguinea
which I put from this observation nest into a smaller
_ experimenting nest together with seven Dinarda

1) See Wasmann, “Dinarda-Arten oder -Rassen,” in Wien. Entom.
Ztg., 1896, 4th and 5th issue, and ‘‘Die Myrmekophilen und Termi-

tophilen,” p. 435 (Extr. du Compte rendu du troisiéme Congrés intern,
de Zool. Leyden, 1896).

58 Chapter II,

Maerkelii, there was but one Dinarda hunter. Whilst
the rest remained perfectly passive towards the
Dinarda, this one ant immediately began an active
hunt. Had I not soon removed her from the small
nest, she would probably have aroused in her com-
panions the inStinct of imitation for a similar persecu-
tion, a fact which I have often observed. But by
remoying this passionate hunter, I preserved friendly
relations between the other ants in the same experi-
menting nest (11 sanguineas, 2 rufibarbis, 2 fusca)
and the Dinarda Maerkelu. In the greater observation
nest, from which I had taken these individuals, the
Dinarda hunt, which had begun with the killing of
Dinarda Maerkelu in March 1896, continued against
D. dentata until November of the same year, when the
ants gradually returned, but only for a short time, to
their former toleration of these guests. The resuming
of experiments in the following spring resulted finally
in the complete extermination of all the Dinardas in
that observation nest. During the following six years,
I never succeeded in securing the existence of even a
single D. dentata in that nest, although in nature this
beetle is indifferently tolerated in all sanguinea nests!
The psychological importance of these phenomena has
been pointed out already in our discussion on the
different forms of learning in the animal kingdom.
Another strange fact quite irreconcilable with the
“blind automatism” of instinct is the behavior of ants
regarding the number of Dinardas and other beetles,

1) “Instinct and Intelligence,” etc. (Herder, St. Louis, Mo., 1903),
p. 157. Also “Die psychischen Faehigkeiten der Ameisen” (Stuttgart,
1899), pp. 84, 88, 93,

Wars and Slavery in the Animal Kingdom. 59

which I introduced into that observation nest during
my experiments on the international relations’ of ant
guests. If previously no Dinarda had been present for
some time, one or two of D. dentata were often quietly
received and tolerated for weeks. But as soon as I
would add some more beetles of this species, the perse-
cution began, at first against the newcomers, and
finally, the passion of hunting having been aroused,
also against the old ones, until all were seized and
devoured. Thus the ants seem to have been led to this
persecution not so much by the perception of one beetle
by itself, but rather by that of their increasing number.
It was this that aroused their hostility.

The sanguineas of the same observation nest once
manifested unmistakably that they desired only a
limited number of their genuine guest Lomechusa
strumosa, otherwise so dear to them. In September,
1898, I had captured in a sanguinea colony? 116 beetles
of that species and placed 30 of them in my observa-
tion nest. After a few weeks, however, 19 of them
had been driven out of the main nest and were con-
fined in the empty annex, where they all perished,
being isolated from their nurses, whilst the remaining
11 were for the whole winter carefully tended by their
hosts and very often fed and licked by them. The
ants, therefore, had not changed their behavior toward

1) “International relations’ I term the relation of ant guests to
different colonies and different species of ants, which are hostile to

. @ach other.

2) No. 191 of my statistical map of the sanguinea colonies near
Exaten. In treating about the education of the Lomechusa larvae by
F. sanguinea, I shall communicate more detailed observations regarding
this colony (in the last chapter, number 3, “Adoption Instincts in the
Animal Kingdom’’).

60 Chapter II.

the Lomechusas as such, but merely toward their
excessive number, which became disagreeable to them.
Perhaps they were unable to feed so many beetles and
had therefore expelled half of them. At any rate,
such phenomena prove to a certainty that ants are
not to be regarded as mere “instinct automatons” or
“reflex machines.” We must ascribe to them sensile
mental faculties, which by way of different percep-
tions and representations cause great variability in the
display of their instinctive impulses. But, beyond this,
nothing is required to explain satisfactorily the psychic
life of the vertebrates. Hence, there is no need of
“animal intelligence,” neither in the case of ants nor
in that of the higher animals.

A beautiful instance of how sensile experiences of
ants lead them to acquire certain individual peculiar-
ities of character, I witnessed in the case of a F. rufi-
barbis of the same mixed colony. She was a worker,
easily distinguishable from the others by her small size.
She used to visit regularly the glass bulb of the feed-
ing tube (see diagram, p. 23), where she would lick
the honey or sugar in order to supply the other ants
in the main nest from the sweet juice stored up in
her crop. Although F. rufibarbis belongs to a very
irritable and pugnacious species, yet this ant had
gradually become so tame that she would allow her-
self to be fed from my hand. As soon as I removed
the cork of the glass bulb, she would come out and
look for food on the outside. I would then present
to her a needle dipped in honey. At first she darted
back, but after a few seconds of hesitation she would
approach, examine the needle with her feelers and

Wars and Slavery in the Animal Kingdom. 61

lick off the honey. Later on I placed the honey on my
finger. The ant had already become so tame that
she was not in the least disturbed by the odor of my
finger, whereas others would have been provoked to
a fight or would have been greatly alarmed. She
would quietly lick off the honey and then, without
resisting or trying to flee, allow herself to be seized
with a pincette by one of her legs and placed back in
her nest. This goes to prove that ants also are tama-
ble in spite of their excitable nature. The tamable-
ness of ants, like that of higher animals, is due to their
possessing the powers of sensitive perception and imag-
ination, upon which the intelligence of man acts to
accomplish his purpose.

Against this parallelism an objection was raised
by Mr. Bethe.t To tame an ant, he says, takes
weeks and months; but a dog may be tamed in a few
days; therefore the above mentioned fact presents no
proof of the existence of psychic faculties in ants!
Whether Mr. Bethe will succeed each time in taming
a vicious dog within a few days, is rather doubtful.
Nor is it at all true, that it takes several weeks or
months to tame an ant, e. g., a F. fusca or rufibarbis,
which are especially suitable for such experiments.
It is but required to mark a certain individual which
comes regularly to the feeding tube. If you are very
careful not to frighten the animal, it is possible to
train it in a few days, in the manner described above.
' But if you wish to reckon the time needed for taming
an ant by beginning with the day on which she was

1) “Duerfen wir den Ameisen und Bienen psychische Qualitaeten
zuschreiben?”? (Bonn, 1898), p. 23.

62 Chapter II.

first deprived of her freedom and placed in the arti-
ficial nest, then the same method of calculation must
be applied in the case of the dog. Mr. Bethe should
not, therefore, take for his experiments a domesticated
dog, but he would have to operate upon a newly-
captured animal of the wild dog species. Then let
us see, which would take longer, to tame a wild dog
or to tame a wild ant!

There are several other interesting analogies
between the psychic life of ants and that of dogs.
A small dog, as long as he is in the company of his
master or of some stronger comrade, will not be afraid
to meet a rival, whom otherwise he would try to avoid.
The same is the case with the small black negro ants
(F. fusca) when they are in company with sanguineas.
In their own colonies they are generally cowards.
As soon as their nest is disturbed, they flee and try to
hide their young, but when they are slaves in colonies
of F. sanguinea, they are the bravest defenders of the
mixed colony, as I have often experienced to my cost.
Just as in the mixed colony of F. sanguinea the
instinctive courage of F. fusca, which is otherwise so
cowardly, is to be explained psychologically from their
perception of the great number of valiant companions
and their consequent sense of solidarity, without sup-
posing any reasonable deliberation on their part, so
also are the different degrees of courage found in
different colonies of the sanguine slavemakers to be
accounted for. If a numerous population inhabits a
rotten fir stump, on the surface of which we find some
of the ants running about, a gentle kick will at once
call forth a whole army ready for the fray. In a

Wars and Slavery in the Animal Kingdom. 63

moment the whole surface of the stump is covered
with thousands of ants furiously hurrying to and fro.
But, if the colony is weak, the same kick, which at
other times calls forth an army, will have the con-
trary effect. The ants which just before were running
about the surface, disappear through the entrances of
the nest as if by magic, and deathlike quiet succeeds,
If in ants this appropriate estimation of the strength
of their own colony is characterized as instinctive,—
and this is, no doubt, the only correct expression,—
then, similar occurrences among higher animals should
also be credited to instinct and not to intelligence.’
Yet, the courage of individual ants in a colony is
dependent not only on the perception of the great
number and courage of their comrades. In populous
colonies also of warlike species such as F. sanguinea,
there will always appear considerable differences in
individual courage, as we have shown above. Some-
times even single, isolated individuals make head
against a numerous foe. Such an example of “hero-
ism” scarcely ever equaled by dogs, lions, and tigers,
was once witnessed by Rothney? in Bengal. A mid-
dle-sized worker of a large black ant species (Cam-

1) Forel (Un apergu de Psychologie comparée [1896], p. 25) men-
tions an instance, where a very strong colony of Camponotus ligni-
perdus, when fighting with F. pratensis, showed a more warlike spirit
-than is the custom with the ordinary colonies of that species. We fully
agree with Forel if he infers therefrom the existence of “plasticity”
in the psychic faculties in ants. Our own observations mentioned
‘above prove the very same. Yet, in considering this plasticity of the
sensitive powers of cognition and appetite to be essentially identical
with human intelligence, he i$ entirely wrong, as we have shown in a
former publication, ‘Instinct and Intelligence in the Animal Kingdom.”

2) “Notes on Indian Ants,” p. 349 (Transact. Entom. Soc. Lon-
don, 1889).

64 Chapter IT,

ponotus compressus) attacked quite alone a whole
colony of small red ants (Solenopsis geminata).
Without moving from her place she remained before
the entrance of the Solenopsis nest from 4:30 p. m.
until night, seized the ants with her jaws as they came
out, and bit them in two. At last she was overpowered
by superior numbers, and after having killed 150 or
200 of her foes she paid for her temerity with her
life. We leave it to the modern worshippers of animal
intelligence to raise a monument to the memory of
this insect Leonidas.

We need not, however, go as far as Bengal to find
‘such examples of “heroism” of single ants. There
are plenty of them in the heaths of Germany and
Holland, and also in North America. During hot
weather strolling workers of sanguinea will frequently
enter into a fierce battle with colonies of Lasius niger
or Tetramorium caespitum living in the neighborhood,
till at length, when too many of their opponents have
clung to their legs, they fall victims of their own fool-
hardiness. A scene, more harmless indeed, yet psy-
chologically not less remarkable, I observed near
Exaten on the afternoon of August 15, 1894. A large
worker of F. sanguinea amused herself for a quarter
of an hour by blockading all alone, a colony of the
small, red stinging ants (Myrmica scabrinodis). She
lurked about the entrance, seized by the neck one red
ant after another, as they came out, carried them quickly
to a distance of several inches, and dropped them in
order to be back again at once at the entrance to seize
the next customer. The Myrmicas scarcely attempted .
any resistance, although several dozens of them were

Wars and Slavery in the Animal Kingdom. 65

_in and around the entrance. Only one or two tried to
get hold of the intruder by one of her legs, but with-
out success. Of course, their horny (chitine) armor
protected them sufficiently against the jaws of the
sanguinea; all the more, as the latter did not allow
herself time to pay special attention to her single foes.
It was exceedingly amusing to see the indefatigable
zeal and hurry of the large ant getting hold of the
small ants one after another and carrying them away,
after which they slowly crawled home again. It is
hard indeed to say what induced the sanguinea to
engage in this odd kind of skirmish. Perhaps it was
mere love of fighting. That she released the single
Myrmicas so soon, might be explained by her fear
of the sting, with which these ants are provided. Yet,
it is more probable, that the marauder took a fancy
to the entrance of the Myrmica nest, and on this
account tried to expropriate the inhabitants. Of
course, her labor was no more successful than that of
Sisyphus, because she did not carry the ants further
than a few inches from the nest, and, besides, the
number of ants coming out of the nest had no end;
but this did not seem to affect her in the least.

It would be ridiculous, arbitrarily to humanize
such instances and to suppose all possible kinds of
“intelligent purposes” on the part of the ant, as is
customary with popular psychology. Yet, on the other
hand, it cannot be denied, that “mechanical autom-
atism” of instinct will never explain them. The only
satisfactory solution psychology can give, is to
ascribe to ants sensitive powers of cognition and appe-

tite, which, under the influence of exterior sense per-
5

66 Chapter II.

ceptions and individual dispositions, are the principle,
from which these various, spontaneous activities result,
with no difference as to whether the actions are per-
formed by ants, or by dogs and apes.

The phenomena described above may be classed
among the “sports” or “games” of animals, as Groos?
terms them. The facts recorded deserve these names
perhaps just as well as the sports and games of the
higher animals; only, it is generally much more diffi-
cult to ascertain the nature of given facts in the case
of ants. Among the heaps of ants that gather on the
surface of the ant-hills of F. rufa and pratensis, as also
in my artificial nest containing sanguineas when
exposed in spring to the warm rays of the sun, I have
repeatedly observed instances of harmless wrestling,
beginning with and accompanied by lively and playful
movements of the feelers. This behavior of the ants
seems to be due to a resuscitation of their powers and
also to an excess of muscular energy after the winter’s
rest. ;

Forel (Fourmis de la Suisse p. 367) has made
similar observations with F. pratensis, and Huber? with
F, rufa and pratensis. I cannot consider these games

1) “Die Spiele der Tiere” (1896), pp. 125 and 135. By the way,
Groos here and elsewhere was too confident in trusting the authority
of Buechner, who has not unfrequently misrepresented Huber’s and
Forel’s observations to suit his own purposes of humanizing the brute.
Forel, in the Etudes myrmécologiques, has expressly protested against
Buechner’s misrepresentations of his observations. The book of Groos
contains in general a great many statements of doubtful value, in spite
of the critical standpoint from which the author maintains to view the
facts.

2) Since Huber in his “Recherches,” p. 151, does not say whether
he means the fourmi fauve 4 dos rouge or that @ dos noir, we are
hardly able to decide which ant it is.

Wars and Slavery in the Animal Kingdom. 67

of ants as an “evident reaction against instinct,” as
Forel did, at least formerly, in his book mentioned
above. They are instinctive no less than the romps
and scuffles of young cubs, or the frolics of lambkins.
They are due, I suppose, to the natural impulse of
exercising the muscles, which is pleasurable to animals
as well as to human beings. At any rate it would be
wrong to ascribe them to intelligence, either in ants,
or in the higher animals.

The erroneous doctrines about the “absolute blind-
ness” and the “mechanical automatism” of instinct,
which are still current, have done a great deal towards
making animal intelligence appear almost indispen-
sable. Assuming instinct to be a mere reflex mechan-
ism, it was of course impossible not to declare as “‘intel-
ligent activities” all the manifestations of the instinct-
ive faculties, which are determined and influenced by
individual sense perceptions and sense experiences.
This, however, as we have shown above,' is an alto-
gether uncritical procedure. The study of ant life
enables us to determine more correctly the nature of
animal instinct, this being the case especially with
F. sanguinea, which in point of so-called “intelligence”
may fairly rank with the highest vertebrates.

4, Slavery among the Sanguine Slavemakers.

The custom prevalent among these red ants of rob-
bing and rearing slave pupae, is of course entirely
instinctive. Even our scientific opponents acknowledge
that it would be nonsense to explain it as an intelligent

1) “Instinct and Intelligence,” etc. (Herder, St. Louis, Mo.), Chap.
II and III. :

68 Chapter II.

invention of some colony of sanguineas, transmitted
by inheritance to all the descendants of the species.
Forel, Emery and Smalian fully agree with our tren-
chant condemnation' of Buechner’s manner of human-:
izing the “slavery” of ants. It does not seem impos-
sible, however, that for the actuation of this instinct
there should be needed special psychic impulses pro-
duced in the young ants by the example and the feeler
language of their older companions. Yet, this assump-
tion is scarcely probable; for the formation of new
colonies is undertaken, as a general rule, by single
impregnated females; but the females of F. sanguinea
are devoid of the enslaving instinct, and cannot, there-
fore, induce others to manifest it. Yet, since it is the
general opinion, that tradition and instruction aid the
exercise of the social instincts in these insects, and that
the high perfection of their community life receives
thereby its full explanation, we will examine whether
in view of the facts this opinion is still tenable.

It is true, in ant communities the instinct of imita-
tion plays a great part, as we may gather from several
observations recorded above. By the example and the
taps of the feelers of their older comrades the younger
ants are often induced to actions, which otherwise,
at least under the same circumstances, they would not
have performed. In this regard, as in fact in the other
salient features of the psychic life of animals, ants and
the higher animals agree in all the essentials; for,
in the latter also the so-called lessons given to the
young by their parents consist only in exciting instinct-
ively in the young the faculty of imitation by the exam-

1) “Die zusammengesetzten Nester,” p. 182.

Wars and Slavery in the Animal Kingdom. 69

ple of their parents. If we interpret “tradition and
instruction” in this sense, it must be acknowledged
that they aid in the exercise of the hereditary instincts
both in ants and in the higher animals. But, on the
other hand, it is equally obvious, that in this case the
terms “tradition” and “instruction” mean something
very different from what modern animal psychologists
wish to insinuate; for, in our case, they do not imply
any intelligent communication of knowledge, but only
the instinctive excitation of the imitative faculty.

But in the communities of social insects not even
the encouraging example of the older companions is
necessary for the first actuation of the young workers’
instincts. We have ascertained by experiments, that
precisely the most remarkable and apparently most
intelligent habits of the sanguine slavemakers, namely
their rearing of slaves and the hospitable care bestowed
by them on the beetle Lomechusa strumosa, are merely
hereditary instincts, for the exercise of which no kind
of “instruction” on the part of the older ants is
needed.’ To prove this we formed a special colony of
“self-taught” young workers of F. sanguinea, by plac-
ing in a glass filled with a sufficient quantity of earth
a number of ants that were newly developed from their
cocoons in my artificial nest. These self-taught ants
not only performed all the works required for building
their nest, just as the other individuals of their species,
but. they also followed the very same line of conduct
. in nursing their young and even in dealing with
strange worker pupae which I introduced into their

1) L. c., p. 202, and “Die internationalen Beziehungen von Lome-
chusa strumosa,” in the “Biologisches Centralblatt,” XII (1892), 592,

70 Chapter II.

nest. The pupae of Lasius niger they would either
devour or throw away, whereas those of F. rufibarbis
were reared by them as auxiliary ants for their colony.
A Lomechusa strumosa which IJ put in, was immedi-
ately received like an old acquaintance, licked and fed,
just as is the custom in the other colonies of F. san-
guinea. In the face of such experiments the beautiful
theory of tradition and instruction among ants van-
ishes into thin air.

That the older ants “lead their newly born com-
rades about the nest and train them to a knowledge
of domestic duties, especially in the care of larvae,”
is a fable originated by Buechnert and unfortunately
taken up on his authority even by Rorhnanes? and
other modern animal psychologists. The truth is, that
the newly developed ants are as yet the objects of spe-
cial care and protection on the part of the others, as
remarked already by Huber. Being as yet rather help-
Jess, they are still, as it were, considered as “wards.”

The same applies to bees. In their case also the
instruction said to be given by the old workers is a
mere fancy, arising in the brain of some anthropomor-
phizing observer. Already Réaumur in his classical
work Histoire des Insectes* remarks: “Scarcely have
all the parts of the body of a young bee become suf-
ficiently dry, scarcely is she able to move her wings,
when she is already acquainted with everything she
will have to do in the whole course of her life.” He
goes on to relate a few observations showing, that

1) “‘Geistesleben der Tiere,” p. 62.
2) “Animal Intelligence”? (6th ed.), p. 59. :
8) Tom. V. part II, mém. XI, p. 278. Amsterdam, 1741.

Wars and Slavery in the Animal Kingdom. 71

young bees from the very first day are as well able to
use their instincts as are their seniors. Of late some
experiments have been made by Kogevnikov' and
Butkewitsch? on self-taught young bees. The results
were practically the same as in the case of the self-
taught ants. It was found that in the workers the
building of combs and the nursing of the young, and
in the queens the love of combat were hereditary
instincts, utterly independent of experience and instruc-
tion. Besides Charles Janet’s excellent observations
on hornets* show, that social insects are ruled only by
hereditary instincts, excited to their natural manifes-
tation by the very first experiences of the young indi-
vidual. The impulse of imitation with its various
incitements is only a secondary factor. This is the
truth regarding the captious shibboleth of “instruction
and tradition” in insect communities.

No doubt, therefore, is left as to the fact that the
slavemaking habits and the military tactics of the san-
guineas, just as the social life of ants in general, are
due to instinct only, not to individual intelligence.
Yet, this instinct is not an absolutely blind impulse,
but is suitably modified according to the wants and
purposes of a given colony. A blind impulse to rob
and to rear slave pupae would be expected to impel
sanguinea colonies to rob the more slaves, the stronger
and more numerous they are themselves. In the most
populous nests we ought to find the greatest number

1) “Zur Frage vom Instinct,” in ‘Biolog. Centralbl.,” Vol. XVI
(1896), No. 18, pp. 657-660.

2) “Russisches Bienenzuchtblatt,” April,1896. | See Kogevnikov 1. c.

8) “Mémoires de la Société Zoologique de France.” T. VIII
(1895). .

72 Chapter II.

of auxiliary ants. In reality the very reverse happens.
The most populous sanguinea colonies do not contain
the relatively greatest but the relatively smallest num-
ber of slaves. We formerly (in “Die Zusammengesetz-
ten Nester” p. 50) alluded to this fact, which shall now
be explained and proved at greater length. In order
to show the connection between the rearing of Lome-
chusa strumosa in. the sanguinea nests and the educa-
tion of a strange, crippled kind of workers, the
so-called pseudo-females or pseudogynes,? I drew up
an accurate statistical map of the sanguinea colonies
in the neighborhood of Exaten. It comprises 410
colonies with more than 2,000 nests.? Regarding the
number of slaves, the statistics showed that in most
colonies the masters were from three to six times more
numerous than the slaves. The most populous colonies
contain scarcely 50 to 100 slaves, sometimes even less
or none at all. In the middle-sized or weaker colonies,
however, the absolute number of slaves amounts in
most cases to several hundred. The average propor-
tion of masters and slaves in the most populous
colonies is from 100:1 to 10:1, in the middle-sized
and weak colonies, however, from 3:1 to 1:1. Nor
are these the ultimate limits assigned to the number
of slaves found in the nests of these ants. In May,
1890, and from 1896 to 1898 I found near Exaten
several strong sanguinea colonies without any slaves.*

1) “Die ergatogynen Formen bei den Ameisen und ihre Erklae-
rung,” in “Biolog. Centralbl.,” Vol. XV (1895), Nos. 16 and 17.

2) A colony of F. sanguinea not unfrequently embraces several
nests, often one or more metres distant from one another, inhabited
all at the same time or alternately.

3) To similar colonies of F. sanguinea of the race rubicunda in
North America we must probably refer the F. sanguinea race aserva
of Forel, who described it lately from Toronto (Canada), (Ann. Soc.
Ent. Belg. XLV, 1901, p. 395).

Wars and Slavery in the Animal Kingdom. 73

A similar colony I recently detected near Luxemburg,
1904. On May 23, 1889, I met with the opposite
extreme, namely, a very weak sanguinea colony, in
which the slaves were about twenty times more
numerous than their masters. These extreme cases
are, however, very rare. Besides, it is plain that the
number of slaves in different colonies changes every
year; and lastly, the number of slaves in the nests of
the sanguine slavemakers depends also on special,
local circumstances. Where slave nests are very
numerous, e. g., in groves of birches and oaks, there
more slaves will be found in the sanguinea nests than
on the open heath, where fusca nests are very scarce.
Yet, ceteris paribus, we find the constant law, that in a
given sanguinea nest, the number of slaves and that of
masters is not in direct but in inverse proportion. My
observations of sanguinea nests in Dutch Limburg,
Rhineland, Vorarlberg, Bohemia and Luxemburg,
everywhere confirmed this law.

How is this remarkable difference between the col-
onies of sanguineas and of Amazons to be explained?
The latter possess the more slaves, the more populous
the colony; with the former we meet the reverse. This
difference is explained by the fact that F. sanguinea
is not, like the Amazons, essentially dependent on her
slaves, but rather regards them, as it were, as a
secondary complement of her own communities. The
sanguineas rob and rear only as many slave pupae, as
' is suitable for their colonies. Weaker colonies thus
feel greater need of supplementing their own deficiency
by adding auxiliary forces, whereas stronger colonies
do not feel the same necessity; so they regulate their

74 Chapter II.

action according to their perception of this deficiency.
It is true, the smaller number of slaves in more popu-
lous sanguinea nests may partly be due to another
circumstance, viz., that in the latter a greater per-
centage of robbed pupae is eaten than in smaller nests.
Nevertheless, this circumstance is far from explaining,
why in the weakest colonies of this marauding ant the
number of slaves even exceeds that of the masters.
The only way of accounting for this fact is to assume
that these colonies try to strengthen their forces by
the greatest possible number of auxiliaries.

To perceive this necessity of increasing their
numbers does not go beyond the limits of the
instinctive powers of ants. A very interesting case of
this kind was observed by me both in the summer and
the fall of 1898 in my artificial nest of F. sanguinea,
already repeatedly referred to. During my absence,
extending over several weeks of July and August, the
nest had been badly cared for and had repeatedly dried
up; consequently many workers of sanguinea and the
greater number of the old slaves had perished. Mean-
while new auxiliary ants (F. rufibarbis) had been
reared from cocoons which I had given to the
sanguineas. Now, I observed in the course of Sep-
tember, that new clusters of eggs, laid by the two
queens, made their appearance in the nest, and that a
number of larvae were reared, some of which were
conspicuous for their rapid development. This is an
exceptional case, generally not occurring in sanguinea
colonies during autumn. In October and the first part
of November I was absent again. On my return I
found, to my great surprise, that in place of the two

Wars and Slavery in the Animal Kingdom. 75

wingless queens there were now four of them in the
nest. This shows that the ants had reared two new
queens from the eggs which had been laid in the fall.*
The mortality prevalent in their colony during summer
must have caused this exceptional conduct. On duly
considering such facts, it can hardly be found strange
that sanguineas try to make up for the deficiency of
their numbers by robbing and rearing the pupae of
other ants; for the enslaving habit serves them as a
secondary means of increasing their colonies.

The manner, too, in which the sanguineas make
their choice between the different castes of alien
Formica pupae, confirms the above explanation; for
this phenomenon is understood only by referring it to
their desire to augment the forces of their colonies.
It is evident that the rearing of slave pupae is not
governed by a “blind instinct of education,” inducing |
the ants to transfer all their care to the strange brood;
for they consume the male and female pupae of
strangers, or kill the sexual ants as soon as they leave
their cocoons, whereas they adopt, at least partly, those
pupae of strangers, which will develop into workers.
They distinguish, therefore, by olfactory perceptions
between their own pupae and those of strangers, and
they likewise distinguish between the different castes
of the latter. Hence, we are compelled to maintain:
the sanguine slavemakers do not rear the worker
pupae of strangers, because they are unable to distin-
guish them from their own, but because they aim at
augmenting the number of their own workers by
~~ 1) The two old queens were still living in the spring of 1900, and
laid eggs in spite of their age of almost ten years. One of them died

last year (1903), thirteen years old. The surviving queen (fourteen
years old) I set at liberty with the rest of the workers in spring, 1904.

76 Chapter II.

adopting strangers as auxiliaries. Of course, they are
far from having an intelligent knowledge of this pur-
pose. It suffices that, on account of the actual need of
workers, the instinctive impulse of the ants to bring up
fresh workers is aroused with greater intensity and,
for this reason, extends to other Formica cocoons.
And this is the only explanation admissible, for we
have proved above, that with F. sanguinea slave-
holding is not due to experience or instruction, but to
hereditary instincts.

Is there anything in the social life of higher
animals, which can rival this strange phenomenon?
We know of nothing. If there had been, then Darwin,
Ziegler and other evolutionists would not have failed
to turn it to account, and to appeal to it as convincing
proof of the “quasi-human intelligence” of higher
animals; for, if an association of animals perceives
the necessity of increasing its strength by adopting
auxiliary forces, and under the influence of this per-
ception actually adopts them, then this action proceeds
from a motive originating in sensitive experience, and
is therefore intelligent, at least according to modern
animal psychology. Nevertheless, ants are said to be
“instinct automatons,’ but higher animals are not!
And this again shows, how utterly untenable, on the
one hand, is the modern notion of intelligence, and,
on the other, how foolish the attempt to place the
“intelligence” of the higher animals on a far higher
level than that of ants.

Bethe,’ indeed, has of late made an attempt to
explain, in a very simple manner, the proportion

1) “Duerfen wir den Ameisen und Bienen psychische Qualitaeten
zuschreiben?” p. 69.

Wars and Slavery in the Animal Kingdom. 77

between the number of masters and of slaves found
in colonies of F. sanguinea. “The correlation,” he
says, “existing between the numbers of masters and
slaves is as little owing to psychic processes, as the
numerical correlation existing between mice and
buzzards, or between certain butterflies and cuckoos.”
Yet, it is hard to see what is proved by this compari-
son, unless it be the very contrary of what Bethe
pretends to prove; for the more mice there are, the
more buzzards will come to the spot, and the more
butterflies, the more cuckoos; however, in the case of
sanguinea colonies just the reverse takes place, namely,
the more masters there are, the fewer slaves they have
in their colonies!

As the sanguineas accommodate themselves to
given circumstances regarding the number of their
slaves, so also regarding their species. Their favorite
slave species is F. fusca. This black ant is found as
auxiliary in the greater number of the above men-
tioned 410 sanguinea colonies near Exaten. In 25
colonies the place of F. fusca is taken by a different
species, viz., F. rufibarbis; 17 colonies have both
species. Near Feldkirch, in Vorarlberg (Austria), I
found side by side with colonies which had the above
mentioned slaves, others with F. cinerea, or with F.
fusca and cinerea. The latter species does not occur in
Dutch Limburg, and for this reason no cinereas are
found there as slaves in sanguinea nests. Yet, the fact
that the sanguineas occasionally invade weak colonies
of the large hill ants (F. rufa and pratensis) to rob
their pupae and to rear them as auxiliaries, proves
that their “blind instinct” does not force them to rob

78 Chapter II.

automatically a certain, fixed species of slaves. In
May, 1890, I found near Exaten such a “natural,
abnormally mixed” sanguinea colony containing,
besides F. fusca, a considerable number of F. rufa.
This colony has since disappeared; for on my return
after a two years’ absence I looked for it in vain.
Since 1895 I found in the same neighborhood of
Exaten four other natural, abnormally mixed colonies
of sanguineas. One of them (col. No. 66) had only
F. pratensis as slaves; the second (col. No. 105)
F. rufo-pratensis, a variety intermediate between rufa
and pratensis, with F. fusca; the other two had (col.
Nos. 84 and 247) F. pratensis and fusca. Three of
these colonies, therefore, possess besides the ordinary
species of slaves, an extraordinary one. In August,
1891, I came on the Arlberg pass (1,800 m.) across a
sanguinea colony that had rufas as slaves. Forel? has
long since recorded some very interesting instances of
natural, abnormally mixed colonies of these marauding
ants in Switzerland, namely a sanguinea nest with
F. pratensis as slaves, and another with rufa. The
sanguineas therefore display the same peculiar uni-
.versality and the same gift of suitable adaptation in
their enslaving habits both in Holland, and in Tyrol
and Switzerland; those qualities are due to that
specific nature of their sensitive cognition and appetite,
which we call “instinct.”

The above observations on these “natural,” mixed
colonies have made it plain enough, why the san-
guineas accept the worker pupae of different, alien

1) “S&tudes myrmécologiques en 1875,” p. 25 (57) and en 1886,
p. 9 (189).

Wars and Slavery in the Animal Kingdom. 79

Formica species and rear them as slaves even when
they are given to them by man. Forel’ and myseif
have made various experiments on this point
with artificial observation nests kept in a room and
with nests found in free nature. It may suffice to
mention one of them. In the summer of 1895, several
times in succession I took a large bag of worker
cocoons from a huge ant hill of F. rufa and emptied it
in the neighborhood of a densely populated sanguinea
nest,? which had but a few fuscas as slaves. In a few
minutes the sanguine ants had put to flight the thousands
of rufas contained in the bag with the cocoons and parts
of the nest, had snatched the cocoons from the mouths
of the fleeing rufas and began to ransack whatever I
had brought of the hostile nest. For hours after, hun-
dreds of these white “ant-eggs” were seen wandering
from the plundered nest to the den of the robbers and
mysteriously disappearing therein. By far the greater
number of the rufa cocoons were reared by the san-
guineas. This artificially mixed colony numbered, in
1896, about 5,000 sanguineas and 8,000 rufas. The
latter were generally busy building on the surface of
the nest and had soon given it the appearance of a
true rufa nest. At the least disturbance, however,
thousands of light-red sanguineas would dart out from
the interior to defend their common home; and thus
the supposed rufa nest was turned into a sanguinea
nest as if by magic. Because ants know no other home
than that in which they have developed from the
cocoon, these rufas, although they are in the majority,

1) “Fourmis de la Suisse,” p. 258 ff.
2) Colony No. 39 of the statistical map.

80 Chapter II.

will faithfully serve their ravishers and natural ene-
mies, without “reflecting” on how they happened to get
into this unusual society.

I have had in my room, for the last twelve years,
an artificial nest of F. sanguinea (see p. 23). This
colony adopted as slaves the workers of all the Formica
species to be found in Holland, viz., F. fusca, rufi-
barbis, rufa and pratensis. The “slaves” have devel-
oped from cocoons, which I had put into the nest
during recent years. In free nature the same Formica
species are found as slaves in the sanguinea nests,
but only one or two of them at a time; in this nest,
however, they were all united to form one colony
under the suzerainty of Formica sanguinea.

Yet, this “sugerainty” and “slavery” in the mixed
colonies of ants is altogether different from what the
same terms imply when applied to human society.
Only authors like Ludwig Buechner might be found
guilty of confounding ideas to such a degree. There
is perfect equality among all the workers of a mixed
colony, no less than among all the workers of a
simple colony. The very same “constitutional laws”
are in force both for masters and slaves; in other
words, the uniform “nest smell,” which adheres to all
ants reared in the same nest, serves them to recognize
one another as members of the same ant community,
the differences in species being totally disregarded.
The so-called slaves live entirely free in the nest of
their ravishers, that is to say, they live according to
the same innate instincts which would have formed
their rule of conduct at home; they work for their
ravishers, supply them with food and rear their off-

Wars and Slavery in the Animal Kingdom. 81

spring, as if they were in their own colony. They are
called “slaves,” only because they are reared from
robbed pupae, live in the nests of strangers and work
for them. On the other hand the sanguineas are
called “masters,” only for the reason that they have
robbed the pupae of an alien species, from which
their auxiliaries originate; and besides, because these
mixed colonies contain not workers only of F. san-
guinea, but also their males and females, whereas
the slave species is represented only by workers. This
is why in mixed colonies the propagation of the
masters is ensured but not that of the slaves.
Therefore it is downright nonsense for Buechner
to place slavery among ants and human slavery on
essentially the same level. By virtue of his intelli-
gence, man possesses the power of reflecting on his
origin and social position; he is gifted with self-
consciousness; accordingly he considers slavery as an
unjustified deprivation of freedom, a state of humilia-
tion, a degradation of his human dignity. With ants
it is different. They have neither intellect nor self-
consciousness, and are thus incapable of pondering
over the obscure question “whence” and “whither.”
As auxiliary ants they follow their social instincts
just as well as in a nest of their own species: they
are as free and independent as any other ant on
earth. Hence among slave ants there are no run-
aways, no revolutionists, no conspirators, no anarch-
ists. He, who seriously points to the complete social-
‘ ism and communism of mixed ant colonies as models
for human socialism and political economy, is sadly

in need of a nerve specialist.
6

82 Chapter II.

On the other hand, in opposition to those animal
psychologists who rank the “mental faculties” of the
higher vertebrates incomparably above those of ants,
it is necessary to emphasize the fact, that no associa-
tion of apes or other mammals can compare, as to
psychic faculties, with the mixed colonies of ants,
especially with the sanguinea colonies. The rearing
of the offspring of closely allied species as useful
members of their own society is an arrangement never
found with apes. The wars, therefore, and military
expeditions of ants addicted to this practice rank
_ much higher than the wars of baboons and other apes.
True, slavery among ants is based only on instinct,
not on intelligence. But anything higher than instinct
is not found in the societies of higher animals either.
In fact, the development of their social instincts is
rather far inferior to that of ants.

5. Other Wars and Alliances of Ants.

The slavemaking expeditions of the Amazons and
sanguineas are indeed the most interesting; but by no
means the only wars waged by ants. There are many
other feuds and skirmishes, both between ants of dif-
ferent species and between different colonies of the
same species. Most of these feuds are caused by dis-
putes about subterranean or open-air boundaries, call-
ing for settlement “at the point of the sword.” When
underneath a large stone there is a “compound ant
nest,” i. e., when two or more different species have
built their respective nests in close proximity, they are
separated by walls of earth. No one ventures into
the neighboring realm, and woe to him, if he does;

Wars and Slavery in the Animal Kingdom. 83

he is seized and put to death. As a rule, only the
dwarfish, yellow, thieving ants (Solenopsis fugax in
Europe and S, molesta in N. America), may pilfer in
the neighboring nest of a larger species. The small,
black lawn-ant (Tetramorium caespitum, in Europe
and N. America) will occasionally do the same. These
thievish little ants are in the nests of larger species,
what rats and mice are in the abodes of man.

To return to the “compound nests.” By turning
over the stone, under which several different ant
species live side by side in separated nests, the par-
titions are suddenly removed, and a fierce battle
ensues with great loss of life on both sides. The
engagement frequently continues for a considerable
time after the stone has been replaced. Only after
the boundary-lines have been perfectly restored, is
there again peace between the neighboring states.
Sometimes on such occasions it becomes evident that
one colony is numerically far superior to the other.
In this case we notice that the weaker is simply driven
out of his nest, which is then entirely or in part occu-
pied by the victor. Thus the war ends with the
“territorial expansion” of the more powerful state.

Above-ground disputes between neighboring ant
colonies, not infrequently lead to fights, which last
for weeks or even months, interrupted by longer or
shorter periods of “armistice.” These battles are most
obstinate and bloody between different colonies of the
. small, black lawn-ants (Tetramorium caespitum).
This species is found everywhere, frequently with
hundreds of thousands of earth-nests within a square
mile. On July 8, 1886, I saw a regular battlefield on

84 Chapter II.

-a sandy road near Exaten. It fairly swarmed with
fighting lawn-ants. The combatants numbered thou-
sands and they covered a space of about 70 cm. by
8 cm. So dense was the battle-array that individuals
could scarcely be distinguished in the mass of war-
riors. They formed irregular clusters of from 2 to 14
individuals all clinging together with their mandibles
and making liberal use of ‘their stings. The summer
heat had inflamed the rancor of the two tribes, long
living too close together. The battle probably ended
with the expulsion or the utter extermination of one
of the communities.

Among men civil wars are generally the fiercest
and bloodiest. The same may be said of the wars
waged between different ant colonies of the same
species. However, only the “heat oppressed brain” of
Buechner or Brehm could detect a closer analogy
between these phenomena. As the males of certain
birds fight for their breeding districts, nor allow
other families of the same species to settle there,’ so
ant colonies are wisely compelled by the laws of nature
to regard the district about their nest as exclusively
their own, on which no other colonies of the same
species are suffered to encroach. Otherwise, their
wants being equal, their food supplies would become
scarce. Hence arises an instinctive hatred between
different colonies of the same species; whereas col-
onies of different species whose mode of life and
means of sustenance are different are admitted much
more easily. The preservation of the species neces-
sitates the fiercest struggles for existence between

1) Altum, ‘Der Vogel und sein Leben,” (6th ed.), p. 128 ff.

Wars and Slavery in the Animal Kingdom. 85

tribes of the same species. Not inordinate greed in
the individuals nor imperialistic tendencies in the tribe,
but higher, natural laws are the mainsprings of these
“civil wars” among ants. The poet indeed, may
exclaim: “There is room on earth for all’ (Schiller) ;
but even in the life of ants this beautiful saying is
often correct only in theory.

Many more accounts of wars and battles among
ants could be mentioned; but we cannot enter upon
them here, since our principal purpose is to call atten-
tion to a few points of comparison between the “intel-
ligence” of ants and that of higher animals and of
man. It remains only to be stated that the wars of
ants sometimes end in an “alliance,” that is to say in
a peaceful union of the combatant tribes into one con-
stitutional body. These alliances are usually formed
between Formicas of the same or different species,
but are most frequent between different colonies of
sanguineas. From Forel’s “Fourmis de la Suisse”
and from my own observations (see Die zusammen-
gesetzten Nester p. 146-157) many instances might be
selected. The chief conditions for such alliances
between hostile ant colonies are, that the two oppo-
nents be closely allied in species, that they be almost
equally populous, and lastly that they be forced to
live in close proximity and are thus unable to avoid
each other. Under such circumstances their original
skirmishes give way to mutual toleration and finally
to friendly intercourse. A superficial observer, of
such occurrences, might be led to believe that intelli-
gent reflection had caused the animals to overcome
their instinctive aversion. He might conclude that

86 Chapter 11.

ants reason thus: ‘Why this useless shedding of
blood? Let us not exterminate each other but live in
peace; our differences are not so great that we can-
not come to terms!” Yet, there is not a shadow of
proof that ants entering into an alliance reason in this
manner. The phenomenon, which is indeed singular
enough, can be explained more simply and naturally
from the laws of instinctive sensation, with special
regard to the feeler sensations. Especially with the
Formica species, and among these, most of all with
the highly endowed F. sanguinea, the hereditary dis-
position of the sensitive powers of cognition and appe-
tite is so plastic, that with parties of almost equal
numbers fear will be stronger in such cases than
love of combat. First, of course, by tapping one
another with their feelers they find out that they are
strangers, and therefore they try to avoid each other;
but if this is impossible, the perception of mutual
similarity will gradually prevail over their mutual dif-
ference. In the beginning, they live together from
necessity only, but they gradually acquire a common
nest-smell which unites them as members of one
colony. From this time, by taps of their feelers, they
recognize one another as belonging to the same house-
hold. The former opponents have united into one
“constitutional body” which is kept together by the
common nest-smell. Strange though this mode of
communication may appear to us who are not pro-
vided with antennae, it alone explains the fact other-
wise wholly inexplicable, how the confederate colony
thus formed will in future hold together even against
former members of their own colony.t An example
~~ 4) On the explanation of the nest-smell see above p. 16 ff.

Wars and Slavery in the Animal Kingdom. 87

of this kind is related by Forel in his “Fourmis de la
Suisse.’ One day he brought a handful of F. pra-
tensis to a confederate colony of sanguineas and pra-
tensis; the pratensis of this colony, two months
before, had been taken from the same pratensis nest
to which the new arrivals belonged. What happened ?
Immediately the new pratensis were fiercely attacked
by the sanguineas, because they were recognized as
enemies by means of the “smell at contact.”? The
old pratensis seemed to recognize their sisters but
faintly. They met them with suspicion and did not
assist them, although on the other hand they did not
take part in the fight. But soon they began to carry
the new arrivals into the confederate nest, as if they
belonged to it. The number of the pratensis was thus
increased considerably, outnumbering even the san-
guineas. Although the latter continued their hos-
tilities against the newcomers for several days, and
mutilated and killed several of them, it never occurred
to the old pratensis to make common cause with their
maltreated “sisters” against their natural enemies.
They allowed the sanguineas to have their own way,
until the survivors had gradually acquired the nest-
smell of the confederacy. In the course of a week
“peace” was restored, and the strangers were treated
henceforth, also by the sanguineas, as inmates of the
same colony.

If ants had the power of rational reflection, if they

1) Both had been taken from the pratensis nest when fully de-
veloped and not as pupae. Otherwise this example would be out of
place when speaking of “allied colonies.”

2) By this term Forel expresses very well the peculiar sensation of
ant feelers,

88 Chapter II.

had any idea of consanguinity, then this behavior of
the old pratensis of that confederate colony during the
maltreating of their sisters would be altogether inex-
plicable. On the other hand, the instinctive nature
of ant sensation will furnish a very satisfactory expla-
nation of this phenomenon, which is in evident con-
tradiction with animal intelligence. Yet, it should
not be forgotten, that societies of apes and other
higher animals have nothing to compare with the con-
federacies of ants. No one has as yet observed, that
wars carried on between different hordes of apes
ended with a peaceful alliance between the combatants.
This chearly shows, how wrong it is to exalt the
societies of higher animals above those of ants in the
matter of psychic endowments.

Evolutionists, therefore, such as Darwin and Zieg-
ler, are sadly mistaken when they point to the battles
sometimes waged between hordes of apes, and adduce
these as conclusive evidence, that the societies of
higher animals are so closely allied to the “primitive”
societies of man, that a little, unimportant “step”
bridges the difference; for first they imagine a
“primitive state’ of human society, which is depicted,
of course, as brutal and as devoid of reason as
possible; then, to match the picture, they exalt the
societies of higher animals to the greatest possible
similarity with reasonable man, and finally, from this
twofold hypothesis they draw the conclusion that
human society has evidently developed from the ani-
mal societies. And this is called the “consistent,
scientific application of the theory of evolution to
man!” If ants were endowed with reason and risi-

Wars and Slavery in the Animal Kingdom. 89

bility, they would surely burst into a hearty laugh at
these evolutionistic “steps”; for, as to the develop-
ment of social instincts, ant colonies bear a far closer
resemblance to the human societies than the hordes of
apes; and yet even the intelligence of an ant would
be sufficient to understand, that animal and human
societies*are as far apart as heaven and earth. The
difference between ant states and human societies is
readily acknowledged; but the difference between
hordes of apes and the primitive states of man cannot
be conceded, because, forsooth, it is against the theory
of evolution!

CHAPTER III.
ARCHITECTURE IN THE ANIMAL KINGDOM.

1. A General Survey of the Building Activity of
Animals.

HE houses of animals are exceedingly simple and

destined for very prosaic purposes; they merely
serve the wants of daily life, the preservation of the
individual and of the species. To their owners they
are necessary helps in the struggle for existence; they
never aim at art for art’s sake.t This clearly shows,
that in the animal kingdom we can speak only meta-
phorically of architecture properly so called. There
is mere mechanical skill, but not art; and if some-
times its productions bear a faint resemblance to works
of human art, the aesthetic effect is never either intended
or understood by the animal. Another essential point of
difference between the artistic skill of animals and of
man is in this, that with animals it is due to an innate,
hereditary aptitude which has not first to be acquired,
as is the case with man. At its birth the animal is
endowed with all its artistic talents. It applies them
without previous experience or instruction, as soon as
demanded by its organic development and by external
circumstances. The caterpillar of the emperor moth
(Saturnia) begins to be an artist only, when the time
has arrived to transform itself into a chrysalis, and to
weave a bottle-shaped case wherein it is to undergo

1) The buildings of the Australian Tectonarchinae are no exception
to this rule, if we divest descriptions of them of all poetical additions.

Architecture in the Animal Kingdom. 91

‘the change. The female of the leaf-rolling beetle
(Rhynchites betulae) becomes an artist, only when, the
time of depositing her eggs having arrived, she is
forced to cut and to roll up a birch-leaf in the shape of
a graceful funnel. The male of this beetle is no more
endowed with this wonderful, technical skill, than the
emperor moth in the state of imago is able to spin a
cocoon; for the artistic talents of animals depend on
their organico-psychic constitution. Their application
affords no trace of reflecting reason or free choice,
because their mode of action is predetermined to the
most minute detail by the laws of vegetative and sen-
sitive life. Again, animals by their very organic con-
stitution possess all the tools necessary for exercising
their natural talents. Bees have their honey-bags to
gather in the pollen and daggers to ward off their
enemies; silkworms have their spinning glands,
beavers use their tails as trowels, and their sharp
gnawing teeth serve them as axe and chisel in work-
ing the wood. There is no need therefore, to invent or
manufacture implements. Moreover, directions for
using their natural instruments are furnished to ani-
mals by the innervation of their bodily organs and by
the corresponding constitution of their sensitive powers
of cognition and appetite. Reason and free choice,
therefore, are utterly superfluous in the exercise of
these animal talents. Since by their organico-sensi-
tive constitution animals are provided sufficiently with
all the necessaries of life, they are denied the higher,
spiritual faculties. Hence, they, who attribute reason
to animals, show that they have but a superficial
knowledge of animal life.

92 Chapter ITI.

The buildings of animals either serve to shelter
the individual, or else they are places for breeding
and rearing the young. To the former class belong
the tunnels excavated in the earth by the serpulas, the
envelopes made of various substances by the moth-
caterpillars and the larvae of the may-flies, as also
the different casings constructed by the larvae of
insects, particularly of many butterfly-caterpillars
before their metamorphosis. To the latter class
belong the regular nest constructions of animals. The
most primitive specimens are found with the parasitic
Nemertine worms. In several orders of insects,
especially among the Hymenoptera? and the beetles,
we meet with instances of ingenious and manifold
development of the same art. Here we find the most
various forms of nests, and made of all kinds of
material. Those elegant little domes of mortar, the
wasps of the genus Eumenes have built for their
offspring. Those breeding burrows, lined with red
poppy blossoms, have been’ excavated by the so-called
tose-bee (Megachile). Those graceful funnels and
barrels of leaves have been rolled into shape for their
young by weevils (Rhynchites, Apoderus, Attelabus).
And that boat adorned with streamers has been spun
by the great water-beetle (Hydrophilus piceus), as a
receptacle for its eggs. Among fishes nest building is
tare. We find examples of it, in the stickleback (Gas-
terosteus aculeatus), and in other fishes provided with
spines. On the other hand, birds are unsurpassed in
the art of nest building as regards variety, both of

1) See “Naturforscher,” 1886, 19th year, No. 50, p. 494.
*) See especially J. H. Fabre, “Souvenirs Entomologiques,” who
has described these buildings with admirable skill.

Architecture in the Animal Kingdom. 93:

form and of the material used by different species.
With the mammals,! finally, nest constructions are, on
the average, far less complicated and artistic than with
birds and insects.

The buildings which serve to shelter and rear the
young, may likewise be used as permanent lodgings
for parents and offspring. This is the case with social
insects and many mammals. Thus the nest develops
into a family dwelling. Only in relatively rare
instances, do animals employ their building skill in
providing other necessaries of life. Many spiders spin
their webs not only as a hiding-place for themselves
or for breeding purposes, but they also, by means of
their spinning glands, manufacture nets wherewith
to catch their prey. In like manner the neuropterous
larva, which goes by the name of ant-lion, uses its
earth-funnel both as a dwelling place and as a trap for
catching its prey, which consists chiefly of ants or
other insects. Among ants, however, we find the most
varied and manifold application of natural architec-
tural skill.

2. The Nests of Ants.

In the first place, ants use their architectural talents
for building nests, in the strict sense of the word.
Everybody is more or less familiar with ant nests, but
few are aware of the immense variety of forms implied
in that apparently simple term. As there is scarcely
any material unfit for an ant nest, so it may assume
all possible shapes and be found in the most unlikely

1) We shall consider more in detail the buildings of the beavers in
the subsequent pages.

94 Chapter ITI.

localities. Here it is the size of a thimble, there, the
pyramids of the ancient Egyptians are like mole-hills
in comparison, if we take into consideration the rela-
tive size of the builders. Some are in the ground, in
clefts of rocks, or concealed by stones, others are
under the bark or in the wood of trees. Others again
are in the hollow stalk of a plant, or in a gall-nut or
in a deserted snail-shell. Now they hang high in the
boughs of a tree, now in forests they rise as domes
from the level of the ground. Such a nest may be
dug, or spun; it may consist of masonry, or of cavities
hollowed out of the earth or of the wood. Sometimes
all these modes of operation enter into the same con-
struction. In short, the variability as to form, style,
or locality is almost unlimited. There is one charac-
teristic, however, common to all ant nests, viz: the
absence of any uniform architectonic pattern; ant
nests are irregular systems of chambers and galleries,
giving shelter to the ants and their offspring, and
communicating by different openings with the outside
world. This very irregularity of their buildings
enables the ants to suitably adapt their nests to any
locality and to employ any kind of material in their con-
struction. The artificial and, as it were, mathematical
regularity of the honey-combs of bees? is entirely

1) N. Ludwig, in an essay, “Der Zellenbau der Honighbiene,” (in
“Natur und Offenbarung,” 1896, 10th issue, p. 598 ff.), has offered a
new explanation of the hexahedral form of the bee-cell and of the three
congruent trhombs forming its pyramid-like base. In his opinion the
peculiar form of the bee-cell is due only to the construction of the
wax combs, each cell being built only in connection with other cells.
For the bees are actuated by the impulse of combining round cell-walls
bordering on one another, into one single wall and to reduce their
thickness by gnawing off both sides as much as possible without peril
to their necessary strength. Hence, the flat walls of the form described

Architecture in the Animal Kingdom. 95

wanting. This difference is very important for com-
parative psychology. As in the social life of ants
the individual independence of the single workers
attains a higher degree than amongst bees, so in their
architecture the same phenomenon may be observed.
Instead of constant uniformity we find great variety;
instead of the monotonous “automatism” of innate
instinct, we meet with a quasi-intelligent arbitrariness
in the exercise of their sensitive cognition and appe-
tite. Scarcely anywhere else is the wonderful plas-
ticity of animal instinct manifested so clearly and con-
vincingly as in the architecture of ants. This was the
reason why ants seemed to furnish such numerous
proofs of the “individual intelligence” of animals;
and, in fact, im the architecture of ants, if anywhere
in the whole animal kingdom, we find a striking
resemblance to human intelligence.

In order to give a full psychological description of
ant architecture, we shall first, compare the nests of
different species, secondly, we shall consider the differ-
ences in the buildings of one and the same species; in
the third place, we shall examine the methods by which
ants of the same colony co-operate in building their
nest, and finally, we shall investigate the various pur-
poses to which ants apply their architectural talents.
However, it would require a volume of considerable
bulax to treat these points exhaustively. Therefore, we

above. The consequently elongated, prismatic form of bee-cells would
be merely the result of the extreme uniformity in the working of the
bees engaged in building the combs. Single cells are built by the same
bees always in a cylindrical form with hemispherical top and bottom.
This form is to be regarded, according to Ludwig, as the proper,
primitive type of bee-cells, which is found with bumble-bees and other
allied Hymenoptera.

96 Chapter ITI.

must confine our discussion to its narrowest possible
Simits, touching chiefly on such features as are of
special interest for comparative psychology.

Great as is the variety of ant nests, still, every
species has its peculiar architecture, differing more or
less from that of any other species. Many ants, e. g.,
our small, blackish garden ants (Lasius niger) and the
small, yellow meadow ants (Lasius flavus)* work
almost exclusively in earth. Their nests are dug in
the ground, but above the subterranean nest they raise
smaller or larger domes of earth, the stalks and blades
of grass, that grow on the spot, serving as natural
pillars and beams. Other species, again, e. g., our well-
known hill ants (Formica rufa)? build so-called “ant-
hills,’ the popular type of ant nests in our northern
hemisphere. These ant hills may be termed mixed
buildings, an under-ground earth nest being combined
above ground with a dome consisting of earth, pine-
needles, scraps of dry leaves and stalks, and other parts
of plants. The different ant species which build such
ant hills follow systems and styles peculiar to each.
Thus any one with a little practice is able to determine
at once the species of the builders. F. rufa builds
differently from pratensis, pratensis from e-xsecta,
exsecta from sanguinea. The universal tool which,
like the human hand, is fit for and skilled in a

1) Both are found also in N. America; the most common yellow
ant there is L. aphidicola Walsh; L. niger is represented by its very
common N. American race L. americanus Em,

2) The N. American species of the rufa group are very numerous;
among them the nests of F. exsectoides For. and obscuripes For. are
most like the European ant hills of F. rufa.—An interesting descrip-
tion of N. American nests is contained in a paper of Father H. Mucker-
mann, S. J., entitled ‘‘The structure of the nests of some North Ameri-
can species of Formica.” (“‘Psyche,” June, 1902.)

_ Architecture in the Animal Kingdom. 97

variety of performances is found in the jaws (mandi-
bles) of the ants. Of course, in digging burrows in
the earth and in constructing earth-works they are also
assisted by their fore-legs, which help partly to scrape
up the sand and partly to hold down and fasten pellets
of earth. In closely allied species the shape of these
instruments, and especially that of the all-important
toothed inner edge (cutting edge) of the mandible,
is as a rule so similar* that the specific differences in
architectural style can be accounted for only by the

Fig. 2. Fig. 3.
Left mandible of Formica rufa. Right mandible of F. sanguinea.
orker.) (Worker.)

instinctive preferment of a particular style on the part
of different ant species. In the case of ants, therefore,
it will never do to resort to the mechanical automatism
of animal activities, and to explain the differences of
instincts merely by differences of bodily organs. The
decisive factpr is the psychic variety of instinctive dis-
positions. By them the bodily organs, in themselves
indifferent, are directed in their various modes of
operation.

It is true, to a certain extent, that the nature of

1) See the subjoined cuts. Both are drawn with the Zeiss’
microscope, syst. A,, and Abbe’s Camera lucida.
7

98 Chapter III.

the exterior organs of ants will also decide the nature
of their architecture. Thus e. g., the large Camponotus
ligniperdus (horse ants) and their allied species pos-
sess larger workers, whose huge head and strong
mandibles enable them to cut galleries in the wood of
decayed or even of sound trees. And therefore these
species are remarkable for wood nests. Others again,
among them the jet black Lasius fuliginosus as the
only one of this kind among the emmets of northern
Europe, build paper nests by gnawing wood-fibre and
gluing it together with the sticky product of their
salivary glands. They thus produce a coarse, brown
papiermaché, in which they establish their nests. Far
more perfect are the paper nests made by several
foreign ants, especially in South America, Madagas-
car and East India. They resemble irregular, brown
or grey-colored wasp-nests, suspended from or fast-
ened between branches of trees. Rev. A. Schupp,
S. J., sent me from Porto Alegre (South Brazil)
several paper nests of Cremastogaster sulcata, one of
which on arriving in Holland still contained several
thousands of live inhabitarits. Similar nests of Cre-
mastogaster Schenki in Madagascar are reported by
Sikora to be sometimes of such size as to accommodate
a full-grown man. From these paper-nests we must
distinguish nests which are spun and do not consist of
a paper-like material but of a texture like cobwebs.
Such webs are constructed, according to Wroughton’s
observations,| by an East Indian ant Polyrhachis
spinigera for lining her earth-burrows. Other Indian
and Australian ants of the genera Oecophylla and Poly-

1) “Our Ants,” part I, p. 25 (“Journal of the Bombay Nat. Hist.
Soc.,”’ 1892). - %

Architecture in the Animal Kingdom. 99

rhachis build their nests on trees by sewing together
clusters of leaves. It is now finally ascertained (see
p. 128-129), whence the Oecophylla procure the
material for their threads. They do not secrete it
from their mandibular glands, as was hitherto sup-
posed,’ but they use their young larvae as “spinning-
wheels,’ moving them to and fro between the edges of
the leaves. This strange fact observed already by
Ridley and Holland? has recently been confirmed by
other scientists.

Another class of ant nests, likewise found in the
tropics exclusively, are the natural cavities in the
stems, thorns and bladder-shaped swellings of the so-
called “ant-lodging plants,’ which invite ants to take
possession of their well furnished lodgings. Several
of these plants, as the American Imbauba (Cecropia
adenopus) offer to the ants, besides the lodging, also
an agreeable food in the form of special honey-bearing
nectaries. In return for their kindness the “ant-lodg-
ing plants” are afforded by their valiant lodgers effec-
tive protection against leaf-cutting ants and other
herbivorous insects. This mutual relationship of the
ants with the plants in question is called Symbiosis
(consociation). It bears, in fact, some similarity to the
associations existing between animals of different

1) E. H. Aitken, “Red Ants Nests” (“Journal of the Bombay Nat.
Hist. Soc.,’”? 1890, Vol. V, n. 4, p. 422), also Forel, “Die Nester der
Ameisen” (Zuerich, 1892), p. 19.

2) E. E. Green, “On the Habits of Oecophylla smaragdina F.”
(“Proceedings Entomol. Soc. of London,” 1896, p, IX.)

8) Fritz Mueller, “Die Imbauba und ihre Beschuetzer’’ (‘‘Kosmos,”
VIII, 109), and A. F. W. Schimper, “Die Wechselbeziehungen zwischen
Pflanzen und Ameisen im tropischen Amerika’ (Jena, 1888). Also
H. v. Jhering, “Die Ameisen-von Rio Grande, do Sul,” in “Berl.
Entom. Zeitschr.,” 1894, 3d issue, p. 354 and 364 ff.

100 Chapter ITI.

species, e. g., between ants and their guests, such as
the club-bearing beetles (Claviger), the tufted beetles
(Lomechusa, Atemeles), etc. But even those cavities
of plants, which are properly not meant to receive and
to lodge ants, are often occupied by them, especially
_ in the luxuriant vegetation of tropical South America.
Aug. Forel in the winter of 1895 and 1896 visited
the savannas of Columbia, where he found that the
nests of by far the most of the species, belonging to
eight different genera, were built indry stalks of grass.*
This led him to think that in the prairies and forests of
tropical America the nests in stalks and in hollow parts
of plants were the typical form of antnests, correspond-
ing to the climate of that country, whereas in our
zones the usual type is the earth nest or else the hill
made of earth and parts of plants.

This cursory comparison of the various forms? of
nests met with in different ant species, shows clearly
enough, that their character is conditioned by the
peculiar shape of the bodily organs of the builders , but
far less than is the case with most of the other artistic
instincts in insects and other animals. The form of
the mandibles, the presence of salivary glands with
gluey secretions or of real spinning glands, indicate
only the general outline of the architectural style
preferred by their owners. Only the different instinc-
tive dispositions of the builders determine more
exactly the specific differences of their nest forms.

1) “Quelques particularités de l’habitat des fourmis de 1’Amérique
tropicale’” (Extr. des Ann, de la Soc. Entom. Belg., XL [1896], 167 ss.)
and “Zur Fauna und Lebensweise der Ameisen im columbischen
Urwald,” in “Mitteil. der Schweiz. Entomol. Gesellsch.,” IX, 9th issue,

2) Forel, “Die Nester der Ameisen,” Zuerich, 1892.

Architecture in the Animal Kingdom. 101

Within the limits of these instinctive, hereditary dis-
positions there is plenty of room for the worker ants
to give full play to their individual powers of sensitive
cognition and appetite. This is why certain ant spe-
cies, particularly those skilled in earth work or wood
work, often take possession of some spot so suitable
for their dwelling as to be practically half-finished.
This they fit up in a becoming manner as a home for
their colony. Such attractive spots are e. g. on the
heaths of northern Europe rotten tree-stumps, in which
for years numbers of bark-beetles and their larvae or
other wood-boring insects were kind enough to prepare
comfortable quarters by carving out an extensive sys-
tem of galleries and chambers. It only remains for
the ants to take possession of the lodgings, devour
the former inhabitants, if there be any left, clean the
- apartments, close up the superfluous entrances with
earth or rotten wood, and with the same materials
construct, if need be, a few partitions for separate
chambers. If, on occupying the residence, they should
perchance discover that part of the stump is already
inhabited by another ant colony, the latter are killed
or turned out without much ado. If they should fail
in this, they make peace and live on good terms with
the rivals, especially if the latter are equal to them in
fighting strength, but differ greatly in size and means
of defense. The nests are separated by partitions, and
the stump is henceforth inhabited by a “compound
nest.”

Such “stolen nests” iorm a large category of ant
nests, particularly in places where there are many
stones; for almost all ant species that dwell in earth

102 Chapter ITI.

nests, have a predilection for building under stones.
This saves them a great deal of work and gives the
whole building greater firmness, and, moreover, in
. such a nest the heat of the sun more easily penetrates
to the interior. In heaths also, where stones are rather
rare, instances of such stolen nests may occur. A nest
of F. sanguinea,’ on which in 1894 I had placed a clod
of heath serving it henceforth as roof, had in 1895
passed into the possession of a colony of Lasius niger ;
in the years 1896-98, it was again inhabited by F. san-
guinea. A short time ago, in the same region near
Exaten, I found a rather extensive earth hill supported
in the centre by a bunch of heather and inhabited by
a large colony of F. rufibarbis. The ants had collected
on the surface a small heap of dry heather-leaves, as
they generally do there. The earth hill itself, how-
ever, judging by its architecture, was an old nest of
L. niger, which subsequently had been occupied by
the rufibarbis. History is silent as to whether the orig-
inal builders of the nest had quitted it before the time
_of the foreign invasion, or whether they were com-
pelled by force to evacuate it.

F, sanguinea are a restless people. They frequently
desert their nests, which are then taken possession of
by other, smaller species (especially Tetramorium
caespitum, L. niger and alienus). I have noted a num-
ber of such instances in my records of the last few
years; it may suffice to mention one of the most
remarkable. A large colony of F. sanguinea (No. 72
of my stat. map), in 1895 and 1896, had inhabited a
group of three nests, distant from one another 3 and

1) Colony No. 155 of the statistical map.

Architecture in the Animal Kingdom. 103

ym. respectively. Since the spring of 1897, however,
they had emigrated and I did not find them again.
In 1898 the northernmost of these nests was empty,
the central one was occupied by L. niger, and the
southernmost by Tetramorium caespitum. This was
the state of affairs until July, 1898. When I returned
on July 14, I found that the sanguineas of colony No.
72, which I could easily recognize by the size of their
workers, had returned to the southern nest which con-
sisted of two little heaps in close proximity. From one
of these the Tetramorium had already been completely
driven out; in the other they still occupied a retired
corner and were surrounded by the sanguinea nest.
Therefore ant nests also have their fates.

Ant nests are stolen not only by ants of different
species, but sometimes by those of different colonies
belonging to the same species. Many instances of this
kind are furnished by the sanguinea colonies in the
neighborhood of Exaten; but for brevity’s sake, it
must suffice merely to mention the fact.

3. The Nests of the Sanguine Slavemakers.

The great plasticity and power of adaptation to
given circumstances in the nest-building instincts of
ants is seen to the greatest advantage by an examina-
tion of the nests of the sanguine slavemakers (F. san-
guinea). With members of one and the same species
possessing the same specific, natural constitution, there
is such a variety of nest construction, that there is no
trace of that “automatism” of instinct, which postu-
lates a completely uniform and monotonous manifes-

104 Chapter III.

tation of the hereditary instinctive activities. As
these ants, in the parlance of modern animal psychol-
ogy, possess a high degree of “individual intelligence,”
because under the influence of their sense-perceptions
and sense-experiences they, are able to adapt their
innate, instinctive dispositions and aptitudes to any kind
of circumstances, so they manifest great adaptibility in
the building of their nests. I have drawn up statistics
of the sanguinea colonies in the neighborhood of Exa-
ten, which show that to my knowledge there are in
this region about 2,000 nests of this ant species,
embracing 410 colonies. The architectural style of the
nests varies greatly. By far the majority of them
are underground, built either below the bare surface
or under a shrub of heather, beneath a loose clod,
under a stone or at the foot of a tree. In connection
with this underground earth-nest there is generally
on the surface a greater or smaller heap of dry leaves
collected from the heather shrubs. This heap, together
with the earth carried out of the interior galleries and
the twigs of the shrubs supporting the whole construc-
tion, forms a sort of protective dome. With large nests
this hill sometimes has a circumference of several
meters and a height of several decimeters (e. g., in
colonies Nos. 208, 216, 118); but often it is rather
insignificant and sometimes it is altogether wanting.
Besides these simple or mixed earth-nests, F. san-
guinea builds also in rotten stumps of fir-trees or oaks,
now under the loose bark, now in the wood, now in the
roots. Sometimes the old stump is surrounded by
earth-galleries, and sometimes the whole nest is con-
fined to the stump itself. The nest of one of our

Architecture in the Animal Kingdom. 105

sanguinea colonies here (No. 112 of stat. map) is
built in a lofty and sturdy oak, in the mould and the
clefts of a hollow in the tree, almost a yard above the
ground. These observations show that the sanguine
ants are able to choose the most peculiar places for
their nest and to adapt its construction to any local
circumstances. This is confirmed by the following
observation. In the immediate neighborhood of Exa-
ten the earth-nests prevail, while two kilometers further
or near the village of Grathem, the greater number of
nests are built in rotten fir-stumps, because in that
region the ground is more turfy and thus renders the
construction of earth-works rather laborious.

Not less variable than the style is the number of
nests constituting a colony of F. sanguinea. Among
the above mentioned 410 colonies which are known
to me in this region, there are but a few that have only
one nest. These are mostly weak tribes which, on
account of the scarcity of members, feel no need of
other nests. In some cases, however, even a very
strong colony has only a single nest built in a specially
convenient place, generally at the foot of a fir
(e. g., col. 208 and 216). In such cases the concen-
tration of the building is of greater advantage than
its division into a number of different nests.
However, by far the majority of the sanguinea colo-
nies have several nests, averaging from two to eight,
either close together or farther off from another, some-
times inhabited simultaneously, sometimes by turns.
The distance between the nests of one colony is mostly
only from 4 to 4 m., but sometimes from Io to 20 m,
or more. The latter is particularly the case with their

106 Chapter ITI.

summer and winter residences, or rather with the
abodes for winter and for spring; for, many but by
no means all sanguinea colonies of this country have
special winter-quarters established in thickets under
the roots of trees or stumps and affording deep and
warm recesses for the cold season. The spring resi-
dence, however, which again often consists of several
single nests, is generally built near the edge of the
thicket. Here, on the first warm days of March and
April, the ants can always be observed changing their
quarters and moving over with the whole family to
the spring residence. In September or at least in the
beginning of October they change again, moving in
the opposite direction.

If it should become very hot and dry in summer,
the colonies move to their winter quarters during the
dog-days, thus converting the winter nest into a mid-
summer nest. When at the end of August, 1898, I
had returned after several weeks of absence, I found
that during the exceptionally hot days of August most
of the sanguinea colonies of this region had aban-
doned their spring nest. What had become of them?
As I was well acquainted with the winter nests of many
colonies by the means of the statistical map I had
drawn up in the preceding years, it occurred to me to
look there in search of them. The result was rather
striking: all of the emigrated colonies possessing
winter quarters of their own, had already occupied
them! This was such a regular occurrence, that, when

1) The country about Exaten consists in its uppermost layer of
light sand, which at once loses all its moisture in places exposed for
some time to the scorching rays of the sun. This condition of the soil
is surely essential in explaining the facts just mentioned.

Architecture in the Animal Kingdom. 107

I found the spring nest deserted, I had only to look
where a given colony had passed the winter; there
they would be found under the clods of their nest.
Strange, you might say, to use one’s winter quarters
as a summer resort! Yet, on considering the shelter
afforded by a winter nest, situated in the brushwood
or at the roots of a shady tree, we may easily under-
stand that the very same place can protect ants both
against the cold of winter and against the scorching
heat of summer. In this winter and midsummer nest
the sanguineas generally stay during autumn and pre-
pare for their hibernation. Whilst the hill ants (F.
rufa and pratensis) are still busy building their hills
and setting out on expeditions to visit aphides, whilst
the small, black lawn ants (Tetramorium caespitum),
the red stinging ants (Myrmica rubra) and the small,
brown garden ants (Lasius niger) are still in full
activity around their nests and are accompanying their
winged sexes on the way to their nuptial flight, there
prevails in the nests of sanguinea a quiet in striking
contrast with the feverish activity of the inhabitants
during the former months. The really active season
for F. sanguinea is from the middle of March to the
middle of August. Therefore their spring nests also
may be called working nests, whereas their winter
nests, which serve for midsummer and autumn, may
be called nests of repose.

Besides these periodical changes of residence, F. san-
guinea will also incidentally move to another nest with-
' in the temperate or hot season. This moving is caused
by special conditions of the weather. If, on account
of long aridity and heat, the sanguine ants begin

108 Chapter ITT.

to feel uncomfortable in their residence on the southern
edge of a fir-plantation, they emigrate, bag and bag-
gage, to the more shady side on the north. Toward the
end of May, 1896, I observed several colonies chang-
ing their nests for this reason. If the weather changes
and continues to be chilly and rainy, they bundle up
once more and go back to the old home. A similar
instance, I noticed on June 20, 1896. There had been
heavy showers for several days back and many san-
guinea colonies that hitherto had lived in earth-nests
were moving into old oak stumps; for these afforded
them a better shelter against the penetrating rains.
Those who designate as “intelligence,” every suitable
change in instinctive activities, caused by sense percep-
tions and sense experiences in animals, can hardly
escape from crediting ants with rather a considerable
degree of animal intelligence; for even in the highest
mammals we hardly find a higher degree of “psychic
plasticity,’ than is manifested in the above examples
by the sanguine slavemakers. However, we have
already shown in a former essay (Instinct and Intelli-
gence in the Animal Kingdom), that it is entirely
wrong to apply the term “intelligence” in this man-
ner; for the phenomena in question can be fully
accounted for by instinctive sensation and, therefore,
they do not supply the least evidence in favor of intel-
ligent, mental activity of the animal. Man, of course,
in observing such phenomena, can attribute to the ants
the following reasoning: “For the last few days it has
been raining a great deal. We and our children have
become dripping wet. Now, we do not want to get
wet again; therefore, we must move to another dwell-

Architecture in the Animal Kingdom. 109

ing, where the rain can not enter; those old oak
stumps, however, are just the thing; therefore we
move to that place.”—We, however, maintain: With-
out admitting animal intelligence the whole affair is
explained much better from the instinctive association
of sense representations. The ants do not like the
old place any more on account of the disagreeable
experiences undergone there, therefore they look for
another. That under these circumstances, just those
dry oak-stumps appear to them to be so very inviting,
follows from the suitable disposition of the sensitive
cognition and appetite. That ants in such cases are -
intellectually conscious of the suitableness of this
change of nests, is an unwarranted assumption to which
we reply: quod gratis asseritur, gratis negatur. In
other words: we are not allowed arbitrarily to attrib-
ute a human course of reasoning to animals in the
sense of “popular” psychology. Such men as L.
Buechner may find a proof of the “high intelligence”
of ants in the fact that, e. g., in low-lands Leptothoraxr
acervorum resides under barks of trees, but in the
Alps under stones.t Although we consider the power
of adaptation manifested by the sanguineas in their
nest-building instincts far more deserving of admira-
tion, yet we are far from regarding even this power as
an instance of animal intelligence, but, rather, of animal
instinct, the various activities of which depend neither

1) Buechner, “Geistesleben der Thiere,” p. 73. In this book the
author calls Leptothorax acervorum erroneously Lasius acervorum.
Romanes in his book, “Die Geistige Entwicklung im Thierreich”
(Leipzig, 1885), p. 268, was surely referring to the same passage of
Buechner, because he still more erroneously calls that ant Lasius
acerborum.

110 Chapter III.

on mechanical automatism nor on individual reflec-
tion of the animal, but on the suitable disposition of
its sensitive cognition and appetite.

True, the plasticity of the building instinct is
greater with the sanguineas than with their allied
species; yet even the latter sometimes perform actions
that go to prove clearly, that the nest-building instinct
in ants is not blind mechanism, but is suitably modified
by their sensitive cognition. I observed a striking
instance of this kind in the summer of 1898 at Lipp-
springe in Westphalia. In a growth of young fir-trees
near the so-called Fisherman’s hut there lay a small
heap of old pieces of tar-paper. This treasure had
been discovered by some Formica truncicola N yl., which
had their nest at a distance of 64 m. in a fir-plantation
on the other side of a broad, sandy road. Their nest
was a normal truncicola nest, a hill of fir-needles and
earth, built around a fir sapling. Now, the ants were
better pleased with the newly discovered place under
the tar-paper, than with their original nest, therefore
they moved over, bag and baggage; and the moving
lasted several weeks. The tar-paper afforded them in
a far higher degree the advantages usually derived
from their surface domes called ant-hills, for under
the layer of tar-paper warmed by the rays of the sun
_ there was a uniformly higher temperature and, at the
same time, an effectual shelter against the rain. There-
fore they established their nest under the tar-paper
without surmounting it by a hill. When I returned
to Lippspringe at the end of May, 1899, the truncicola
nest was still under the tar-paper, no indications of
any building being visible above ground. Their former

Architecture in the Animal Kingdom. 111

hill in the forest, however, was in ruins, a sign that it
had been completely abandoned by its builders.

Neither experience, nor instruction, nor “profound
thought” could have taught the ants, that tar-paper
possesses in a high degree the qualities which allowed
them to dispense with an ant-hill; for it was a material
which ants are not wont to meet with. It is not, there-
fore, “intelligence,” but the instinct of ants that rightly
accounts for a change of nests seemingly so wise. The
first ants that happened to find the paper were de-
lighted with the place; their senses were impressed
with its comfort and security. This led them to bring
over some of their nest-mates, and since these also
were pleased, the whole colony finally emigrated and
settled at the new place. The sensitive powers of
cognition and appetite also explain very easily, why
the ants did not raise a hill above the tar-paper. Under
the artificial roof they felt safe and warm enough
without a superstructure, therefore they saw no need
of any additional construction. Other ants, too, often
omit to build above ground, when they meet a
stone, that furnishes sufficient warmth and protection
for the underground parts of the nest. Even of F.
rufa, which is the most typical “hill-ant,” I found a
series of nests under stones, near Goebelsmuehl,
(Luxemburg) in July, 1904.

As ants, and in particular the sanguine slave-
makers, are able to adapt their skill in building nests
_ to the most varied localities and conditions of season,
so the same instinct manifests great plasticity with
regard to sudden emergencies, e. g., against the attacks
of their various enemies. A weak colony of F. san-

112 Chapter III.

guinea selects for its nest a more hidden place than a
stronger one does. This is especially the case, if there
are hostile ant species in the neighborhood, liable to
pay them a visit at any time. Thus colony 166 of my
statistical map of Exaten, being harassed by neigh-
boring pratensis, had at last hidden itself so well in
the earth, that I was obliged to spend a considerable
time before finding it. I myself have often enough
been vexed to see that colonies of the same sanguinea,
on being disturbed even by the hand of man, emigrate
and seek elsewhere a new place of settlement. Some-
times even a clod of heather placed on their nests will
cause weaker colonies to emigrate. This is all the
more strange, since our sanguinea is very fond of
using such clods as a roofing for her nests. Other
colonies, indeed, did not emigrate on account of my
repeated visits, but tried to retire farther into the
interior. A striking instance of this kind was offered
by colony No. 36, which was of moderate dimensions.
Originally the exterior of the nest looked like any
common earth-nest of the same numerical strength.
The clod which I had placed on it was used as a roof
and was covered by a heap of dry heather leaves. I
often visited the nest and each time I lifted the clod
to have a view of the interior of the building. In con-
sequence the ants blocked up the former entrances;
they dug new ones at a greater distance from the
nest, and came to the surface much more rarely. Even
the small heap of materials gathered from plants
gradually disappeared from the top of their nest.
After a time it was blown or washed away, nor did
the ants renew it. It was evidently instinct that led

Architecture in the Animal Kingdom. 113

them to withdraw from these repeated disturbances by
closing up and concealing their nest.

Professor Aug. Forel’ relates that he had brought
home from Algeria a colony of Myrmecocystus altis-
quamis and placed it in his garden at Zuerich; but
owing to the trouble caused them by the small ants
Lasius niger and Tetramorium caespitum they grad-
ually modified their usual manner of nest construction.
Under normal circumstances this Myrmecocystus
species has wide open nest entrances; in this case,
however, they were contracted to afford greater pro-
tection against the thievish visitors, and finally they
were almost entirely closed up. This instance is sim-
ilar to the one recorded above of colony 36 of F.
sanguinea, and is psychologically to be explained in
. the same way. The repeated disagreeable experiences
caused to the ants by the troublesome strangers
induced the Myrmecocysti, contrary to their former
habits, to close up and to conceal their nest. As
Forel points out, these facts afford irrefutable evidence
of the great plasticity of ant instinct. For, this
instinct is not merely a nervous mechanism forced to
operate along uniform lines; it includes sensitive cog-
nition and appetite, which are not only of an organic
but also of a psychic nature. Thus animals are
enabled, by new sense perceptions and experiences,
to adapt their wonted mode of action to the require-
ments of circumstances. This does not, however,
_ compel us in the least to attribute to animals a power
of cognition essentially the same as human intelligence ;

1) “Les Formicides de la Province d’Oran” (Lausanne, 1894), p.
8; see also “‘Apergu de Psychologie comparée,” p. 24, by the same
author.

114 Chapter III.

in fact, we cannot even do so, if we wish at all to
proceed scientifically. Popular psychology may,
indeed, perceive a “spiritual power of reflection” in
those activities of animals, in which sense experiences
enter as additional factors. And of course, this
uncritical procedure forces them, in consequence, to
ascribe to ants at least the same degree of “individual
intelligence” as to the highest mammals; for, with
the latter great plasticity of instinct is rarer than with
ants. These conclusions, which are declared absurd
by modern evolutionists themselves, clearly prove the
untenability and’ self-contradiction of modern animal
psychology.

4. How do Ants Build Their Nests?

In spite of its irregularity every ant nest is always
a unit, consisting of one or more chambers, galleries
and entrances, by which ants communicate with the
outside world. And now we are confronted with the
question: How do the members of a colony co-oper-
ate in building their nest?

Almost a hundred years ago Peter Huber? atten-
tively observed the ants building their nests, and
described in a masterly manner the skill and assiduity
of these small animals. Any one can satisfy himself
of the correctness of these observations by watching
wood ants (F. rufa) building their hills on some
sunny day in spring, or by looking at the small, black
garden ants (Lasius niger) constructing their earth-
nests during some warm spring shower.

1) “Recherches sur les moeurs des fourmis indigénes” (1810).
Nouvelle édition, 1861. Chap. I. :

Architecture in the Animal Kingdom. 115

In building nests the single workers co-operate
differently in different species; nowhere, however, do
they co-operate with the regularity of a machine or
according to a rigid pattern, but each ant with evident
liberty follows her own impulse and her own plan. It
is above all the instinct of imitation, which neverthe-
less causes a uniform result, a nest consisting of com-
municating galleries and chambers. As a rule the
most zealous and skillful worker is imitated most;
her zeal is catching, so that she directs the activity of
the others into the same channel. This mode of
co-operation effected by means of the instinct of imi-
tation is predominant in hill ants (F. rufa) and the
small, blackish garden ants (Lastus niger). F. fusca,
however, the greyish-black slave ant or negro ant,
(Huber’s fourmi noir-cendrée) belongs to those
species, in which the independence of the several
workers in building is especially remarkable. The
same is observed with the closely allied F. rufibarbis
(Huber’s fourmi mineuse). I.often witnessed in
these two species, how the pellets of earth, which one
worker had just put down for building a wall in a
certain place, were carried away by another one in
order to apply them elsewhere in a manner more suit-
able to her own taste. To observers, who know the
habits and doings of ants but superficially, and are
wont to transfer their own thoughts into the brain of
the animal, such occurrences may appear as though

an ant wished “purposely to correct” the work of
~ others. And, as a matter of fact, one of P. Huber’s
observations! bearing on this head has been actually

1) Lic., p. 48.

116 Chapter III.

thus interpreted by writers on popular science, and
made to serve as a beautiful proof of animal intelli-
gence. This kind of arbitrary misrepresentation of
the plainest facts can certainly lay no claim to any
scientific value.

How, then, do ants build their nests? They adapt
themselves to given situations and prudently take
into account the various circumstances. When in
spring a gentle, warm rain begins to render the dry
soil soft and manageable, immediately the earth-
working ant species are kindled with new zeal for
building. These assiduous little animals will then
sally forth from their nests by hundreds and place
pellets of earth upon pellets to build new galleries and
chambers, availing themselves of blades of grass,
twigs of heather, pieces of leaves and other natural
props as pillars or vaults. For the same purpose such
auxiliary materials are also first dragged to the spot.
In this connection the sanguine slavemakers give evi-
dence of eminent skill in combining timber-work with
masonry.

It is especially remarkable with earth-working ants,
that they accommodate their instincts to the changes
of temperature and moisture. This could be ascer-
tained constantly in glass nests, in which I kept under
observation small colonies of Lasius niger, Tetramo-
rium caespitum and Myrmica scabrinodis. If the mois
ture of the nest became too great, the ants would set
to work and pile up the earth in the shape of a dome
perforated by innumerable openings, making it appear
like a sponge; thus the evaporation of the water was
facilitated. But when the moisture decreased too

Architecture in the Animal Kingdom. 117

much and the nest threatened to dry up, a change to
the opposite was effected; the nest was constructed as
flat and as low as possible with very few openings on
the surface. The propriety of such proceedings is cer-
tainly striking. Often enough the same can be
observed also in nature, both in those species that
build only domes of earth, and in those that build
regular ant-hills. It is a fact even noticed by farmers
and ascertained by myself repeatedly, that in dry and
hot summers the hills of wood ants are lower and
flatter than in moist and cold summers. The first way
of building is for the purpose of reducing evaporation
to the lowest limit and to offer to the hot rays of the
sun but a small surface to shine upon; on the other
hand, the higher and the more vaulted the hills are,
the easier is the drainage in case of rains, and the
greater are the evaporating. and heating surfaces.
Indeed, ants would have to be very intelligent, if their
own reflection should lead them to modify their nests
so prudently. Yet, considering that the dome shape
for the nest includes in principle these quasi-intelli-
gent adjustments to the variations of temperature and
moisture, which can be traced, in consequence, to the
specific manner of building proper to the ants in
question, it is clear that instinct and not intelligence
is the guiding principle; for, the specific plan of the
building is, no doubt, instinctive, as is acknowledged
at least by scientific authors. In adjusting, however,
their instincts to changing circumstances, the animals
are influenced by sensitive perceptions and experi-
ences, which are nothing else than the natural exercise
of the same instinctive power of cognition, on which is

118 Chapter III.

based the specific mode of building in any species of
ants. Why, therefore, introduce a foreign element,
called intelligence, between this hereditary disposition
and its changeable application? I should think it is
far more simple and natural to account for the whole
activity of an animal by one and the same principle.
Unless you mistake instinct for mechanical automa-
tism, it is by no means necessary to assume animal
intelligence in order to explain the above phenomena.

Let us now compare the architecture of ants with
that of birds. There are several important differences.
The nests of birds are more artistic! and regular;
yet they are stamped with the unmistakable marks of
monotony and uniformity within the same species,
they are products of instinct in the strictest sense of
the term. Moreover, as Altum? has admirably proved,
the architecture of birds is a function of their
breeding instinct. It begins at a certain stage
of the development of this instinct; both reach
their climax at the same time, and then grad-
ually vanish together. On this account the nests
for the first hatching in spring are, as a rule, built
better than those later in the season. In this case
practice does not form the master but the bungler.*
With ants, however, architectural skill is found in the
-workers throughout their life, it is suitably carried into

1) By the way, this art has often been exaggerated. See ‘Die
Baukunst der Voegel auf ihren wahren Wert zurueckgefuehrt,” in
“Jahrbuch der Naturwissensch.”’ I (1885-1886), 198.

2) “Der Vogel und sein Leben” (6th ed.), p. 163 ff.

8) Something similar obtains also among the Coleoptera in the
ingenious nest-building of the leaf-rolling beetle (Rhynchites betulae).
See Wasmann, “Der Trichterwickler, p. 78 ff.

Architecture in the Animal Kingdom. 119

operation under the most different circumstances, and
is even to a certain degree improved by sensitive
experience, i. e., it is modified according to new per-
ceptions. True, many birds, too, are able to adapt to
changing circumstances’ the places and the materials
they choose for their nests; nor can their instinct be
called mechanical automatism, because its activities
are governed by the sensitive cognition of the animal.
Many species of birds change the place and the
material of their nests according to locality; besides,
the single individuals are not rigidly bound to a cer-
tain material for building their nests, but they fre-
quently employ scraps of paper, horse hair, cotton and
other materials, which happen to come in their way.
Those birds that build nests more or less open to view,
instinctively avoid materials the coloring of which
would set their nest into striking contrast with the
surroundings. In this they are evidently governed by
their power of sense-perception. Sometimes this pro-
tective resemblance of the nest with its background is
effected quite of itself, when birds use the material
for their nests, which is usual and natural to them;
but sometimes also unusual materials found by chance
serve the same purpose. Thus I was informed by a
friend:? “In Blyenbeck (in the northern part of Dutch
Limburg) I had occasion to observe how chaffinches in
building their nest ‘ingeniously’ imitated the greyish-
white color of lichen which covered the tree on which
they built. They used small scraps of paper, and thus

1) Several examples of the same are found in ‘“‘Westfalens Thier-
leben,” Vol. II. Besides also in Darwin’s posthumous essay on instinct
(Romanes, “Evolution in the Animal Kingdom,” appendix).

2) Rev. L. Dressel, S. J.

120 Chapter III.

they disguised their nest.’”* The visual resemblance
between the white tree-lichens and the paper-scraps,
which impressed the sensitive power of perception of
those chaffinches explains quite naturally their seem-
ingly intelligent proceeding.

He, who concurs with Darwin? in attributing intel-
ligence to birds when their actions are influenced by
sensitive cognition, must credit ants with a still higher
degree of intelligence; for it cannot be denied that
birds, in building their nests, show far more specific
uniformity than individual variability, whereas in ants,
as a rule, the contrary is the case; with them the
“psychic plasticity’ of the nest-building instinct is no
doubt much greater.

Yet the chief point of excellence, which distin-
guishes the architecture of ants from that of birds is
the number of uses to which it may be put. Birds
build nests to serve as places for hatching their young
only ; except during the pairing season, birds do not
know their nests, nor does it ever occur to them to
use them as dwelling-places. With ants, however,
nests serve as permanent abodes for the whole family
and often also for strangers of different species, which
are hospitably received as guests. Finally, ants use
their architectural skill for many other purposes. But
before entering into particulars on this head, we would
like to draw a comparison between the architecture of
mammals and that of ants.

In as far as the buildings of many mammals are

1) Ch. Darwin, too, mentions already a nest of a chaffinch described
by Hewitson, in which, instead of lichens, shreds of paper likewise

had been employed (l. c., p. 417).
2) L.c., p. 414.

Architecture in the Animal Kingdom. 121

not only nests but also permanent dwellings both for
the parents and the young, they bear a closer resemb-
lance to the nests of ants than to those of birds.
Instances are plentiful; for the burrows of badgers,
foxes and wild rabbits are well known. In some cases
the same buildings serve also for storing provisions,
as is the case with German marmots and moles. These
buildings, therefore, by their greater universality of
purpose, are more similar to ant-nests than to bird-
nests. Another point of similarity with the former
is in this, that they show, with some species at least,
a greater individual variability and less specific uni-
formity than the latter. Nevertheless, in all these
respects ant-nests by far excel the buildings of mam-
mals. The very highest vertebrates, the anthropoid
apes, scarcely manifest a trace of building instinct or
of -its intelligent application, unless you wish to
mention the sleeping-places somewhat resembling
regular nests, which Orang-Utans* are wont to build
on trees. Although the brain of apes most resembles
that of man, yet the most “intelligent” architects
among mammals are found, not among the apes but
among the rodents, which in development of brain
are far inferior. Beavers are the.only higher animals
whose architecture can bear comparison with that of
ants.

The buildings of beaver-families consist of an
underground chamber and burrow resembling those
of other mammals, and of a so-called “lodge.” The
manner in which the latter is built was ably described

1) See Buettikofer, ““Zoologische Skizzen aus der Niederlaendischen

‘Expedition nach Central-Borneo” (Compte rendu du 3me Congrés
international de Zool.), p. 224.

122 Chapter III.

by Friedrich in a recent publication.t The beaver-
lodge is nothing but an accumulation of brushwood
above the opening of the underground chamber, which
is the real center of the whole dwelling. Wherever
beavers are living in colonies and when circumstances
favor the full development of their instinctive skill in
building, they construct their well-known dikes? to
dam the water, and sometimes they even build canals
for the transportation of timber. Although these
works are the result of the co-operation of several
families, yet each pair works only for its own pur-
poses; there is never any division of labor like that
in ant colonies. Of course, observers like Lewis H.
Morgan,* who mistake for intelligence every action
due to sensitive cognition, discover many proofs of
high intelligence in the doings of the American
beavers. However, this so-called “free intelligence” is”
nothing else than the power of adapting their buildings”
to the changes of situation. This power is possess
also by ants in at least an equal degree. If Mo
and Romanes,‘ e. g., regard it as an infallible proof”
of the intelligence of beavers that they regulate the

level of their ponds by widening or narrowing “the

orifices of their dams as the case may be,” they should

admit the same in ants, which regulate the degrees of

moisture and temperature of their nests by changing

1) “Die Biber an der mittlern Elbe” (Dessau, 1894), p. 20 ff.

2) According to Friedrich they also occur at some places on the
banks of the middle Elbe, although indeed on a smaller scale; there-
fore they are due to an instinct common both to the European and
the American beaver.

8) “The American Beaver and his Works” (Lippincott & Co.,
1868).

4) “Animal Intelligence” (6th ed), p. 377 ff.

Architecture in the Animal Kingdom. 123

their domes accordingly; and, just as beavers use
their architectural skill not only in building nests but
also in constructing dams and canals, so ants use theirs
for far more various purposes. Those who concur
with Romanes in maintaining that “the adaptations
of pure instinct have reference only to conditions that
are unchanging’, and that meeting of continual vari-
ation of conditions cannot be accounted for but by
intelligence, must, indeed, attribute to beavers a con-
siderable degree of individual intelligence, but no less
to ants. However, this conception of instinct and
intelligence is untenable. Even Romanes is loath to
ascribe to beavers such high psychic faculties, yet
with his false notion of intelligence this conclusion is
unavoidable. If beavers in modifying their buildings
are guided by their own reasoning, they must be like-
ise credited with an intelligent knowledge of the
inciples of their architecture; for the former without
atter is impossible; thus, we have, instead of
building instinct, the highest order of human,
rchitectural intelligence! This is evidently untenable.
Those, however, who explain the architectural instinct
of beavers from the hereditary disposition of their
sensitive cognition and appetite, are able to explain
from the same principle any given modification of their
architecture, without resorting to “animal intelli-
gence.”

5. Other Purposes for Which Ants Employ Their
Architecture.

Some, ants having populous colonies often establish -
temporary stations at the foot of trees and bushes,

it ee Ae i sy

124 Chapter ITI.

where they visit their plantlice and scale-insects to
“milk” them by caressing them with their feelers. A
few European ants, namely Formica rufa, pratensis
and Lasius fuliginosus build regular streets, clearing
away from their path all vegetable growth to a dis-
tance of sometimes from 20 to 50 m.t_ These streets
lead from their nests into woods and bushes and thence
branch off to the pasture-grounds of their “cattle.”
Other ants, in particular Lasius niger and Cremas-
togaster scutellaris build covered roads or tunnels of
earth, by which their nests communicate with trees
and bushes that are inhabited by aphides or scale-
insects. These they occasionally surround with earth-
ramparts, in order to keep them together and to pro-
tect them from other ants by whom they might be
coveted. Such a “plantlice-pavilion,’ an earth con-
struction the size of a hazel-nut, is in my collection;
it was built by Myrmica scabrinodis at the top of an
oak-twig in the neighborhood of Exaten. Other ants,
e. g., the African Dorylus species, dig subterranean
tunnels, where they go for their prey, consisting
chiefly of insects and worms. The harvesting ants of
Southern Europe, Western Asia, Northern Africa,
America and India establish granaries in their nests,
where they store up their provisions for winter or sum-
mer. The Afta of tropical America, feeding on mush-
rooms, use a number of subterranean chambers as vege-
table gardens and hot-houses, in which the mushrooms

1) Among foreign ants there ate especially the larger species of
the American leaf-cutting ants (Atta) which, according to Belt, Brent
and Forel, build similar roads, but frequently of a still more consider-
able length and breadth. The same ant-roads we find in North America
with ants of the group of F. rufa, especially with Formica exsectoides.

Architecture in the Animal Kingdom. 125

are grown.’ Ants employ their building skill also to
protect themselves against enemies. They raise ram-
parts and barricades to keep off foreign invaders; and
unwelcome visitors which cannot be got rid of in any
other way, are simply walled up with earth, and are
thus kept at a distance. Thus, in one of my observa-
tion nests of F. sanguinea a salamander introduced
by me was in a short time entirely walled in. The
slaves (F. fusca), past masters in the art of building,
were most zealous in this work. An occurrence far
more amusing took place in a nest of Lasius flavus,
to whom I had given a Lomechusa strumosa as guest.
The small, yellow ants were not at all pleased with
the unwieldy fellow, and tried to get rid of his impor-
tunity in the following droll manner. From all quar-
ters they brought together pellets of earth and heaped
them up on the back of the unfortunate beetle, until
nothing was to be seen of him but the tips of his
feelers.”

Ants are even supposed to have their cemeteries
and burial-places. Superficial observers have circu-
lated many fables on this subject. In the book of a
certain Reverend White (Ants and their Ways, Lon-
don, 1883), I found a touching story by Mrs. Lewis-
Hutton, of Sidney, which is really too characteristic of
this kind of natural history to be passed over in silence.
One of her children had sat down on an ant-nest and
had been assailed by the enraged inhabitants. At the

1) Moeller, “Die Pilzgaerten einiger suedamerikanischer Ameisen’”’
(Jena, 1898), and Forel, “Zur Fauna und Lebensweise der Ameisen
im Columbischen Urwald” (Mitteil. der Schweiz. Entom. Gesellsch.,”’
IX, 9th issue), p. 406.

2) “Die internationalen Beziehungen von Lomechusa strumosa,”’
in “Biol, Centralbl.,” 1892, p. 653.

126 Chapter III.

cries of her child the mother ran to the spot and killed
a few score of ants. A short time after she saw the
corpses surrounded by a number of their companions.
The burial ceremonies began. A deputation of ants
was despatched to the nest to fetch the train of
mourners. They marched in due order two by two to
the scene of disaster. They took up the corpses,
marched slowly in procession to a sandy place in the
neighborhood and buried them one by one. A few of
the gravediggers which tried to escape this doleful -
duty by flight, were pursued by the other ants, over-
taken and summarily sentenced to death. The sen-
tence was immediately carried out, and the criminals
were all interred in a common pit. The said lady
maintains to have witnessed similar proceedings more
than once. Gerstaecker in his “Report on the scientific
results obtained by Entomology during the year 1861”
mentions this burial story (p. 156) with the follow-
ing ironical remark: “To render the mystification
complett, nothing more was lacking than a funeral
sermon held by one of the ants.” Strange to say,
Perty? attempts to defend the imaginative lady against
Gerstaecker by saying: “There seems to be some
truth in it, anyway, for Dupont also maintains that
ants have common graveyards at some distance from
their buildings, whither they carry their dead.”
Ernest André? was far more correct about those burial
ceremonies of ants in calling them phantastic misrep-
resentations of the commonest occurrences. It seems
scarcely possible, that such an anecdote should see

1) “Seelenleben der Thiere” (2d ed.), p. 328,
2) “Les fourmis” (Paris, 1885), p. 176.

Architecture in the Animal Kingdom. 127

the light in a highly scientific publication, in the Trans-
actions of the Linnean Society of London (1861).
We are scarcely able to understand, why George
Romanes, who incorporated the Australian burial
anecdote in his work “Animal Intelligence,” did not
even in the sixth edition (1895) entertain any doubts
about it, but continued to regard it as_ sufficiently
authentic. Something, to be sure, was true in the
whole affair; but this “something” is confined to the
fact that ants remove their dead from their nests to a
certain place where they collect all their rubbish, and
sometimes cover it with earth. These places, how-
ever, would no doubt be far more correctly called
“dumping grounds” than graveyards; for they are
nothing but places where they deposit everything
that displeases their instinct of cleanliness.1 These
phenomena have nothing to do with “Chinese venera-
tion for the dead,’* as can be easily ascertained in
artificial observation nests. In these nests the dryest
spots farthest away from the interior of the main nest
are used for collecting the refuse. In my large obser-
vation nest of F. sanguinea the place bearing the namg
“refuse nest” is the one in which the corpses of dead
ants, remnants of dead flies, wings of dismembered
dragon-flies, empty cocoons of ants and other rubbish
is finally stored, just because the ants wish to get rid

1) Forel, “Ameise und Mensch oder Automatismus und Vernunft.”

2) W. Marshall, in his ‘“‘Leben und Treiben der Ameisen” (p. 26),
says, ‘the American forms seemed (as regards the treatment of their
dead) to be more affectionate than those of the old world.” We should like
to know why? At least nothing of the kind follows from the observa-
tions, which he mentions, of McCook. See “Die Honigameise des
Goettergartens” (“Stimmen aus Maria-Laach,” XXVII, 1884, 282),

where we have dwelt more at length on the “affection” of the Ameri-
can honey-ants.

128 Chapter III..

of such disagreeable objects. It is utterly useless to
embellish the life of ants with fabulous anecdotes like
that Australian burial story. The cold facts are inter-
esting and wonderful enough.

6. Is the Architecture of Ants Guided by Intelligence?

The building instinct of ants proves to be such a
universal faculty, and its application to various pur-
poses is in many cases seemingly so intelligent, that
we are finally confronted by the question: Why
should we not call this an intellectual faculty? The
following discussion will probably throw some light
on the subject.

Would it not be a proof of intelligence, if ants,
not themselves provided with spinning glands,
employed their larvae for manufacturing threads, by
means of which they build a nest of leaves? Accord-
ing to W. D. Holland’s observations this is done by a
large, reddish-yellow ant of Eastern Asia called
Oecophylla smaragdina, whose nests he studied in
Ceylon.t. With their mandibles the ants first bring into
the proper position the leaves to be connected and keep
them there; then others approach in large numbers,
each carrying a larva in its mouth, with which they
begin to move across the leaves from side to side.
Wherever the mouth of the larva touches the leaf, a
thread appears sticking to the leaf. This process is
continued, until the leaves are attached to one another
at their edges by a firm tissue of threads, and finally
a viscous, paper-like stuff is formed consisting of
innumerable threads crossing each other in all direc-

1) E. E. Green, “On the habits of Oecophylla smaragdina F.”’
(“Proceedings of the Entomological Society of London,”’ 1896, p. IX.).

Architecture in the Animal Kingdom. 129

tions. These ants use their larvae as “spinning
wheels,” not only for building their paper-nests, but
also, according to Holland, for protecting their nests
against the invasions of small ants, with whom they
are in continual warfare. Around the trunk of the
tree containing their nest they sometimes put a belt
a foot broad, formed of threads, which serves to
entangle the small ants and prevent them from climb-
ing the tree. In manufacturing this protective tissue
the ants come forth from their nest, each carrying a
white pellet in its mouth, and move to and fro on the
trunk. Upon closer inspection it was found that those
little white lumps were again their larvae!

These curious phenomena needed to be corrobo-
rated by further investigation. Forel in his “Die
Nester der Ameisen” (p. 20) already called attention
to the fact that the mandibular glands of Oecophylla
-- are strongly developed and possess large and numer-
ous cells. From the analogy with other ant species
(Cremastogaster, Dolichoderus, etc.), which secrete
a certain glue from their mandibular glands for manu-
facturing their paper-nests, it might seem probable,
that the spinning material of Oecophylla proceeds
from the ants’ mouth. But, on the other hand, Pro-
fessor Chun has recently shown in his splendid book
“From the Depths of the Ocean,’’! that the spinning
glands of the Oecophylla-larvae are far more developed
than those found in other larvae of ants. We must.
conclude, therefore, that Mr. Holland’s statements
were quite exact, and that the spinning glands of the
larvae, not the salivary glands of the ants themselves,

1) “Aus den Tiefen des Weltmeeres,”’ 2d ed., Jena, 1903, p. 129.
9

180 Chapter III.

deliver the threads employed in the marvelous archi-
tecture of Oecophylla.

In adopting this supposition, that ants employ
their own children as a kind of “spinning wheel,’ we
are confronted by the extraordinary fact that animals
make use of an instrument, other than any bodily
organ, for building and defending their nests, an
occurrence unparalleled in the whole animal kingdom,
even among higher animals. But can we account
for the proceedings of this Indian ant on the score
of intelligence, that is to say, of her own, individual
reflection? Just as little as in the case of the other
specific arts and talents of animals; for they are all
the hereditary property of certain species, not invented
or learnt by independent individuals. And, therefore,
also the spinning talent of Oecophylla, even if it is
done by means of the larvae, is due to hereditary
instinct, not to the individual intelligence of the animal.

To obtain proofs for attributing the architecture
of ants to their own intelligence we sliould have to
look for instances, in which, in consequence of indi-
vidual experience and reflection, these animals modify
their innate instincts in such a manner as to invent
new means of accomplishing their purpose. This is
the third form of independent learning, which, as we
have shown in a former publication, is a real proof
of the intelligence of the learner. We must, there-
fore, examine, whether the building activity of ants

1) In the chapter on the different forms of learning in ‘Instinct
and Intelligence” (Herder, St. Louis, 1903). A more detailed discussion
of this point will also be found in our publication, ‘Die psychischen
Faehigkeiten der Ameisen” (Zoologica,” 26th issue, Stuttgart, 1899),
pp. 82-114.

Architecture in the Animal Kingdom. 131

contains any facts belonging to this class of psycho-
logical actions.-

Patrons of animal intelligence have repeatedly
maintained, that ants build bridges with the intelligent
purpose of overcoming obstacles placed in their way.
Even a century and a half’ ago it was noticed by
Cardinal Fleury, that when he tried to keep ants from
climbing a tree by smearing it with bird-lime, they
gradually covered the bird-lime with earth and thus.
paved a road across. He informed the famous Réau-
mur of this observation, who immortalized it in his
Mémoires pour servir a l’histoire naturelle des Insectes
(1734-1742). And because ants have not become
weaker-minded since the days of Réaumur, many
other friends of nature had the occasion, from that
time on, to observe and.to record similar “bridges”
built by these insects. One of these reports, best
known in recent times, is the following.t Professor
Leuckart in Giessen (Germany) wishing to keep ants
from frequenting a certain tree, surrounded it with a
cloth soaked in tobacco-juice. The ants above the
cloth upon meeting the impediment turned back and
after awhile let themselves drop to the ground from
the branches, but those which were ascending to visit
the aphides, after-having in vain tried to cross the ill-
smelling cloth, descended and after a short time were
seen coming back, each with a pellet of earth in its
mouth. These they pasted over the tobacco-juice, and
soon a passable road was constructed. William Mar-
shall recording this observation of_Leuckart in his
“Leben und Treiben der Ameisen” (p. 40) adds the

1) Buechner, “Geistesleben der Thiere,” p. 116,

182 Chapter III.

following momentous reflections: “All the philos-
ophers together of ancient and modern times, and all
the theologians moreover, will not impose upon me
by asserting that we have to do here with the action
of an unreasonable creature. If this is instinct, then
the invention of the steam-engine is instinct, too! No,
both mean a clever profiting by given circumstances,
due to reflection !”

Many an unwary reader may, possibly, be over-
whelmed by this spirited appeal to the steam-engine
on the part of Marshall. However, if we do not allow
ourselves to be imposed upon by the boldness of his
oratorical flight, we shall arrive at different results,
without being exactly philosophers or theologians.
We can observe any day, that on the part of the ants
ill-smelling or sticky objects are simply covered with
earth, if they cannot be removed from the nest.
Nobody will be inclined to maintain seriously, that
ants, by so doing, make use of any “intelligent gift of
invention” transcending their power of instinct. Out-
side of their nests also, and governed by the same
instinct, they occasionally adopt the same procedure.
Now, in the above mentioned case the ants found, that
the road which led them to their aphides on the tree,
had been covered with an ill-smelling, sticky substance.
What was more natural than to fetch pellets of earth
and to clear a passage by a method so familiar to their
instinct? Hence, we are justified in drawing the fol-
lowing conclusion: That by this pretended “bridge-
building” these ants have given infallible evidence of
reflection, inventive genius, and intelligence, is a state-
ment worthy only of popular, uncritical psychology.

Architecture in the Animal Kingdom. 133

One thing, however, is made evident by these
and similar observations, viz.: that ants are not mere
reflex machines, but beings endowed with sensitive
cognition and appetite, and with the power of employ-
ing in the most various manner their innate, instinc-
tive faculties and abilities under the influence of
different sense-perceptions. And just on this account
it is altogether superfluous to admit “animal intelli-
gence”; for, the complex representations of sensitive
cognition, as we have shown in the above example,
afford a simpler and better explanation of whatever
is not mere fiction in those supposedly intelligent
actions of animals.?

Another example of bridge-building, which, by the
way, is merely vouched for by a Mr. Theuerkauf in
Buechner’s “Geistesleben der Thiere” (p. 117), is
still less corroborative of ant intelligence than the
former. In this case the ants used a different means
for bridging over a circle of tar smeared around a
tree. The ants were descending from the top; on —
arriving at the obstacle some stuck fast, others
returned to fetch plantlice from the twigs; they put
them oh the tar and thus constructed a bridge. Sir
John Lubbock? remarks in explanation of this story,
that he had his doubts as to the interpretation of the
fact. “Is it not possible that, as the ants descended
the tree, carrying the aphides, the latter naturally
stuck to the tar, and were therefore left there? In
the same way I have seen hundreds of bits of earth

1) Cf. on this point, “Instinct and Intelligence in the Animal King-
dom” (1903), p. 109 and 137 ff., where we have shown, that not even
higher animals may be credited with formal consciousness of purpose.

2) “Ants, bees and wasps’? (London).

134 Chapter III.

deposited on the honey, with which I fed my ants.”
In fact, only trivial observers could maintain that in
this case the ants had intentionally employed their
aphides as bridge-building materials. The correct
explanation might rather be the following: the ants,
becoming uneasy about their precious aphides living
on the tree, tried to save them by carrying them down.
By this attempt, however, the aphides literally “got
stuck in the mud.” It was, no doubt, merely by
chance, that the aphides adhering to the tar formed
a sort of bridge for the ants.

These two famous stories are, therefore, far from
furnishing any evidence in favor of ant-intelligence.
We have to investigate other examples to determine
whether or not ants are able by “reasonable reflection”
to invent new means for fulfilling their designs.

Sir John Lubbock? has made a number of experi-
ments with ants in order to test their intelligence ; some
of the more important only can be mentioned here.
For the ants of a nest of Lasius niger he arranged a
bridge made of a piece of straw or a slip of paper, by
which they could get at their larvae. After the ants
had become sufficiently familiar with this pathway,
he slightly moved the bridge, ‘“‘so as to leave a chasm,
just so wide that the ants could not reach across. They
came and tried hard to do so; but it did not occur to
them to push the paper bridge, though the distance
was only about one-third inch, and they might easily
have done so.” Another experiment he relates as
follows: “I suspended some honey over a nest of
L. flavus at a height of about half an inch, and accessi-

1) “Ants, bees and wasps,” Chap. IX.

Architecture in the Animal Kingdom. 136

ble only by a paper bridge more than ten feet long.
Under the glass I then placed a small heap of earth.
The ants soon swarmed over the earth on to the glass
and began feeding on the honey. I then removed a
little of the earth, so that there was an interval of
about one-third of an inch between the glass and the
earth; but, though the distance was so small, they
would not jump down, but preferred to go round by
the long bridge. They tried in vain to stretch up from
the earth to the glass, which, however, was just out
of their reach, though they could touch it with their
antennae; but it did not occur to them to heap the
earth up a little, though, if they had moved only half
a dozen particles of earth, they would have secured
for themselves direct access to the food.” It is evident
from this, that the ants had not the least idea of em-
ploying even this simple means. In all his experi-
ments Lubbock obtained entirely negative results.
There was no indication of the much-vaunted intelli-
gence of ants. Strange to say, William Marshall has
not mentioned these experiments of Lubbock in his
“Leben und Treiben der Ameisen,” although they
could not have been unknown to him, as he translated
them himself into German; perhaps it was, because
the results did not fit in with his enthusiastic praise
of the “reflective faculty” and “inventive genius” of
these animals.

This latter experiment of Lubbock was repeated
of late by Albrecht Bethet in a somewhat different
form. Over a well-frequented pathway of Luasius

1) “Duerfen wit den Ameisen und Bienen psychische Qualitaeten
zuschreiben?” (Bonn, 1898), p. 66.

136 Chapter III.

niger he fastened a strip of tin with some honey.
After the ants had for a long time been allowed to
visit the honey, the strip was gradually raised by a
screw, until from their pathway the ants could no
longer get at the honey. Though it would have been
easy enough to heap up a little earth under the strip of
tin, it never occurred to the ants to do so; the honey
remained beyond their reach. This experiment, there-
fore, had the very same results as Lubbock’s, namely,
that the ants were not capable of forming the simplest
intelligent conclusion, which would have led them to
employ their building skill for the purpose of getting
at the honey.

I may add here a few observations and experiments
of my own. Since it might be objected against Lub-
bock’s results, that he took for his experiments some
ant-species “little endowed with intelligence,” namely,
Lasius and Myrmica, I chose the most intelligent ants,
namely, Formica sanguinea* and her allied slaves as
subjects for experiments, of which only a brief extract
is here presented.

In the front-nest of my above mentioned observa-
tion-nest (see p. 23) a piece of wood formed a com-
modious bridge, over which the ants could pass to the
rim of the glass and thence into the top-nest. By their
earth-constructions in the front-nest the ants had
gradually lowered the bridge, so that the distance

1) Bethe infers from this experiment that ants do not even possess
sensitive perception and cognition. This inference is too far-reaching,
and is owing to his mistaking intelligence for sensitive cognition. See
“Die psychischen Faehigkeiten der Ameisen,” p. 73.

2) Forel also (‘‘Fourmis de la Suisse,” p. 443) states that F.
sanguinea deserves the palm for intelligence,
’

Architecture in the Animal Kingdom. 187

between the top-end of the bridge and the cork of the
glass was about 2 cm. When exposed to the sun the inside
of the glass was generally covered with moisture, and
the ants found great difficulty in passing the intermedi-
ate space and in getting into the top-nest. Although
this state of affairs lasted for weeks, and the ants con-
tinued to encounter the same difficulties, it never
occurred to them to connect the broad, upper end of
the stick with the roof of the front-nest by a bridge
of earth. They connected the sides of the stick with the
glass by a wall of earth; gradually they also covered
the whole glass-wall with pellets of earth to protect
themselves against the rays of the light; but they
never built a bridge at the critical spot, where one was
evidently needed and of the greatest importance. The
pellets of earth accidentally fastened there were, on
the contrary, continually thrown down by the ants
that crawled up the road. Thus exactly that spot
over which the ants, if they were endowed with any
power of thought and reflection, would be expected to
build a bridge, was left slippery and smooth, and was
still the same after half a year.

In 1884 I repeatedly made the following experi-
ment: In a tiny vessel I suspended some honey or
ant-larvae over a nest of F. sanguinea, contained in a
large “crystallisator”,’ so that the ants could touch the
vessel with their antennae only, but could not reach
it except by a very circuitous route. It should have
occurred to them to heap up a little earth or some
pieces of wood underneath the vessel, in order to form
a “bridge” leading directly to the wished for goal.

1) A low, round glass bowl, covered by a glass plate.

138 Chapter 111,

But neither the sanguineas nor their slaves (F. rufa
and fusca), living in the same nest, ever hit upon this
obvious method, although it would have sufficed to
raise the surface of their nest at the spot in question
just by 1 cm.!

A more wonderful result was obtained in another
experiment on the same nest of sanguinea. On June
16, 1884, I filled a large watch-crystal with water and
in the center upon a kind of island I placed a little
shell filled with ant-cocoons previously taken from the
same colony. This artificial pond with its island was
then introduced into the nest. The ants soon noticed
the cocoons and stretched out their feelers towards
the island; but getting into the water at every attempt
to approach, they retreated again and again. I
began to think they would never be able to overcome
the difficulty, when suddenly a sanguinea began to
throw into the water pellets of earth, bits of wood,
dead ants and similar solid materials. Others followed
her example and they soon had built a road over the
water! In the space of an*hour, counting from the
minute I started the experiment, they had fetched all
the cocoons from the island by means of this “floating
bridge.” The very last cocoon having been secured
by the ants, one of them returned to the island and,
finding it empty, she squatted on her haunches, passed
the spur of her fore-feet through her mouth and then
combed her feelers with the spur, sitting there for
several minutes in a most provoking attitude, as if she
were saying to me: “Ah, my dear, who has won the
game now?” Js this fact not a staggering proof,

1) This very last instance, as many others in this translation, was
added by the author from his original notes.

Architecture in the Animal Kingdom. 139

that at least the sanguineas are endowed with a good
quantity of reflective power and of intelligent con-
sciousness of purpose?

In order to answer this question I tested the above
observation by the following experiment: After some
time I again placed the same watch-crystal filled with
water in the nest, but this time without island and
without cocoons. Now, supposing that the ants in
the previous case had really intended to build a bridge
for the sake of getting at the cocoons, there was no
longer any reason for them to repeat the procedure.
However, this second time also, after several had
accidently got their feet wet, they soon started to fill
up the lake. Although this time there were no cocoons
to be obtained, nor any island in sight,’ yet they again
covered the water with all kinds of materials in the
same manner and almost in the same space of time as
they had done before. Hence, we are allowed to con-
clude, that even the first time the ants had not intended
to build a floating bridge, but only to get rid of the
disagreeable moisture that barred their way. If, there-
fore, we maintained, that in the first experiment the
ants had by intelligent fore-thought invented a means
for regaining possession of their cocoons, we would
be guilty of uncritically humanizing the brute.

From all the observations made and noted down for

1) I wish to lay emphasis on this circumstance, because Prof.
Charles Sajé (in “Prometheus,” 1899, No. 486, p. 284) believes the
ants had hoped to find some treasure on the island this time also. The
ants could easily notice from the margin of the watch-crystal, that
there was no island. The eyes of F. sanguinea being rather large and
sharp and capable of distinguishing the form of small objects at a
distance of several centimeters, the ants could undoubtedly see that
there was no island,

140 Chapter III.

the last twenty years, I could record here still many
an iftteresting occurrence, which, like the above exam-
ple, impresses a superficial observer as an intelligent
action. Yet, closer examination invariably proves
that such facts are accounted for much more easily
and naturally by the instinctive combinations of sense-
representations ; therefore, no “ant-intelligence,’ and
in fact no “animal-intelligence’ at all is required.
Indeed, the higher mammals ranking next to man
in brain development are far from supplying more con-
vincing proofs of “intelligence” than ants. In them
also the whole process of cognition is confined to the
mere connecting of sense representations and sense ex-
periences according to the inborn laws of instinctive
association of representations, which ordinarily regu-
late their lives. The psychic endowments of dogs
and monkeys go no farther. Unless a dog has been
specially trained, it never occurs to him to open a
door, the knob of which he is unable to reach, by
fetching for instance a foot-stool to gain a higher
level; he may have seen children, his play-fellows,
doing the same thing a hundred times; the relation
between means and end, though so natural and obvious
in this case, will forever remain hidden to the canine
soul. Hence the dog is not a whit more intelligent
than the ants, that failed to notice, that a little heap
of earth would have sufficed to secure them an easy
passage to the honey suspended in a saucer above their
nest.
Neither do apes possess the power to invent by
their own reflection new means of accomplishing their
end. Even these highest mammals are confined ex-

Architecture in the Animal Kingdom. 141

clusively to the instructive association of sense-repre-
sentations. We have shown this in our first chapter,
where we discussed the wars that take place in the
animal kingdom, and pointed to the fact that apes are
unable to invent the simplest weapons and implements
even. The same holds good as to the use of fire. If -
a troop of apes in the forest hits upon the remnants
of a fire lighted by the hands of man, they will cer-
tainly gather around it and enjoy the comfortable
warmth. But it has never, hitherto, occurred to any
ape to supply it with fuel.t And yet it would be such
a simple and natural combination of representations,
requiring but a low degree of intelligence. Why do
apes, in spite of the “high plasticity” of their quasi-
human brain, never hit upon such a simple means?
Because they possess no spiritual soul and therefore no
intelligence. ‘The “plastic neurozymic activities” of
the simian brain are essentially different from human
intelligence; like those of ants and all lower animals
they prove to be functions of mere sensitive instinct.
It is wrong, therefore, to describe ants as instinctive
automatons, in order to safeguard the intelligence of
the higher animals. The psychic actions of all animals
are due to automatism, as far as they are unable to
attain the level of reasonable reflection and free self-
determination. For the rest, however, there is no
question of automatism either with lower or with
higher animals, because it is sensitive cognition and not
mere reflex activity, which prompts them to act. It
is true, that instinctive actions have a certain auto-

1) Cf. Tylor (in Ranke, “Der Mensch,” II, 1st ed., 486) and

Charles E. v. Baer (in Stoelzle, “Karl E, v. Baer und seine Weltan-.
schauung,” pp. 304, 314).

—:'142 Chapter ITT.

matic character, inasmuch as they are, to a certain
degree, predetermined by the natural constitution of
the animal. However, inasmuch as they are governed
by sensitive cognition and appetite and, therefore,
capable of more or less modification within the prede-
termined natural limits, they are not of an automatic,
but of a spontaneous character.

In conclusion, let us sum up the results of our
comparative discussion on architecture in the animal
kingdom. They are: Ants surpass all animals, both
lower and higher, by the quasi-intelligent variability,
the spontaneous self-determination and the power of
suitable adaptation, manifested in their architectural
skill. Nevertheless it is as certain of them as of any
other animal, that they are not endowed with intelli-
gence properly so called.

_ This corroborates the views advanced in our dis-
cussion on the different forms of learning. Ants are
able to accommodate their buildings to the most varied
conditions; hence they are able to “learn” how to
modify their buildings according to given circum-
stances. But this learning takes place only in so far as
sense-experience gives rise to new combinations of
representations; as soon, however,-as the modifica-
tion of their activity would require intelligent reflec-
tion, or the drawing of conclusions from former con-
ditions to the present ones, then both ants and higher
animals, without exception, are all at once incapable
of further learning. This shows to evidence, that the
doctrine of “animal intelligence” is utterly untenable.
1) “Die psychischen Faehigkeiten der Ameisen,” pp. 82-114; ‘‘In-

stinct and Intelligence in the Animal Kingdom” (Herder, St. Louis,
Mo., 1903), Chap. VIII.

CHAPTER IV.
CARE OF THE YOUNG IN THE ANIMAL KINGDOM,

1. A General View of the Breeding Instincts of
Animals,

UMAN community life naturally evolves from the
family, which must always remain the founda-
tion of the state. Animal societies have a similar
origin and basis, though this similarity does not go
beyond mere analogy. Wherever we meet permanent
associations of animals, they are seen to depend, with
very few exceptions, such as for instance the mixed
colonies of ants, on the ties of common descent. The
purpose of this social co-habitation is the preservation
of the race and species. All other animal instincts are
by natural law subordinated to that higher end, which
is also the reason for the existence of social instincts
of animals.

One of the most important means for preserving
the species is breeding, with the various instincts sub-
servient to it. The different forms of breeding in the
animal kingdom form one of the most interesting
chapters of comparative animal psychology; in this
place, however, we must confine ourselves to some of
its more prominent features.

With those lower animals, which reproduce with-
out sexual generation by fission or by budding, there
can be as little question of breeding instincts as with

148

144 Chapter IV.

plants. The new being comes into existence already
fully endowed with the power of subsistence according
to the laws of merely vegetative nature, whether it be
separated from the mother-organism, or remain united
with it as a new part of a polyzoal colony. Here,
therefore, it would be altogether useless to possess
breeding instincts for the propagation of the species.
Even among such animals as propagate through
sexual generation, we meet with breeding instincts
only where they are required for the preservation of
the species; and the parents care for their off- -
spring only in as far as it is necessary for that purpose.
Within these limits, however, we find a wonderful
adaptation of means to the end, and at times a mar-
velous sagacity of animal instinct, which appears
nowhere else to such advantage.

But also nowhere else in the whole animal psy-
chology are manifested so palpably the impotence of
so-called animal intelligence and the unsoundness of
the modern tendency of humanizing animal. life. How
should the ephemera know by her “own intelligence,”
that she may without any apprehension drop her eggs
into the water? Does she perhaps still remember, that
her “mother” once upon a time dropped her also into
the water as an egg? Or has she perhaps by the
study of zoology gained the knowledge that ephemeras
need no hatching? According to Brehm’s psychology
we ought to give the ephemera a thorough scolding
for showing so little motherly love towards her dear
offspring and for not caring for their welfare. But
scientifically speaking, such a scolding is as nonsen-
sical as if we were to blame an oak tree for bearing

Care of the Young in the Animal Kingdom. 145

acorns instead of pumpkins, or a hen for laying eggs
instead of begetting live chicks. Ephemeras flit
about over stagnant pools and drop their clusters of
eggs into the water; pearl-flies carefully attach their
eggs to a flimsy peduncle consisting of a sap hardened
by exposure to the air; ichneumon-flies deposit their
eggs in the body of a caterpillar by means of their
ovipositor; gall-flies introduce theirs under the rib of
an oak leaf, from which later on the gall-nut is to
grow, serving both as dwelling and as provision store
of the young larva; the blue-bottles place their eggs
on putrefying flesh, whereas a certain species of wasps
(Pompilus viaticus) glue theirs to the bodies of
spiders which they paralyze by skilful thrusts of their
sting without killing them, so as to enable the growing
larvae to feed upon live flesh; the common cabbage-
butterfly deposits her eggs on cabbages, the hawk-
moth on poisonous spurges, the large clavicorn water-
beetle (Hydrophilus piceus) weaves for its eggs an
ingenious boat with a little streamer on top to float
about on the surface of the water, whilst a smaller
allied species (Spercheus emarginatus) carries its eggs,
as many spiders do, in a bag attached to its abdomen ;
the leaf-rolling beetle (Rhynchites betulae) cuts a
birch-leaf in a manner implying a difficult problem in
applied mathematics, and rolls it up into the shape of
an ingenious funnel, in which it deposits its eggs;
whilst Rhynchites pubescens saws a cradle for its eggs
in the wood of an oak-twig, the ear-wig hatches its
eggs like a hen, whilst Lomechusa strumosa, just like
cuckoos, confides its brood to the care of ants; they
all ae aes duty with equal prudence, but all too are

146 Chapter IV.

ignorant of the prudence of their actions, and uncon-
scious of duty. Under the guidance of sensitive cog-
nition and perception they follow the mysterious in-
stinctive impulse arising from their organic develop-
ment, which suggests to them the means necessary for
preserving their species. But they do not understand
the appropriateness of these means and need not reflect
how to use them to advantage.

Wherever in the animal kingdom the care of the
young requires “family life,” i. e., a regular co-habita-
tion of the parents and of the young, this task
takes place only as far as the preservation of the
species renders it necessary. The same organico-
instinctive laws, to which breeding is subjected in
general, also determine the existence and firmness of
family ties as well as the extension of the family circle
with different species of animals. There is no room
for individual reason and liberty; and to postulate
them is not only wholly superfluous, but also contra-
dicted by innumerable facts. As birds associate in
pairs only during the mating season for the preserva-
tion of the species, so in building their nest and in
hatching their young, the two mates co-operate like-
wise only as far as is necessary for preserving their
species; and the pairs remain together and in com-
pany with their young no longer than the same purpose
requires. Altum, in his excellent book “Der Vogel
- und sein Leben”, has supplied us with a number of
striking instances, proving how ridiculous and unten-

1) We have so minutely proved this fact in the case of insects that
live single, in our book, “Der Trichterwickler, eine naturwissenschaft-

liche Studie ueber den Thierinstinct’” (Chap. IV. ff.), that there is no
need of rereating the proof here.

Care of the Young in the Animal Kingdom. 147

able it is to apply to birds the notions of marital and
maternal affection as it exists among men. As a
matter of fact, there is no more “‘marital affection” in |
the human sense of the term to be found with a loving
couple of parrots than with spiders, where the smaller
male must be on its guard not to be devoured by the
larger female immediately after mating. And by
devouring her “husband” the female spider sins as lit-
tle against morals, as she acts conformably to them in
carefully protecting and carrying along her egg-bag
or in spinning a protecting web for her young. And
the female cuckoo smuggling her eggs into nests of
strangers acts as little against morals, as the foster-
parents of the young cuckoos act conformably to
morals in feeding and rearing these changelings.
There is no room for reason and morality in the breed-
ing instincts of animals; for they are exclusively deter-
mined and regulated by the laws of organico-sensitive
life.

The same holds good for mammals, the anthropoid
apes not excepted. As long as young dogs, cats, and
apes need the care of their parents, they will not be
forsaken. But no sooner are they old enough to shift
for themselves, than their parents no longer know
their once so “beloved” offspring. As the mates know
each other only for sexual intercourse, so also they
know their young only as helpless beings, whose
behavior stimulates the nursing instinct of their parents
. to action. As soon as this instinctive impulse ceases,
then the mates and the young are completely estranged,
having no regard for each other in the .relentless
struggle for existence, for food and rut, just as if

148 Chapter IV.

they had never belonged together. This is a genenal
law of nature, ascertained by science throughout the
whole animal kingdom, setting at naught all the fine
phrases and sentimental talk of Brehm on marital and
parental love among animals. And this gush, the
outcome of erroneous notions and misplaced sentiment
is dubbed by thousands of its votaries modern animal
psychology!

Care of the young in its most primitive form is
found among the Echinoderms, namely in a few spe-
cies of star-fishes (Asterias Muelleri, rugispina, Cri-
brella oculata).1 According to Perrier the female ani-
mal, by bringing her arms near to the body, forms
a kind of breeding cavity, in which the young, hud-
dling together, are enclosed. In the different classes
between the Echinoderms and the vertebrates care of
the young assumes very different forms, which we are
unable to discuss here. Of peculiar psychological
interest, however, are those animals, among whom the
males and not the females are entrusted with the care
of building nests and rearing the young. The best-
known example of this kind among fishes is the
stickle-back (Gasterosteus aculeatus).* In this species
the females are regular “cannibal stepmothers,”
whereas the males are models of “affectionate fath-
ers.” How ridiculous such facts are, when couched

1) See H. Ludwig, “Sitzungsber. der Niederrh. Gesellsch. fuer
Naturk.” (Bonn), 1896, 1st half, p. 104; besides in ‘‘Zoolog. Anzeiger,”
1897, No. 534, p. 217 and No. 535, p. 237.

*) Also among amphibias cases of male hatching have been ascer-
tained. See Fr. Werner in “‘Verhandl. der Zoolog.-botan, Gesellsch.
von Wien,” 1898, 1st issue, p. 11 ff. See also R. Wiedershein,
“Brutpflege bei niederen Wirbelthieren” (Biolog. Centralbl. XX, 1900,
Nos. 9 and 10).

Care of the Young in the Animal Kingdom. 149

in anthropomorphic language, goes without saying.’
Some instances of taking care of the young occur also
among amphibias. The female of the Surinam toad
(Pipa dorsigera) carries her young in the cavities of
her dorsal skin; whereas in a frog species (Arthro-
leptis seychellensis) inhabiting Central America the
young hold on to the back of the male.? But quite
universal and commonly known is the care bestowed
by birds and mammals upon their young. Yet its
highest perfection, connected with the most perfect
form of community life in the animal kingdom, does
not occur with the higher mammals, but with the
social insects, in particular with ants. Here this degree
of perfection is made possible by the organic division
of the female sex into females proper and into nurses
(workers) incapable of generation. And although
these are not the mothers of the children they nurse,
the psychic development of their breeding instinct
reaches the greatest perfection in the whole animal
kingdom. Before discussing, however, this aspect of
- the breeding instinct of ants, we must first explain its
connection with the laws of their organic develop-
ment.

The bodily differentiation of the members of an
insect-state into classes and castes, their co-habitation
in a common abode, their nest-construction, acquisition

1) The following amusing quotation will do for the purpose: “The
greatest danger threatens him (Mr. Stickleback) from the mothers of
his own children. Eager to devour their own offspring, they ‘are con-
tinually dashing in unison against the nest, in which the young are
guarded by their watchful father, and but too often the latter pays the
penalty of his polygamy” (Thilo, ‘‘Umbildungen an den Gliedmassen der
Fische,” in ‘‘Biolog. Centralbl.,’”? 1897, 1st issue, p. 24).

2) “Zoolog. Jahrb. Abth. fuer Systematik,” XII (1898), 89 ff.

150 Chapter IV.

of food, their whole life and activity have for their
object the care of the offspring, and thereby the pres-
ervation of the species. The animal colonies of bum-
ble-bees, wasps' and hornets represent a lower stage
of community-life, than the perennial colonies of
honey-bees, ants and termites. In the case of the
former the insect-families and the entire community-
life have to be established anew every year by some
hibernating female; but with the latter the original
families last several years and often much longer, thus
giving their community-life a character of stability
and also of greater variety and perfection.

The organic foundation of insect-states, as already
shown above (p. 14), is polymorphism, or the separa-
tion of the individuals into sexual animals and “neu-
ters”? or workers. The prime duty of the former is
generation, this being the direct means for preserving
the species, whilst the latter perform all the work
required for the welfare of the family, thus in their
turn indirectly contributing to the same end. Without
this appropriate division of labor insect-states would
be impossible; and, as a rule, the more perfect this
division, the more perfectly developed is the insect
community.2 In bees, with whom the workers are

1) According to H. v. Jhering (“Zoolog. Anz.,” Vol. XIX, 1896,
No. 516, p. 449) a large number of the Brazilian social Vespidae
(Polybia, Chartergus, etc.) form perennial colonies for several years,
not annual ones as our native wasps.

2) We have already in Chap. I, No. 2, referred to the fact, that in
reality they are not, properly speaking, sexless.

3) The greatest importance must be attached in this place to the
differentiation between sexual individuals and workers. Thus e. g. in
the annual colonies of bumble-bees there is a dimorphism of workers,
and hence a more marked division of labor than in the perennial
‘colonies of our honey-bee. (On the bumble-bees see esp. E. Hoffer’s
excellent observations on the bumble-bee of Styria). Nevertheless the
bee-states are more perfect than those of the bumble-bees on account of
the greater difference between their workers and genuine females.

Care of the Young in the Animal Kingdom. 151

winged and not unlike the real females, polymorphism
and, in consequence, community life is less differen-
tiated than in ants, where the neuters are devoid of
wings and,—with many species,—are again divided
into different castes, namely, workers and _ soldiers.
The greatest variety of bodily differentiation, how-
ever, obtains in termites, which belong to the insects
with imperfect metamorphosis. In ‘their case the
larva resembles the imago and is transformed grad-
ually with little change of outward appearance. There-
by termites present the organic foundation for a still
more abundant and variable division of castes, the
formation of which may embrace not only sexual indi- —
viduals on the one hand, and workers and soldiers on
the other, but within these two categories again several
different classes or forms.*

Not even the most ardent defenders of modern
animal intelligence would venture to attribute poly-
morphism, which is the fundamental law in the con-
stitutions of insect-states, to the “intelligence” of the
animals themselves. It is evidently based on the
hereditary laws of organic development. Just as it is
not owing either to his own intelligence or that of his

1) Cf. Hagen, ‘Monographie der Termiten” (“Linnaea Entom-
ologica,” X-XIV); Grassi e Sandias, “‘Constituzione e sviluppo della
Societa dei Termitidi,’”? Catania, 1898. (‘‘Atti dell’ Accademia Gioenia
di Scienz. nat.,” 4, VI and VII); Wasmann, “‘Einige neue Termiten
aus Ceylon und Madagaskar,” in ‘‘Wien. Entom. Zeitung,” 1893, 7th
issue; “Neue Termiten und Termitophilen aus Indien” (‘‘Annali del
Museo Civico di Stor. nat. di Genova,” 2, XVI, 1896, 613-630);
“Termiten von Madagaskar und Ostafrika” (“‘Verhandl. der Senkenberg.
Naturf. Gesellsch.,” XXI, 1897, 1st issue); G. D. Haviland, ‘“‘Observa-
tions on Termites” (‘“‘Linnean Society’s Journal, Zoology,” Vol. XXVI,
pp. 358-442). Dr. F. Silvestri “Ergebnisse biologischer Studien an
suedamerikanischen Termiten” (‘‘Allg. Zeitschr. f. Entomol.,’” WIT,
No. 9 ff.).

152 Chapter IV.

“mammy,” that a young rooster became a rooster
and not a jackdaw, so no intelligence of the ant is
responsible for the fact, that from the egg and the
larva which she nurses, there emerges not a bee but an
ant. All this is self-evident. Yet, the peculiar organic
laws of development not only form the material
of the breeding instincts, but also their directing
principle. The breeding instincts of the different
species are so well adapted to the hidden laws of
organic growth, that no reflection or intelligence on
the part of the animal, nay, not even the keenest human
reason could ever succeed in inventing them. More-
over, they are exercised by the workers completely and
perfectly, without previous experience or instruction;
they are innate in the animal, and grow with it,
and when the young ant has reached the perfection of
its organic development, they, likewise, are just as
perfectly developed. Hence they must spring from
_ the same source as the organic growth, that is to say,
they spring from the organico-psychic laws of devel-
opment of a given species, and have nothing to do
with individual reason and free determination. As
it is by organic development, that the male of an
ant-species receives also the psychic endowment of a
male, thus it is with the females and the workers. The
distribution of psychic endowments in the different
castes of an ant-state is regulated by the same laws
as their bodily polymorphism. And this alone accounts
for the fact, that within one and the same species the
males are the most stupid members of the whole state,
possessing the smallest brains, whilst the workers are
endowed with many marvelous instinctive talents and

Care of the Young in the Animal Kingdom. 153

even surpass the females proper in perfection of
instincts and brain development.t Those females,
which are destined for generation, are provided by
their organic development with perfect ovaries, whilst
their brain and instincts are far less perfect. The
workers, on the other hand, which on account of their
small ovaries may be called undeveloped females, are
compensated by a more perfect development of the
brain and the instinctive endowments. Hence, the
astonishing prudence displayed by the worker ants
and their consequent social leadership are merely a
function of their organic development. ‘This is the
so-called “intelligence” and “intellectual life” of ants,
viewed in the light of genuine science!

We have thus far been considering the breeding
instincts of ants from their organic side; let us, in°
the subsequent discussion, turn our attention to their
psychic aspect.

2. Care of the Young among Ants.

The hereditary disposition of the sensitive cogni-
tion and appetite of animals, called instinct, has in the
case of ants a wide range and great variety of actions,
and especially so with regard to the breeding instincts,
wherein ants surpass even the highest mammals. The
instinctive disposition is no mechanical automatism,

1) As to the peduncles of the ant brain, the significance of which
with regard to psychic life we have,already pointed out in a former
essay, Forel says: Les corps pédonculés sont énormes chez les ouvriéres
du genre Formica, qui renferme les fourmis les plus intelligentes; et,
chose trés remarquable, ils sont plus petits chez les femelles et
beaucoup plus petits chez les males du méme genre (‘‘Fourmis de la
Suisse,” p. 123). My own observations confirm Forel’s statements; see
“Instinct and Intelligence in the Animal Kingdom,” p. 130 ff.

154 Chapter IV.

but is guided and influenced by various sensitive
affections and perceptions. In bee-hives the eggs are
simply deposited by the queens in the cells previously
prepared by the workers, and the young bee-larva
goes through the successive stages of development in
one and the same cell. With ants breeding shows far
greater variety and independence. The eggs laid by
the queen are received by the workers and gathered in
clusters of various dimensions. Then from all sides
they are licked again and again with the utmost care,
and begin to increase by the endosmosis of the nour-
ishing juice.t This is the first stage in the rearing of
the young in ant-communities. As soon as the egg
has developed into a larva, there follows the second,
the feeding and nursing of the larvae. When the time
for entering the state of a pupa has arrived, the ant-
larvae are carried by their nurses to a spot covered
with damp earth, whereupon each larva is surrounded
by a case or little dome of earth, within which it spins
its cocoon, enwrapping the whole body. From time to
time some worker comes to see, whether the cocoon is
finished. As soon as it is, it is carefully cleansed of
adhering earth, and is then stored up in a neat little
heap in company with others that have reached the
same maturity. With those ant-species, whose larvae
do not spin cocoons, the larvae are not encased in earth
before their pupation. On that account the ex-
tremely tender skin of the pupa unprotected by a
cocoon requires all the’*more care and caution, lest
grains of sand or mould should enter between the

1) On the growth of ant-eggs see Forel, “‘Fourmis de la Suisse,”

p. 388; it is of minor importance, whether or not the increase in volume
of ant-eggs be called growth in the proper sense of the term.

Care of the Young in the Animal Kingdom. 165

tiny segments of the body, or lest in transportation
any part of the soft pupa be too sharply pinched by
the hard mandibles of the ants, which in this case serve
as hands.

It requires a great deal of attention and skill on
the part of the workers employed in nursing, only to
keep neat and clean thousands of eggs, larvae and
pupae. In earth-nests the moist, soft skin of these
small beings is in continual danger of being soiled by
sand or other foreign matter, and, besides, it is excel-
lent soil for the growth of injurious fungi. ‘ Never-
theless, the ants always keep their brood perfectly clean
so that even under a magnifying lens not a speck of
dust can be discovered. In spite of the damp and
mouldy atmosphere, they are able to prevent entirely
the growth of fungi both on their brood and through-
out the nest. It might perhaps be suggested, that this
is, for the most part, to be attributed to the antiseptic
effect of the formic acid contained in the poison
glands of the ants. But, in reality, the chambers in
which the larvae are kept, show an alkaline reaction,
as was of late pointed out by Ch. Janet,’ who explains
this phenomenon from the fact, that the secretion of
the epidermic, and especially of the salivary glands
of ants is of a basic nature.

The cleaning of the young is only a secondary
occupation in the ant-nursery. But even in this sec-
ondary office the ants surpass all other animals in
care and skill. No cat by licking will wash her kittens
with such exactness and affectionate attention, as ants

1) “Réaction alcaline des chambres et galéries des nids de fourmis”

(“Extr. des Comptes rendus hebdomadaires de l’Acad. des Sciences,”’
CXXVII, 1898, 1380).

156 Chapter IV.

clean the larvae entrusted to them. Therefore we are
right in expecting, that the same perfection of the
breeding instincts of ants will be manifested also in the
other branches which are not less important for the
preservation of the species, namely, in the suitable
regulation of the temperature and in the proper nour-
ishment and defense of the brood.

In the care of the young it is of the utmost import-
ance to regulate the conditions of temperature in a
manner most advantageous to this development. In
bee-hives the position of the brood is determined by
the shape of the comb. Throughout its development
the young bee remains in the same cell, in which it
was placed as egg, and it is therefore constantly sub-
ject to the same conditions of temperature. It is quite
different with ants. Here the temperature has to be
altered and regulated by the workers according to
the different stages of development. The eggs and
the larvae in their earliest stage are generally stored up
in the lowest chambers of the nest, where the air is
cool and damp. Further above the half-grown larvae
are lodged, whilst the uppermost stories are occupied
by the full-grown larvae and the pupae; for these
latter require for their development greater heat, which
is found immediately beneath the surface of the nest
struck by the rays of the sun. If out of doors it
‘grows chilly and rainy, the pupae and. elder larvae
are immediately carried into the lower chambers, where
they are better protected from cold and moisture. This
regulation of the conditions of temperature alone,
according to the necessities of different stages of
development, implies astonishing sagacity, such as even

Care of the Young in the Animal Kingdom. 157

men would be able to acquire only after years of
observation and study. Yet, in ants every single worker
is endowed with it as soon as she is drawn from her
cocoon and has become dry. This is because their
sagacity is instinctive, essentially different from intel-
ligence and reflection. Ants are in their every action
guided directly by sensitive perceptions, not by intel-
lectual ideas. The enigma, therefore, is satisfactorily
explained by the innate adaptation of their sensitive
cognition and appetite, whereas the hypothesis of ani-
mal intelligence is unable to offer any solution.

But now we come to the most puzzling and mys-
terious question in the nursing of ants, namely, the
influence of the education of the young larvae on the
development of different castes in ant-states. Science
has but just now begun to divine the mysteries hidden
here; but it is still far from having fathomed their
depths. We shall very briefly place before our readers
what is certain or at least probable concerning this
matter. It will fully suffice to prove, that the nursing
instincts of ants, bees and termites are far superior
to those of birds and mammals.

According tothe older opinion, thus far commonly
held, and based chiefly on Dzierzon’s classical obser-
vations on bees, the sex of their posterity is determined
by the instinctive choice of the oviparous queen, and
not by the workers that rear the brood. Because it
has been observed, that with ants, bees and wasps
unfertilized eggs produce males only, it is assumed
that also the normal males of these social insects are
always hatched from unfertilized eggs. The queen,
when depositing her eggs, is supposed, by either open-

158 : Chapter IV. :

ing or closing the connection between the oviduct and
the seminal vessel, to control, under the influence of
her oviparous instinct, the fertilization of the egg, and
therefore to decide, whether it would develop into a
male or a female. This instinct of the queen is aroused
to suitable activity by the peculiar nature of the cell,
into which she puts her head before oviposition; in
drone-cells she puts an unfertilized egg, in those of
a future queen or worker a fertilized one. Probably
she is led to make this difference not so much by the
touch-perception of the different shapes of the cells,
but rather by the smell of the salivary gland secretions
employed by the workers in their construction.t The
peculiar odor of the cells, however, as well as their
size and form is due to the architecture of the workers ;
thus, in bees the sex of the offspring is indirectly
at least controlled by the instincts of the workers.
With ants it is different, because their queens do not
deposit the eggs in cells, but simply suffer them to
be received and carried away by the workers. The
instinctive self-determination of the oviparous female
seems, therefore, to be greater with ants. Of course,
it is very probable, that the nourishment and treat-
ment of the queen on the part of the workers indirectly
also influences the oviposition; but in what manner,
is as yet entirely unknown.

To pass from the queen to the workers, it was ascer-
tained long ago, that, with social wasps, bees and ants,
also workers, which have small ovaries with a reduced
number of ovarial tubes, and are, besides, unable to

1) Cf. on this point N. Ludwig, ‘‘Futtersaft oder thierische Veran-

lagung,” p. 32; and p. 57 of the publication of Ferd. Dickel mentioned
below.

Care of the Young in the Animal Kingdom. 159

mate, nevertheless sometimes lay eggs capable of
development. This :phenomenon has been called by
Ch. v. Siebold,t to whom we are specially indebted
for its discovery, virgin-generation, or parthenogene-
sis. Under natural conditions parthenogenesis occurs
with ants principally in colonies which have lost their
queen, and therefore try to rear their posterity from
eggs laid by workers. In several observation-nests
of Polyergus rufescens, Formica sanguinea and rufi-
barbis the queen was missing, and I observed that the
workers,—with Polyergus the slaves,—selected an extra
large worker of the dominant species as a substitute
for the queen, treated her with greater care, gave her
more food, and thus induced her to parthenogenetic
oviposition. This shows that under certain circum-
stances the instinct of ants is able to effect by special
treatment the development of the ovaries of even
adult workers, so as to make them capable of laying
eggs ;? but these unfertilized eggs can only produce

1) “Wahre Parthenogenese bei Schmetterlingen und _ Bienen,”
Leipzig, 1856. Quite recently Ch. Janet has discovered parthenogenesis
also among hornets (Sur Vespa Crabro, Extr. des Mém. de la Soc.
Zool. de France, 1895, p. 75).

2) This form of parthenogenesis, which is spontaneouly caused by
the workers themselves, must be carefully distinguished from another
form caused by artificially raising the temperature, and quite independent
of the ants’ instinct. See my observations in “Biolog. Centralbl.,” XI
(1891), No. 1: “Parthenogenesis bei Ameisen durch kuenstliche Tem-
peraturverhaeltnisse.” The experiments made by E. Bickford (“Ueber
die Morphol. und Physiol. der Ovarien der Ameisen-Arbeiterinnen,”
in “Zool. Jahrb. Abth. fuer Systemat.,” IX, 1895, 1st issue) with
Lasius fuliginosus (p. 19; Sep., p. 23) belong rather to the second
category than to the first, since she too employed artificially raised
temperature. At any rate, they do not approach natural conditions as
closely as my observations mentioned above. On the latter cf. ‘‘Stett.
Entom Ztg.,’’ 1890, pp. 303-305, and ‘‘Biolog. Centralbl.,” 1895, pp.
609 and 610.

160 ’ Chapter IV.

males.t. This phenomenon manifests the marvelous
sagacity and quasi-intelligent plasticity of animal
instinct, which can hardly be styled “automatism.”
Neither can it be identified with intelligence properly
so-called, for this would suppose rational knowledge
of the internal laws governing the growth of the ant-
organism, a knowledge far surpassing even the intelli-
gence of man and entirely beyond the reflections and
experience of ants. Only the appropriate disposition
of their sensitive cognition and appetite can account
for the fact, that the perception of a given want is
followed by a corresponding modification in their
nursing instinct, by which the defect in question is
remedied.

According to Dzierzon’s views, which we men-
tioned above, it is the oviparous instinct of the queens,
that controls the sex of the bee developed from a
given egg; in this supposition the worker-bees are
assigned a merely indirect influence. Of late, how-
ever, another theory on the differentiation of castes
in bees has been advanced, which assigns to the nurs-
ing instincts of the workers a far more extensive
sphere of action. The originator was an Italian priest,
Lanfranchi by name, who published it in 1894 in the
“Apicoltore.” In Germany it was developed and
confirmed by new experiments, principally by Ferd.
Dickel,? the editor of the “Noerdlinger Bienenzeitung.”

1) More recently H. Reichenbach has published some observations
(in “Biol. Centralbl.,” 1892, p. 461 ff.) which seem to prove, that with
Lasius niger the parthenogenetic eggs laid by workers may give origin
also to workers. But further confirmation will be required before
accepting this statement. In North America Prof. W. M. Wheeler has
lately published some interesting reports on parthenogenesis in ants.

2) “Das Princip der Geschlechtsbildung bei Thieren geschlechtlicher
Fortpflanzung, entwickelt auf Grundlage meiner Bienenforschungen.”
Noerdlingen, 1898; cf. especially p. 20.

Care of the Young in the Animal Kingdom. 161

Dickel says: “Under normal circumstances the fer-
tilized mother-bee lays only fertilized eggs; it is the
workers, that influence and control the fate of these
homogeneous eggs.” According to this new opinion,
also those eggs, which in normal bee-hives produce
drones, are fertilized; and not only the differentiation
between queen and worker, but between queen and
drone, and between worker and drone, is due to
the influence of the salivary gland secretions of the
workers on the eggs previously deposited in the cells.
Hence, Dickel regards both queens and drones only
as the foundation for the development of the sexes, the
workers, however, as the really determining factors.
According to Dickel, certain salivary glands of the
workers contain the secretions determining the sex,
and the differentiation of all the castes in bee-hives
depends on the instinctive application of these secre-
tions, when the workers are licking the eggs.
Although several biological experiments of other
authors seemed to confirm the theory that under nor-
mal conditions all the eggs in a bee-hive are fertilized,’
we must add, nevertheless, that the very exact micro-
scopical studies of Paulke and Petrunkewitsch on the
existence or non-existence of spermatozoids in the
eggs of bees rather corroborate the old theory of
Dzierzon, according to which the eggs giving origin
to drones develop without containing any spermato-

- 1) Cf. N. Ludwig, ‘Neues ueber Ernaehrungs- und insbesondere ueber
Fortpflanzungsverhaeltnisse der Honigbiene” (‘‘Natur und Offenb.,”
XLIV, 1898, 12th issue, pp. 705-719), ““Weiteres zur neuen Lehre ueber
die Geschlechtsbesti g der Bienen” (ibid. XLV, 1899, 3d issue, pp.
140-148; “Weitere Ergebnisse ueber die Fortpfl gsverhaeltnisse der
Biene” (‘‘Natur und -Offenbarung,” 1901, 7th issue, pp. 426-430).

162 Chapter IV.

zoids. It is best, therefore, to suspend our judgment
on this problem, until it is definitely solved.

If the new theory of the fertilization of all the eggs
in a bee-hive, under normal conditions, should prove
true, it ought to be extended also to the ants. Hence,
in ant-colonies, too, it would be the workers, who by
their nursing instincts determine, whether a given ant-
egg is to produce a worker, a winged female, a soldier,
or a male.

We intend here, however, to consider only the
wonderful influence exercised by nursing on the dif-
ferentiation between females and workers. This is an
established fact, quite independent of the new theory.
Why is it that one and the same kind of egg now
produces a queen with complete power of generation,
now a worker devoid of generating powers, but com-
pensated, as it were, for this loss by psychic endow-
ments all the more perfect? Here we enter a mys-
terious region, where the breeding instincts of social
insects reign supreme, an instinct which for its creative
power is unparalleled in the entire animal kingdom.

It is a well-known fact, that with honey-bees a
worker larva can be developed into a queen by increas-

1) W. Paulke, in ‘“‘Anatomischer Anzeiger,” Vol. XVI, 1899; A.
Petrunkewitsch, “Tie Richtungskérper und ibr Schicksal im befruchteten
und unbefruchteten Bienenei,’” (Zool. Jahrbuecher,” Abtl. fuer
Anatomie, Vol. XIV, 1901); Aug. Weismann, “Ueber die Partheno-
genese der Bienen” (“‘Anatom. Anzeiger,” Vol. XVIII, 1901, Nos. 20-21);
H. v. Buttel-Reepen, ‘‘Ueber die Dzierzon’sche Theorie” (‘‘Bienen-
wirtschftl. Centralbl.,”’ 1901, No. 1); “‘Der Abschluss der Freiburger
Eiuntersuchungen” (Ibid., 1901, No. 19); “Die Parthenogenesis bei
der Honigbiene” (“Natur und Schule,” Vol.I, 1902, 4th issue); P.
Bachmetiew, “Ein Versuch, die Frage ueber die Parthenogenese der
Drohnen zu lesen” (“‘Allgem. Zeitschr. f. Entom.,”’ 1903, Nos. 2-3).

Care of the Young in the Animal Kingdom. 163

ing its cell and giving it different food.t Also in the
case of termites, so we are informed by Grassi and
Sandias,? the various methods of nursing the larvae,
and especially the different salivary gland secretions of
the nurses are of great importance in the differentia-
tion of the castes of one and the same species. The
same probably holds good with ants,* and, indeed, with
far more variability than with honey-bees, although less
than with termites. This is indicated by the numerous
intermediate forms between the females and workers
of ants. Of course, wherever these appear, they are
exceptional forms, yet none the less they throw some
light on the origin of the normal differentiation into
females and workers; for, their existence is most
intelligible on the supposition, that the difference of
caste is not predetermined in the egg, but that it will
depend on the nursing, whether the fertilized egg will
bring forth a winged, perfect female, a normal worker
or perhaps some intermediate form.

Of course, the specific development peculiar to
every ant-species is the necessary foundation for the
differentiation of the normal castes and for the origin
of certain abnormal, intermediate forms. Where there

1) Cf. N. Ludwig, “‘Futtersaft oder thierische Veranlagung.”—
According to Planta’s tables the food of queen-bees contains a far
larger amount of fat. But according to Ludwig it is especially the
different quality of the saliva, added by the bees to the nutrifying
juice, which is of decisive importance.

2) “Costituzione ¢ sviluppo della Societa dei Termitidi” (Catania,
1893), pp. 75-106.

8) Emery, ““Le Polymorphisme des fourmis et la castration alimen-
taire,’’ Leyden, 1896 (Extr. du Compte rendu des Séances du 3me
Congrés internat. de Zool., p. 395 ff.).

*) Wasmann, “Die ergatogynen Formen bei den Ameisen und ihre
Erklaerung” (‘‘Biolog. Centralbl.,’’ 1895, Nos. 16 and 17).

164 Chapter IV.

is no possibility of developing into certain forms, there
is, of course, no basis, no material for the exercise of
the nursing instincts of the workers. From this it
becomes clear, why fixed intermediate forms between
females and workers occur with certain species, dif-
ferent forms with other species, whilst with others
again there are none at all. But, within the limits of
this natural disposition for further development, there
remains to the ants a wide range for exercising their
nursing instincts.

These intermediate forms between females and
worker ants I have grouped into six classes, but here
we are concerned with but one or two of them. In
some of these “ergatogyne” forms it appears almost
at a glance, how they came into existence, namely,
whether the larva, reared up to a certain stage to be
a worker, was later on cared for so as to become a
female, or whether the opposite took place. In the
first case the intermediate form makes the impression,
that the worker-character had been developed to
excess; in the second case, that the female character
had been stunted; in the former the so-called worker-
like (ergatoid) queens are the result, in the latter a
kind of female-like workers, which I have named
pseudo-females (pseudogynes). The former combine
the vaulted thorax of females with the small and
abdominal development of queens; the latter unite
the vaulted thorax of females with the small and
stunted abdomens of workers. Especially the latter
form, the pseudogynes, are apparently best accounted
for on the score of education rather than by a peculiar
disposition inherent in the egg, from which they are

Care of the Young in the Animal Kingdom. 165

hatched.1 I know some colonies of Formica san-
guinea, near Exaten, in which these pseudogynes sud-
denly made their appearance, became more numerous
in the course of the next few years, and gradually
decreased later on, or disappeared entirely. One
colony (No, 21) in 1895 reared every possible pseu-
dogyne intermediate form between normal workers
and normal queens! Since the queens which lay the
eggs in these nests are unable to change at will the
nature of their ovaries from year to year, but are
always compelled to lay fertilized eggs, capable of
equal development, the origin of those intermediate
forms is probably due to changes in the manner of
nursing, and to modifications in the very nursing
instincts of the workers. This supposition is con-
firmed by the fact, that in F. sanguinea there is a
certain causal relation between the origin of pseu-
-dogynes and the education of the larvae of a genuine
ant-guest, the beetle Lomechusa strumosa. I have
ascertained this mysterious connection by means of
my statistics embracing 410 sanguinea colonies within
a radius of several kilometers around Exaten; these
statistics will be published later on in some scientific
periodical.? Here it may suffice to mention, that the
centres of propagation of the pseudogyne forms and
of the Lomechusas are always together in the same

1) See my recent publication, ““Neue Bestaetigungen der Lomechusa-
Pseudogynen Theorie’ (‘“Verhandl. der Deutsch. Zool. Gesellsch.,’’ 1902,
pp. 98-108 and Pl. II), where this theory is extended also to North
American ants. See below (the following section, p. 179 foll. and the
plate opposite p. 181).

2) The beginning of those statistics dates back to 1895 (“‘Die

ergatogynen Formen bei den Ameisen und ihre Erklaerung,” “Biol.
Centralbl.,’”? 1895, Nos. 16 and 17).

166 Chapter IV.

or at least in neighboring nests. The number of
colonies, in which I found Lomechusas (100), is more
than ,three times as great as that of nests containing
pseudogynes (33); these are the centers, from which
the Lomechusas gradually spread to the neighboring
nests, where by and by they cause the birth of pseu-
dogynes. It is scarcely possible, that the presence of
these beetles and of their larvae, which are fed by the
workers, should have a modifying influence on the
ovaries of the queens, but, probably they do so on
the nursing instincts of the workers. My recent
observations and experiments until 1904 have con-
firmed this solution of the interesting problem.
Some of the above mentioned intermediate ant-
forms are useful for the preservation of the colony
and the species, whilst others are more or less indif-
ferent, and still others positively hurtful, being prob-
ably pathological deformities. The rearing of worker-
like, wingless queens among the Amazon ants (Polyer-
gus rufescens), for instance, is very appropriate,
because their colonies are rather rare and far distant
from one another. Therefore, the probability is very
slight, that on their nuptial flight the winged sexes
will meet with those of other colonies; and besides,
this species has to encounter exceptional difficulties in
founding new settlements by means of single fertilized
females, since the Amazons are entirely dependent on
the help of their slaves. The wingless queens, on the
other hand, cannot go far from their nests; and after
they have been impregnated by the winged males,
some strolling slaves can easily find them in the
neighborhood and bring them home again; hence it

Care of the Young in the Animal Kingdom. 167

is very suitable, that just Polyergus should so often
rear ergatoid queens. Nor is this arrangement in
any way due to the “intelligence” of the Amazons; for
the education of their offspring is entirely committed
to the care of their slaves (mostly F. fusca) ;+ these
slaves, however, have been robbed from colonies which
do not rear ergatoid females, and neither reflection
nor experience could have given them intellectual
knowledge of the requirements peculiar to the nursing
of Polyergus. Here animal intelligence is entirely
powerless. If F. fusca, the slaves of the Amazons, rear
the offspring of their masters in a way suited to the
preservation of exactly this species, then we must
admit, that the nursing instincts of the slaves are
influenced and modified by the peculiar sensations
caused by the Polyergus-nests.

But what shall we say of the rearing of pseu-
dogynes with F. sanguinea? This combination of
female and worker is decidedly injurious to the
preservation both of the colonies and of the species.
The pseudogynes are stunted beings, neither workers
nor females, unfit to participate either in building the
nest or in nursing the young,’ in defending the colony
or in propagating the race; in fine, they are down-
right failures. It is evident, that their origin is not
due to the “individual intelligence of the ants”; for,

1) Near Exaten, Holland, all Polyergus-nests, I met with, contained
F. fusca as slaves; those which I found in Bohemia (Mariaschein),
Austria (Vienna) and in Luxemburg, had F. rufibarbis as slave species.
The ergatoid queens I met hitherto only in nests with fusca-slaves.

2) It happened only very seldom (among several hundred observa-
tions only five times), that, on the nest being exposed to the light, a
pseudogyne seized and carried away an ant-larva, whilst workers are
always wont to do so,

168 Chapter IV.

if they had but a spark of intelligence, sad experiences
would have enlightened them long ago on the folly
of this mistake. Nay more, if the pseudogynes owed
their origin to the normal nursing instincts of ants,
we should have to despair of the fitness of animal
instinct and even of the wisdom of the Creator. What
is the key to this mystery? It is the beetle Lomechusa
strumosa. According to our hypothesis the rearing
of the pseudogynes is an aberration of the breeding
instinct of ants, caused by the continuous education
of Lomechusa-larvae. In the economy of nature it
is the duty of this beetle, to check the excessive
increase of the ant-species, whose hospitality it enjoys.
For this reason its larvae not only consume countless
ant-eggs and ant-larvae,—the ants calmly looking on
‘the while,—but by destroying the offspring of the
ants, and by the care which the ants bestow on them,
they cause the degeneration of the normal nursing
instincts of the workers, resulting in the education of
crippled pseudogynes.t. To account for these facts on
the score of “individual animal intelligence” would

1) These expositions will probably suffice also to refute an objection
raised by Dr. G. Adlerz, who, misunderstanding my psychological ex-
planation of the rearing of pseudogynes, says in the third part of -his
valuable “Myrmecologiska studier’ (Stockholm, 1896), p. 51: ‘With
regard to this Wasmann seems inclined to credit ants with an exag-
gerated power of reflection, which he otherwise is unwilling to do.”
Besides, the pathological degeneration of the breeding instinct explains,
why the rearing of pseudogynes is still continued, even when colonies
have been deprived of their Lomechusas. By the way, let me repeat
a remark formerly made, that the causal connection of pseudogynes with
the Lomechusas is not to be confounded with the explanation of this
connection. The former seems to be firmly established by direct observa-
tion, the latter is still an hypothesis, but an hypothesis, strongly con-
firmed by recent experiments of myself and of Vichmeyer. See ‘Neue
- Bestaetigungen der Lomechusa-Pseudogynentheorie” (‘‘Verhandlungen
der Deutsch. Zool. Gesellsch.,’’ 1902, p. 98 ff.).

Care of the Young in the Animal Kingdom. 169

involve us in endless and hopeless contradictions.
They are explainable only from the standpoint of a
higher, teleological consideration of nature, which
does not presume to replace the wisdom of the Creator
by the “intelligence of animals.”

The phenomena in the nursing of ants mentioned
above can be ascertained only by close, scientific
observation. But some other features are known to
every amateur in the study of ants, and do not escape
even the most casual observer. The first thing that
strikes our attention is the great attachment displayed
by the. workers for their charges. They carefully
guard them against every disturbance, and at the risk
of their own lives they exert all their strength in ward-
ing off hostile invaders. The whole colony is seized
with frenzy, if an attempt is made to rob them of
their larvae and pupae.t. Merely thrust your stick
into a hillock of wood-ants! At once there ensues a
tumultuous uproar and masses of workers rush forth
to rout the enemy. But if you happen upon a chamber
filled with pupae and attempt to take away the cocoons,
the fury of the ants reaches its climax. Like an army
of raging furies they fall upon the assailant, viciously
biting and ejecting their poison. Hundreds and thou-
sands are crushed by the enemy, but other hundreds
and thousands are eager to face the carnage. No
lioness, no she-monkey ever defends her young with
the heroism displayed by ants. Workers will rather

1) This is the case with species otherwise very peace-lovirig, e. g.,
with the large American leaf-cutting ants of the genus Atta. On open-
ing a nest of Atta sexdens in Rio Frio, Forel even had an artery of his
little finger pierced by a large-headed worker. See Forel, “Zur Fauna

und Lebensweise der Ameisen im Columbischen Urwald” (“Mittheil. der
Schweiz. Entom. Gesellsch.,”? Vol. IX, 9th issue, p. 407).

170 Chapter IV.

suffer their heads to be torn off, than yield to the
enemy the pupae they carry in their jaws. And yet,
it is not even for their own children, that they ‘‘sacri-
fice themselves so unselfishly ;’ their charges are but
their foster-children. But that higher natural law,
which has made preservation of the species the fore-
most instinctive commandment implanted in the ani-
mal soul, this natural law, I say, also constrains the
worker-ants to risk life and limb in behalf of beings
begotten by others. This commandment they observe
faithfully, not led by any sense of duty or by noble
forgetfulness of self, but by an irresistible, instinctive
impulse implanted in them by Another, and to which
they yield obedience, not intelligently or voluntarily,
but urged on by a blind necessity of nature!

To credit animals with intelligence, to ascribe to
them ever so faint a trace of intellectual knowledge of
the purpose of their actions, will necessarily lead to
extolling the self-sacrifice of the single workers for
the welfare of the colony and especially for the young,
as a high degree of quasi-human, nay of superhuman
virtue. And in fact, L. Buechner, E. Haeckel, Th.
Eimer, O. Zacharias and other modern animal psychol-
ogists have actually ventured such assertions.1 Of
course, their only commendation is their boldness, but
it is a boldness leading to the greatest absurdities.

What is it then, that impels the ants to such heroic
devotedness and self-sacrifice for the offspring of
their colony? Is it perhaps “motherly love’? No;
for the workers are but the sisters or aunts of their

1) Wasmann, “Die zusammengesetzten Nester und gemischten
Kolonien der Ameisen,” pp. 190 and 191,

Care of the Young in the Animal Kingdom. 171

charges, since under ordinary circumstances the eggs
are laid only by the impregnated females. Their
affection, therefore, as sisters or aunts is the psychic
impulse of their nursing instinct. According to
modern animal psychology, which ascribes to animals
besides their instinct at least a modicum of genuine
intelligence, it cannot be doubted, that the worker-
ants fulfill their duties as sisters or aunts “very know-
ingly,” that they are aware of the importance of the
work allotted to them in their social economy, and
that they apply themselves to nursing the offspring
of another with the consciousness of doing their duty.
But to what degree will the love of the workers for
the young be advanced, if to their affection as sister
or aunt there is added “motherly love’ in the full
sense of the term? Must we not expect that the love
of the workers for their own young should attain to
an exalted, an unutterable degree of tenderness? For,
in the whole creation no love is stronger than that of
a mother.

Indeed, our expectations would be justified, if ants
were endowed with intelligence and self-consciousness.
But what do we find in reality? The workers gen-
erally devour most of the eggs, which they have laid
themselves.1. Is this the climax of noble, self-sacri-
ficing motherly love? Or shall we call these workers
abominable cannibals regardless of duty? The psy-
chology of Brehm and others of that ilk may decide
this question. In our opinion, however, such facts
ought to lead reasonable people to perceive the obvious
contradictions, in which all the talk about “animal

1) See my observations-in “Biolog. Centralbl.,” XI, 1891, p. 21 ff.

172 Chapter IV.

intelligence” and ‘animal ethics’ is hopelessly
involved,

The nursing instinct of ants with all its “devoted-
ness and unselfishness,” is therefore nothing else than
a purely instinctive impulse guided and determined
in its operations only by sensitive impressions and not
by intellectual concepts. Under normal circumstances
this instinctive impulse is appropriately regulated, and
manifests itself as the product of “self-sacrificing
sisterly love.” But, if the abnormal irritation of the
nervous system of the ants, caused by parthenogenesis,
has disturbed the normal sphere of sensitive impres-
sions, then sisterly love is not changed into motherly
love, but into “barbarous, unfeeling cannibalism”’!

Modern animal psychology evidently toys in a
rather frivolous manner with the term “motherly
love,” by applying it to the nursing instincts found
among animals. Nor can the plea be advanced, that
with higher animals matters are quite different than
with ants; for, we have proved above, that the nurs-
ing instincts of ants far surpass in perfection those of
birds and mammals, not only by their quasi-intelligent
self-determination in the method of education, but
also by the great unselfishness manifested in nourish-
ing and defending their young. If there should be
any difference at all, it is in this, that in the care of
their offspring the higher animals betray far less
“intelligence” and far less “individual liberty,” than
is found in ants. Moreover, it is a well-known fact,
that domestic pigs not seldom devour some of their
litter; yet pigs are “higher animals.” In such cases,
however, the sow sins as little against good morals,

Care of the Young in the Animal Kingdom. 178

as worker-ants do by devouring their own eggs; for,
morality presupposes reason and free-will, reflection
and consciousness of duty, all of which are wanting
throughout the animal kingdom, being the exclusive
privilege of man.

That animals in caring for their young are not led
by reason, but only by sensitive emotions and repre-
sentations, becomes evident especially from the phe-
nomena of adoption in the animal kingdom. There-
fore these shall form the subject of the following
section.

3. Adoption Instincts in the Animal Kingdom.

The tendency to adopt the offspring of strangers
is shown by all those animals which, to preserve their
species, are forced to bestow great care on their own
progeny. This tendency is found among ants not
only with regard to the eggs, larvae and pupae of
other colonies of their own species or of allied species,
but also with regard to members of altogether dif-
ferent orders of insects, living in their communities.
These adoption instincts are responsible for the mixed
colonies of slave-making ants, the robbed pupae of the
slave-species being nursed as carefully as others, either
by the masters or by the slaves already present in the
nest. To the same instinct of adoption must be
referred the care bestowed by the ants on their gen-
uine guests or other nest-mates belonging to different
orders of insects, but above all, the solicitude
with which they rear the larvae of certain beetles
(Lomechusa, Atemeles, Xenodusa) and the eggs of
several kinds of plantlice. The adjoining illustration

174 Chapter IV.

shows the beetle Lomechusa strumosa so often referred
to, and one of its larvae, magnified to four times their
natural size.

Fig. 4. Fig. 5.
Lomechusa strumosa F. Fullgrown larva of Lomechusa
(Magnified. ) strumosa. (Magnified.)

The same adoption instinct occurs also with birds,
although not so seemingly intelligent as with ants.
The best-known example is the hen, that readily
hatches eggs of other hens, ducks, geese, turkeys,
etc., and extends to all her adopted children the same
“motherly care,” she would show to her own chicks.
G. Romanes? succeeded even in making a hen the
foster-mother of some young ferrets, which he had
substituted for the artificial eggs, on which she was
hatching. The numerous species of birds, which
tend the young cuckoos, follow the same line of con-
duct, the only difference being, that they lavish still
greater care on these changelings, because they open
their mouths wider in crying for food than their own
nestlings. The adoption instinct, finally, is met with
among mammals, the most blood-thirsty carnivores not
excepted. Though it is a fable, that ancient Rome

1) “Mental Development in the Animal Kingdom.”

Care of the Young in the Animal Kingdom. 175

owed its origin to the adoption instinct of a she-wolf
that gave suck to Romulus and Remus, yet similar
facts are fully authenticated; for instance, that suck-
ling cats adopt young rabbits and squirrels.1 Espe-
cially among apes this instinct is quite prominent, but
much more characteristically unreasonable than with
any other animal. Of course, for a certain kind of
modern animal-psychology, which is accustomed to
the most superficial observation, such occurrences are
noble manifestations of human compassion, and of an
abundance of “motherly tenderness” lavished on the
offspring of strangers. But accurate, scientific obser-
vation and critical investigation prove the very con-
trary, namely, that throughout the animal kingdom
the nursing instinct is but a sensual impulse, unac-
companied by individual intelligence or individual
morality.

If a hen calmly continues trying to hatch pieces of
limestone or links of iron chains put in place of her
eggs, she can hardly be said to be actuated by
“motherly love.” Animals merely endeavor to satisfy
their instinctive breeding impulse; the higher pur-
poses of their action are altogether unknown to them.
William of Reichenau relates that a bitch, being rob-
bed of her pups, fetched an old pair of slippers and
tried to suckle them.?, Whether she thereby intended
to allay her pangs of conscience as to the fulfillment
of her “maternal duties,” animal psychologists a la
Brehm will be better able to decide than we, to whom
such facts merely prove, that the nursing instinct in

1) Cf. W. Herd, in “Scottish Naturalist,” 1872, p. 155.
2) Cf. “Kosmos,” 4th year, VII (1880), p. 217,

176 : Chapter IV.

animals is a sensual impulse, not guided by reason
and reflection.

This organico-sensitive nature of the nursing
instinct also explains, why-it often extends to the
helpless offspring of other species, whose instinctive
behavior is somewhat similar to that of the animal’s
own progeny. The sense-perception of these helpless
beings stimulates the nursing instinct of the old ones,
and therefore they “adopt” the young. ones of
strangers. The smell of the larvae of Lomechusa
strumosa is especially attractive to the sanguine slave-
makers; besides these larvae instinctively mimic the
attitudes and behavior of the ant-larvae, and although
they possess six feet, they do not make use of them,
but conduct themselves like helpless ant-larvae. For
these reasons they enjoy the most careful attention on
the part of their hosts. And as these beetle-larvae,
when fed by the ants, grow much faster than the ant-
larvae, they impress the instinctive nursing impulse of
the ants far more favorably than the latter, and hence
are the objects of “greater tenderness.” At any dis-
turbance of the nest the workers first care for their
“adopted children” and bring them to a place of safety,
before they attend to their own offspring; yea, they
even neglect the rearing of the latter, their only care

1) That the ants do not confound those coleopterous larvae with
their own on account of their shape and color, I ascertained, in May,
1897, by experiments with larvae of Anthonomus pomorum, which are
far more similar to ant-larvae than those of Lomechusa. The Antho-
nomus-larvae were instantly seized as prey and torn to pieces by the
sanguineas of my observation nest. On the whole, it must not be
imagined that the Lomechusa-larvae make the same impression as their
own on the sensitive perception of the ants; the impression is at most
similar, but more agreeable, which probably explains why the ants prefer
the adopted larvae to their own.

Care of the Young in the Animal Kingdom. 177

being their Lomechusa-larvae, so dear to them on
account of their quicker growth and their better
appetite. It does not affect the ants in the least, that
the Lomechusa-larvae again and again devour the
eggs and young larvae of the ant colony by the whole-
sale; on the contrary, they even carry these change-
lings to the clumps of eggs and larvae to facilitate
their work of destruction. Unless a superior Wisdom
had provided that the ants themselves, by their stupid
affection, prevent the excessive increase of the Lome-
chusa population, the number of these guests would
become so large as to destroy all the sanguinea-
colonies. But there is no danger of any such calamity ;
for, the ants deal with the Lomechusa-larvae during
their pupation just as they do with their own, imbed-
ding them carefully in a vault of earth. After a
short time, the ant-larvae having meanwhile spun
their cocoons, are again removed from the earth.
This latter measure applied to the Lomechusa-larvae
proves fatal to them. The larvae of these beetles do °
not spin a solid cocoon, but only an extremely flimsy,
silken web, which tears as soon as they are unearthed ;
soon after the Lomechusa-larvae are again carefully
imbedded at some other place, then they are taken
out again, carried about, again imbedded, until at
length they become dry and perish. In this manner
the folly of the ants causes most of the Lomechusa-
larvae to die before they are changed into pupae; and
even those, which have fortunately entered the state of
a pupa, are often unearthed by the ants and devoured, .
—perhaps from an excess of affection? According
to my long continued observations on the development
12

178 Chapter IV.

of Lomechusa strumosa in normal sanguinea-colonies
only those larvae escape destruction, which, after hav-
ing been imbedded in their little cave, are forgotten
by the ants; all the rest are hopelessly doomed; of
100 larvae, therefore, at most about Io reach the stage
of imago, sometimes scarcely one. For instance in
the observation nest illustrated on p. 23 in May, 1896,
about 150 Lomechusa-larvae, the offspring of Io
Lomechusas, were reared under the most favorable
conditions both of nutrition and temperature; from
these 150 larvae I obtained—one single Lomechusa!
For thousands of years and in thousands of normal
colonies F. sanguinea year after year repeat the same
senseless performance: first, with the greatest
devotedness they nurse the Lomechusa-larvae, even
allowing their own offspring to be devoured by them;
then, their stupid affection does not allow them to
leave the larvae in peace during the time of pupation,
and finally they devour the pupae. They cannot see
that during their pupation Lomechusa-larvae are to
be treated differently from those of ants; but this is
their salvation; for otherwise their care of the Lome-
chusas would long ago have brought about the ruin
of their own race. One and the same superior Wis-
dom has designed, that on the one hand the increase
of the ants be checked by their inconsiderate love for
Lomechusa strumosa and for their larvae, and that on
the other hand the spread of this beetle be kept within
limits by the very same unreasonable affection of the
ants. By these means so gentle and yet so effective,
Divine Wisdom is able to maintain the equilibrium in
nature. In the face of such phenomena, the defenders

Care of the Young in the Animal Kingdom. 179

of animal reason and animal ethics stand utterly
bewildered.

We have seen so far, how the Lomechusa-larvae
are treated in normal sanguinea-colonies, in which the
education of Lomechusas dates but one or two years
back. According to my observations during the last
years, the case is, however, different in colonies, where
the influence of the parasitic larvae of that beetle has
caused the appearance of those crippled, intermediate
forms between females and workers, which we have
called pseudogynes. For the reason mentioned above,
only a few Lomechusas are developed, as a rule, in
the normal colonies, whereas in colonies containing
pseudogynes, many more of the beetles pass their
pupation successfully, because the pupae generally
remain undisturbed. This accounts for the fact, that
the sanguinea-colonies containing pseudogynes are the
centers from which the rapidly multiplying Lomechusa
infests also the neighboring nests. One instance may
be mentioned here. Colony No. 191 of my statistical
map in the middle of May, 1898, had from 2 to 3%
of pseudogynes; in August, however, the number of
pseudogynes newly developed during summer had
risen to 30%! In order to ascertain, how many beetles
had been reared in this colony in 1898, I dug up the
nest at the end of September and found among the
ants 116 Lomechusas' snugly ensconced in their winter-

1) From this number the 30 Lomechusas were taken, which I put
in my sanguinea nest at home and the greater part of which were again
driven out by the ants (see p. 59). This surprising conduct is some-
what accounted for by the fact that the colony of my observation nest
belonged to the normally developed class, and not to such as contained
pseudogynes,

180 Chapter IV.

quarters. The rapid increase of pseudogynes in that
colony, therefore, was in proportion to the number of
beetle-larvae which had successfully developed in the
same nest that year.

It is, therefore, necessary to assume, that in san-
guinea colonies the frequent rearing of Lomechusa-
larvae gradually modifies the normal nursing instinct
of the ants. This modification is manifested partly by
the production of the crippled pseudogynes, partly by
the more appropriate treatment of the Lomechusa-
larvae which, after having been imbedded in their
cradles, remain undisturbed. Ants, therefore, grad-
ually learn to modify their nursing instinct. Is not
this a proof of intelligence? True, their sensitive cog-
nition guiding their instinctive activities may furnish
the immediate occasion for that two-fold modification.
But we have proved in a former essay, in discussing
the different forms of learning,t that not every modi-
fication of the hereditary instinct, occasioned by sense-
experiences, is due to intelligence, but only that, which
manifests a knowledge of the appropriateness of a
given action. If ants were gifted with intelligence,
they could not help understanding, that by improving
their treatment of the Lomechusa-larvae, they cause
their colony but to perish the sooner, just as they con-
demn it to utter destruction by rearing pseudogynes.
The latter modification of the nursing instinct, which
leads to the rearing of cripples, can only be a
pathological symptom, pointing to a morbid dis-
turbance of the normal, organic condition of that

1) “Die psychischen Faehigkeiten der Ameisen,” p, 111; ‘“In-
stinct and Intelligence in the Animal Kingdom,” Chap. 8.

Fig. 1.

Form. san-
guinea Ltr.,
subsp. 7udz-
cunda Em.
Normal dea-
lated queen
from Prairie
du Chien,
Wisconsin,

U.S.A.

Fig. 2.

Form, san-
guinea Lir.,
subsp. ~udz-
cunda Em.
Normal wor-
ker, from the

same __ nest.

Fig. 3.
Formica san-
guinea Lirs

subsp. rudbicun-
da Em, Pseudo-

gyne from the

same nest.

Le)

(All figures magnified to seven times their natural size.)

Fig. 4.

Xenodusa cava
Lec., whose lar-
val
caused the rea-
ring of Pseudo-
gynes in
same colony of

education

the

ants.

Care of the Young in the Animal Kingdom. 181

instinct. Hence, the improved education of the
Lomechusa-larvae which is invariably attended by an
impaired education of their own larvae, is not owing
to intelligence on the part of Formica sanguinea,
but to a disturbance of their normal, instinctive dispo-
sition, occasioned by nursing the strangers.

What we have said here of the rearing of our
European Lomechusa strumosa by the sanguinea, has
its exact counterpart in North America, where the
larvae of Xenodusa cava, a species nearly allied to
Lomechusa, are educated by the North American race
of F. sanguinea, which Emery has named F. rubi-
cunda. Rev. H. Muckermann, S. J., of Prairie du
Chien (Wisconsin) has succeeded in finding also the
pseudogynous ant-form in an infested nest of F. rubi-
cunda. In the Verhandlungen der Deutschen Zoo-
logischen Gesellschaft(1902 p. 98-108) I have given
an account of these observations, which are illustrated
on Plate II of the “Verhandlungen.” We give here
a copy of this plate, to show more clearly what pseu-
dogynes (fig. 3) are, and how they differ from the
normal queen (fig. 2) and the normal worker (fig. 1)
of the same ant-species. The malefactor, Xenodusa
cava, is photographed in fig. 4.

To be sure, at the first glance the care bestowed
by ants on other animal species, their guests or nest-
mates, often looks like intelligence. This explains to
a certain extent, why modern animal psychologists
attempted to utilize these occurrences as arguments
for the great intelligence of ants. This attempt was

1) Of late Father Muckermann himself has published an illustrated

account of his discovery in the ‘Entomological News,” (Philadelphia),
December, 1904.

182 Chapter IV.

made, e. g., by William Marshall in his “Leben und
Treiben der Ameisen” (p. 102), where he speaks of
the care given by ants to the eggs of plantlice. Sev-
eral ant-species of the genus Lasius collect the eggs
of certain Aphides in their nests. Being carefully
protected during winter, the young aphides in spring
are carried to the plants on which they find their food.
Thence Marshall infers, that the ants tend the eggs
with the intelligent purpose of enjoying later on the
sweet secretions of the aphides. “This is surely a
very strange phenomenon,” he says, “which proves
perhaps better than anything else the high degree of
intelligence attained by ants. We must credit them
with a considerable power of observation, and we must
own, that they have studied, to a certain degree, the
habits of their domesticated animals,” etc. Yet this
bold conclusion is entirely unfounded. How does
Marshall know, that the ants gather the eggs of the
aphides with the intelligent purpose of rearing
aphides? That.there is some connection between the
eggs of the aphides and the aphides themselves is,
indeed, for many ants a subject of sensitive knowl-
edge and experience; but it is unwarrantable to mis-
take this process of instinctive association for intelli-
gence proper. Even if ants in reality tended the eggs
of aphides only on account of a combination of their
sensitive experiences, this would be as yet no proof
of their intelligence, but merely of their memory.
In reality, however, the case is different. Take a few
newly developed workers of a Lasius nest and unite
them to form an autodidactic colony, restricted to its
innate instincts without a shadow of experimental

Care of the Young in the Animal Kingdom. 183

knowledge as to the development of aphides. Entrust
them with eggs of those aphis-species, which their
congeners are wont to rear and to nurse. They will
treat them as though they had previously “studied”
the habits of these aphides! Hence the fondness of
certain ant-species for the eggs of aphides is a merely
instinctive impulse, which, of course, can be strength-
ened by sensitive experience. It was rather rash for
Mr. Marshall to proclaim it boldly as “a faculty of
taking the future into account.’ Alfred Espinas was
far more correct in calling the aphis-nursing of ants
an “intelligence non réfléchie,” i. e., merely analogous
to human reason, having but a faint similarity to
intelligence proper, the difference being not merely of _
degree but of kind.t| This analogum rationis is simply
an instinctive association of representations, assisted
by-sensitive experience.

In spite of the perfection attained in their nursing
of plantlice, the Lasius species are far inferior to the
Formica species in what modern animal psychology
erroneously styles intelligence, viz: in the ability to
profit for the future by past experiences. It will be
interesting, therefore, to examine, in how far the latter
ant-species, in taking care of their offspring, “con-
sciously foresee the future.”

Whenever care is taken of the young, then also the
future is instinctively taken into account, above all
in the rearing of the female ant-larvae; for it
depends entirely on modifications in the nursing,
whether the fertilized egg will produce a female
proper or a worker. But only uncritical popular

1) “Sociétés animales” (2d ed.), pp. 157, 188, etc,

184 Chapter IV.

psychology is able to confuse instinctive intention and
instinctive foresight with intelligent intention and
intelligent foresight. This is made evident by the
following facts. The beetles of the genus Atemeles
have their larvae reared by certain Formica-species,
Atemeles emarginatus by F. fusca, Atemeles paradoxus
by F. rufibarbis, Atemeles pubicollis by F. rufa, Ate-
meles pratensoides by F. pratensis. The young Ate-
meles having successfully reached their full develop-
ment, either quit the Formica nests or are driven out of
them. They then move over to Myrmica rubra’ and
spend the greater part of their lives in the nests of
these ants, by whom they are licked and fed. Only in
spring, in the mating season, they return to their
respective Formica species, where they allow their
offspring to be reared at the expense of the ant-larvae.
For whom, then, are these Formica species nursing
the young Atemeles? Not for themselves, but for
the Myrmica species. The only consequence of their
adopting the Atemeles-larvae is the immense damage
inflicted on their own eggs and larvae by these vora-
cious changelings. Where now is “the faculty of
intelligently taking the future into account,’ with
which Marshall credits his ants? For thousands of
years the Formica again and again have had the sad
experience, that the pains bestowed on these beetle-
larvae are but “love’s labor lost.” I believe that if

1) This older collective name comprises Myrmica scarbrinodis,
laevinodis, ruginodis, sulcinodis and rugulosa.

2) The same applies to the education of the larvae of the North
American Xenodusa in the nests of Formica species; for, the Xenodusa
are found as fullgrown beetles with other ants, especially of the genus
Camponotus,

Care of the Young in the Animal Kingdom. 185

Mr. Marshall had not been ignorant of the develop-
ment of these myrmecophilous beetles, he would
scarcely have extolled the great intelligence of ants.

Lomechusa strumosa spends her whole life with F.
sanguinea which, no doubt, is the “most intelligent”
of European ants. She remains with her hosts after
having attained the state of imago, and even, as a
general rule, abides in the same nest, in which she
was reared. In this case, therefore, the ants really
derive some profit from rearing the Lomechusa-larvae.
They not only have the pleasure of gratifying their
nursing instincts by the rapid growth of these adopted
children, but also later on they enjoy an agreeable,
narcotic stimulant obtained by licking the yellow
hair-tufts of the beetle. But if the ants had the
faintest trace of intelligence, would they really be
foolish enough to rear the Lomechusa-larvae merely
for the sake of this sensual gratification? They
experience again and again, that these changelings
are their worst enemies, that they destroy their brood
and moreover cause the birth of the merest cripples,
namely the pseudogyne workers. Hence ants ought
to have perceived long ago, that by rearing the Lome-
chusa-larvae they are guilty of a folly little short of
suicide. But alas, the very contrary is the case. The
longer Lomechusa-rearing has been going on in a
given sanguinea-colony—which time can be ascertained
by the increasing number of pseudogynes,—the more
care is bestowed on these beetles; and the percentage
of beetle-larvae, which, after being imbedded in the
earth, are left undisturbed, is continually increasing.
The experiences made only serve to entangle the ants

186 : Chapter IV..

more and more in the toils of their treacherous guests.
In the face of such facts “animal intelligence” is alto-
gether untenable. On the other hand, these facts
furnish a new argument proving the correctness of
our explanation of the psychic activities of animals.

Birds which nurse the unfledged cuckoos, do not
behave a whit more reasonably than the ants with
regard to their Lomechusa-larvae. Because the young
cuckoo opens its bill wider, makes more noise and
wiggles its stumpy wings more energetically, its
“foster-parents” feed it with special devotedness, and
rather suffer their own young to starve. Moreover
they calmly look on, whilst the young cuckoo pushes
their own offspring over the edge of the nest to make
them fall to the ground; indeed it has been observed,
that the foster-parents assist in this work.1 Among
birds, too, the nursing and adopting instincts are due
to the very same laws of sensitive life as in ants.
There is no discrimination between their own off-
spring and that of others, no idea of “consanguinity,”
of “parents” or “children,” but everywhere we wit-
ness the same unreasoning dependence on instinctive
sense-impressions, the appropriateness of which for
the welfare of their own or of strange species escapes
the sensitive knowledge of the animal.

This is manifest also in the care bestowed on their
young by the highest mammals, the apes. Just as
within the same species of ants eggs, larvae, and
pupae are a kind of international property, and are
therefore received and nursed also by other colonies;
as the eggs of eider ducks, of hens and other birds

2) “Westfalens Thierleben,” II, 22,

Care of the Young in the Animal Kingdom. 187

have the same international character, extending even
to the rearing of the young developed therefrom; as
in many ants and birds the instinct of adoption, which
is founded on the external resemblance between the
nurslings of strangers and their own, is now and then
extended to entirely different species (Lomechusa,
cuckoo): so there is in apes a similar instinct owing
to the same psychological causes, which proves to
evidence the lack of intelligence in animals. “It is a
well-known fact,” as the third ed. of Brehm’s Tierleben
(p. 52) has it, “that apes, without much ado, adopt
the children of any other species, protect them with
the utmost tenderness, and can scarcely be separated
from their dead bodies. When our shepherd-dog
Trina would present us again with young puppies
swarming with fleas, we used to put them into a cage
of marmosets. There they were heartily welcomed,
cleaned and fondled with care and tenderness, whilst
from without the old dog was watching with a know-
ing look (sic). But as soon as we deprived them
of their nurslings, the monkeys would set up a pitiable
screaming: they had distributed the pups among
their number and evidently intended (sic) to keep
them.” The anthropomorphism, with which modern
fanatics in the matter of animal intelligence try to
varnish over the true character of these adoption phe-
nomena, must be mercilessly exposed by genuine, crit-
ical psychology. We wish to picture the psychic life
of the animals such as it is in itself, and not as it
exists in the imagination of would-be psychologists.

That the inclination of apes to adopt the offspring
of other apes, of dogs, cats, rabbits, Guinea-pigs, and

188 C. hapter IV.

even of man is an entirely instinctive impulse devoid
of intelligent reflection, is so evident to logical minds
and so plainly expressed in the facts, that further
proof seems superfluous. Since with apes the two
sexes differ far less in psychic endowments than with
ants, it can hardly be surprising, that not only the
females but also the males have an instinctive nursing
inclination, and try to gratify it by nursing any young
animal. But how do they do it, especially if the young
ones belong to another species? Alfred Brehm says,*
“Here the ape often appears to be an inexplicable
puzzle. He nurses his adopted favorite to the full
extent of his power, hugs him, cleans him, continually
keeps an eye on him, but generally does not supply
him with any food. Without pangs of conscience
(sic), he keeps for himself the food destined for his
nursling, and even carefully keeps him away from the
pot, whilst he himself is eating. This I have observed
with baboons, who had picked up young dogs or cats
as their foster-children.”

Is this really an “inexplicable puzzle’? Only for
those who are unwilling to understand the correct
solution, because they are blinded by their monomania
on animal intelligence. The solution of the puzzle is
as clear as day-light. The instincts both of nursing
and of eating are purely sensitive inclinations, unat-
tended by reason and reflection. The faculties of
sensitive cognition and appetite are so appropriately
disposed in animals, that with regard to their own
offspring the nursing instinct is stronger than hunger,
but only so long as the young of that species, under

1) Ibidem, p. 51.

Care of the Young in the Animal Kingdom. 189

normal circumstances, require nursing. Encaged she-
apes often dispute every bite with their offspring,
although they themselves have enough to eat; yea,
they would even allow their “darling children” to
starve, unless they were prevented by force, or unless
the young possessed the strength and agility to get
at the food in spite of the envy and greediness of their
“dear mamma.”

Critical psychology cannot but regard these phe-
nomena as evident proofs, that even the highest
animals are unreasonable beings, whose actions are
guided only by instinctive impulses. Under normal
conditions, the nursing instinct which serves to pre-
serve the species, is, in animals, with regard to their
own young, stronger than hunger, which provides for
the preservation of the individual; the higher law of
preserving the species demands so. Hence, in the
beginning the mother-ape suckles and feeds her young
with unrivalled “unselfishness,” whereas later on she
grudges it every bite; hence apes hug and nurse the
young of strangers with every sign of affection,
whilst at the same time they deny them food and
cruelly allow them to starve; hence the worker-ants
nurse with motherly tenderness the eggs not laid by
themselves, whereas for the most part they devour
the eggs, which they have laid by way of partheno-
genesis. Their natural duties being those of nurses
and not of mothers, these loving aunts then become
cannibals and monsters of. cruelty, because their
instinct of eating is subordinated to that of nursing
not by intelligence or consciousness of duty, but by
the appropriate disposition of their sensitive appetite.

190 Chapter IV.

Let us briefly sum up the results of our discus-
sion on the nursing instinct of animals. In this respect
all animals obey the same psychological laws. Every-
where the inclination of nursing and rearing the young
proves to be a sensitive instinct, entirely different from,
and even excluding, individual reflection and conscious-
ness of duty. This is the case both in the highest
mammals and in ants; for the latter even far surpass
the highest mammals by their quasi-intelligent freedom
of choice in rearing the different castes, and by an
attachment to their charges verging on heroic unsel-
fishness. With all animals the care of the young is
directed exclusively by sensitive impulses and percep-
tions, which, under normal circumstances, are suitably
regulated both for preserving their own species and
for maintaining the equilibrium between different
species. Yet this appropriate correlation is far beyond
the ken of the animal; hence, in the nursing of ani-
mals there is no question of any “consciousness of
duty.” Man alone by virtue of his intellect perceives
the relations of consanguinity and the connections
resulting therefrom; he alone has an intellectual
notion of “parents” and “children”; only with him
can there be question of the moral duties of parents
toward their children. True, also in man motherly
love is founded on a sensitive instinct; but, at the
same time, it is spiritual, because the mother knows
that she is the mother of this child, and because this
knowledge with the resultant consciousness of the
duty of attending to the welfare of the child, lasts for
life. In man the love of parents toward their chil-
dren and the care they bestow on them rises far above

‘Care of the Young in the Animal Kingdom. 191

the sphere of sensitive instinct into the province of
spirituality and morality: and because the love of a
mother is a rational love, conscious of duty, there-
fore it is the highest and noblest love existing in
nature. To ascribe such motherly love to animals,
as do modern psychologists, is nonsense, scientifically
speaking, and morally speaking it is a degradation of
human dignity.

CONCLUSION.

OT to exceed the limits of this publication, we

must refrain from pointing out other parallels
existing between the psychic life of ants and that of
the other animals. In particular, the extremely vari-
ous ways of gaining a living, prevalent in ant-com-
munities, would furnish plenty of material. But in
this essay we had to confine ourselves to a few stray
remarks on that subject (p. 38). What we have
dilated upon may suffice, however, to furnish a posi-
tive and reliable answer to the question, with which
we introduced our essay, namely: Are animals en-
dowed with instinct only, or also with intelligence?

We have already proved in a former publication
(Instinct and Intelligence in the Animal Kingdom),
_of which the present study is a confirmation, that
modern animal psychology influenced by so-called
popular psychology, has inverted and confused the
notions of sensitive cognition and of intelligence.
That which. is popularly styled animal intelligence,
in as far as it is based on real facts and not on fables
and anecdotes, is nothing but the faculty of the ani-
mals of forming complex representations of their
Sensitive experiences and of acting appropriately in
accordance with them. But this power as well as the
immediate instinctive cognition is due to the innate
laws of associations of sensitive representations and
affections; hence it belongs to the sphere of sensitive

192

Conclusion. 198

instinct, not to spiritual intelligence. By a critical,
psychological analysis we were led to define instinct
as the appropriate disposition of the sensitive powers
of cognition and appetite. Hence, any action result-
ing therefrom must be called instinctive, whether
experience is concerned in it or not. But only those
actions can be called intelligent, which presuppose the
understanding of the relations existing between the
sensitive representations, and which cannot be ex-
plained by any other supposition. Intelligence, there-
fore, exclusively signifies the power to act with deliber-
ation and self-consciousness. Only this power can be
called a spiritual faculty; the sensitive power of repre-
sentation and memory cannot be so called, notwith-
standing the efforts of modern animal psychologists.
The pretended “spiritual life’ of animals, about which
popular psychology continues to make such ado, is
based on the confusion between sensitive and spiritual
faculties.

Modern animal psychology splits up the psychic
life of animals into two different factors, styled
instinct and intelligence, between which an artificial
contrast is established. Our explanation of the psychic
life of animals is more consistent and more natural.
What is erroneously termed animal intelligence, we
have traced to the same source as_ the instinctive
actions strictly so called, namely, to the suitable
hereditary disposition of the sensitive cognition and
‘appetite, which we call “instinct;’ for, this dispo-
sition has a twofold aspect, one automatic, the other
plastic. It is automatic, inasmuch as it is determined
by ata and therefore induces the animal to per-

194 Conclusion.

form certain actions, which are independent of indi-
vidual experience and are more or less the same in
all individuals of a given species. It is plastic, inas-
much as within this limited sphere, the powers of
cognition and appetite in the animal are given more
or less play for variously modifying their activities.
The narrower the limits within which they are con-
fined, the more automatically their instinct will cause
them to act; the wider those limits, the more plastic
their instinct. Both elements, automatism and plas-
ticity, are found in different proportions -with all
animals from the highest to the lowest. In the lower
orders automatism, as a general rule, largely prevails,
whereas in the higher vertebrates plasticity is, on the
average, more predominant. Ants, too, more than
dogs and apes, are bound by hereditary laws to the
performance of certain activities. The varying
influence, which individual sensation brings to bear
upon the performance of hereditary instincts, is greater
and more variable in the latter than in the former,
and in this respect the psychic life of ants is more like
“automatism” than that of mammals. But, on the
other hand, the plasticity of the instinct is, also in
ants, often highly developed, and not rarely it is
manifested in a more quasi-intelligent form, than even
in the highest vertebrates.

In the present essay we have reviewed a number
of the most prominent phenomena of the psychic life
of animals, and everywhere we found, that, what
modern animal psychology styles animal intelligence,
is met with also in ants and in many cases, in fact, in
a higher degree than with the highest mammals. In

Conclusion. 195

the community-life of ants, which with suitable
co-operation for the welfare of the colony combines
a manifold independence of action on the part of the
single workers, in their mutual communications and
mutual services, in their wars, in their slave-making
expeditions and their confederations, in their nest
construction and in the manifold application of their
building skill to various changes of circumstances,
finally, in their breeding and nursing, embracing
various methods of education left to the choice of the
workers and manifesting, at the same time, the highest
degree of “self-sacrificing attachment” to their help-
less young ones: in all these points combined we must
rightly consider the life of ants as the climax of
development in imstinctive life throughout the animal
kingdom. As regards the perfection of the nervous
system and of the sense-organs, the higher mammals
are indeed far closer to man, than the ants; but as
regards the quasi-intelligent actuation of animal
instinct under the influence of sense-perceptions and
experiences for the various purposes of community-
life, ants no doubt approach nearer to man than even
the anthropoid apes. Indeed, neither of them pos-
sesses intelligence proper, that is to say, the power
of acting with deliberation and self-consciousness, of
inventing new means for attaining various purposes
and thus making progress in civilization. Still, the
chasm between the psychic life of animals and that
of man, is, in many respects, wider between ape and
man, than between ant and man.

Of course, the results of our study are very dif-
ferent from, and indeed altogether contrary to, the

196 Conclusion,

aprioristic postulates of modern evolutionism,? accord-
ing to which man is nothing but the highest brute,

1) We cannot enter here on the general question of the develop-
ment of instincts. Cf. for this purpose my former publications: “Die
Entstehung der Instincte nach Darwin” (“‘Stimmen aus Maria-Laach,”
XXVIII, 333), “Die Entwicklung der Instincte in der Urwelt” (ibid.
papi 481; XXIX, 248, 383); “Zur Entwickl hichte der

lischaften” (‘‘Die zusammengesetzten Nester und gemischten
Kolosien der Ameisen,” III section, 2 Chap.).

As to the explanation of the genuine guest-relationship (symphily)
by the Darwinian theory of evolution cf. “Zur Entwicklung der In-
stincte” (“Verhandlgn. der Zoolog. Botan. Gesellsch.,” Wien, 1897, 3d
issue, pp. 168-183). Of late Dr. K. Escherich has tried to solve the
contradiction, which we proved to exist between the facts of
symphily and the principles of natural selection (“Zur Anatomie und
Biologie von Paussus turcicus, Zugleich ein Beitrag zur Kenntnis der
Myrmekophilie,” in ‘‘Zoolog. Jahrbuecher” Abth. fuer Systematik, XII,
1898, 27-70). He insists that symphily is not a separate instinct
totally different from the breeding instinct of ants, but that the two are
in causal relation to each other. We formerly (see the first German ed.
of present essay, p. 107 below, and p. 108 above) pointed out the same.
Yet Escherich is wrong in believing that natural selection has been
unable to prevent the development of symphily in spite of the damage
done by it to the ants, because symphily is so closely connected with
the breeding instinct. Natural selection must counteract not only the
development of an entirely new instinct which proves injurious to the
possessor, but also the extension to injurious objects of an already
existing useful instinct; hence selection was just as little allowed to let
the breeding instinct of ants extend jits activity to Lomechusa, Atemeles,
Paussus and other noxious objects, as it was allowed to let the feeding
instinct of animals extend its activity to palatable but poisonous herbs
or to nutritious plants covered with parasites. (Cf. “‘Die psychischen
Fachigkeiten der Ameisen,” 1899, p. 124.) To this Escherich again
objected (in “‘Zool. Centralbl.,” 1899, No. 1, p. 17), that many sheep
are killed by feeding on plants covered by “cercaries” (i. ¢., the
capsulate form of undeveloped trematodes). But what would Mr.
Escherich say to the following, if within the whole species of sheep,
or within a certain race of them, there should develop a special liking
‘for feeding on plants covered by those parasites? Would not such a
phenomenon evidently contradict the theory of natural selection? But
this is exactly the case with the rearing of Lomechusas by the
sanguine slaveé-makers. Therefore Escherich’s objections but confirm
the truth of our assertion: The fact that ants by nursing their guests
rear their greatest enemies, ts equally incompatible with the principles
of natural selection and with the principles of modern animal psychology.

Conclusion, 197

and human society but a gradual evolution from that
of the higher mammals. But scientific research can-
not be hampered by such aprioristic theories; if they.
are incompatible with facts, they are to be abandoned.
It is an undeniable fact, that between the soul of man
and that of the brute there yawns a chasm, which
cannot be bridged over by any evolutionistic specula-
tion.1 Man is, as a matter of fact, the only being in
the visible universe, who is gifted with reason, with
a spiritual soul, and with morality. On account of
the essential difference between sensitive and spiritual
life, it is simply impossible, that in the course of
nature an animal should ever develop into man. True,
we can daily witness, how from instinctive sensations
children gradually arrive at spiritual reasoning; but
this development is possible only because from the
outset the soul of the child is a sensitivo-spiritual soul.
The development of its spiritual faculties must be
preceded by sensitive instincts, because these furnish
the foundation and the materials for the spiritual
faculties. The animal, however, which never mani-
fests spiritual faculties, cannot be credited with any-
thing beyond a sensitive soul, which is essentially dif-
ferent from the sensitivo-spiritual soul of man, and
which makes the animal, be it ape or ant, a being
devoid of reason, and belonging to a lower order!
Hence, so-called popular animal psychology, which
denies the essential difference between the human
spirit and the animal soul, and which appeals in favor

1) Even evolutionists like Wallace have well understood this, and
therefore they protest against applying Darwinism to the psychic part
of man. Cf. Wallace, “Darwinism.”

198 Conclusion,

of this theory to the results of biological research,
must im the first place, be branded as unscientific ;+
for it mistakes sensation for spiritual life, and
instinct for intelligence, thus being diametrically at
variance with the principles of critical psychology.
Secondly, its assertion, that the brute is gifted with
reason and consciousness of duty as well as man,
although in a different degree, is an evident falsehood,
which is given the lie by the actual biological facts.
But this popular psychology is not only unscientific
and untruthful; it is far worse. To be candid, it is
demoralizing and fraught with moral danger to the
human social order. Hence we must do more than
merely shrug our shoulders in contemptuous pity, we
must take a decided stand against it and combat it
with all our might.

By denying the existence of the essential differ-
ence between animal and human psychic faculties, this
psychology not only raises the brute to the level of
man, but degrades man to the level of the brute.
Would to God that this were done in theory only;
but, alas, the practical consequence of this false theory
is the demoralization and brutalization of man. This
is the goal aimed at by those books and pamphlets,

1) Let me once more protest, as I have already done in Chap. I of
“Instinct and Intelligence in the Animal Kingdom,” that there is no
wish on my part to identify the scientific representatives of our modern
zoological psychology with the champions of animal intelligence like
Brehm, Buechner, etc. This would be an injustice to very many
sober-minded naturalists, who condemn just as we do the humanization
of the animal. Nor do we in any manner intend to pass judgment on
the personal motives of Brehm, Buechner and other defenders of
' animal intelligence and animal morality, but we only judge of their
writings. This remark is added here expressly to avoid misunder-
standings.

Conclusion, 199

which describe the sexual impulse of the brute as
essentially the same as human conjugal love, and the
care of the young among animals as essentially iden-
tical with parental love in man. Such men as Alf.
Brehm and L. Buechner were not ashamed to come
forward as “apostles of free love” and to decry as
antiquated and ridiculous the moral bounds estab-
lished for man by reason and divine law. With them
the humanization of the brute, consciously or uncdn-
sciously, aims at degrading man so far as to make
him cast off his reasonable nature and to follow with-
out reserve the sensual inclinations, which he has in
common with the unreasonable brute. On this account
they deny the difference between sensitive and spirit-
ual faculties, between the animal soul and the human
spirit. Hence we do not consider it too harsh a judg-
ment to say: Those, who humanize the animal, not
only trifle with scientific psychology, but they also
drag into the mire the dignity of man. Every well-
meaning naturalist, therefore, ought resolutely to
oppose these unprincipled doirgs of so-called popular
psychology.

Now-a-days, there is, and rightly so, a widespread
agitation against the use of alcohol and other drugs
injurious to the nervous system, because the bodily
and spiritual welfare of humanity is endangered. But
to counteract the ravages of spiritual venoms, which
under the glittering name of modern science are spread
through all classes of society, little or nothing is done.
If the moral principles of Brehm and Buechner should
later on become the common property of humanity,
then the society of the future from the highest to the

200 Conclusion,

lowest, would resemble a herd of unreasonable ani-
mals, whose “spiritual life’ would consist in the
unbridled gratification of the meanest lusts and pas-
sions. Hence our concluding appeal: Do away with
all books, pamphlets and periodicals, whose only pur-
pose is to raise the brute to the level of man!

UNIVERSITY OF CALIFORNIA LIBRARY _
Los Angeles
This book is DUE on the last date stamped below.

“Bromedirahinoarery
Biomedical Library
JAN 2 4] 1992

JAN 24 1992 _|
RECEIVED

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