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EVOLUTION BY ATROPHY:

IN BIOLOGY AND SOCIOLOGY.

By Jean Demooe, Jean Massart, and Emile Vanderveldb.

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EVOLUTION BY ATROPHY

IN BIOLOGY AND SOCIOLOGY

JEAN DEMOOE

agrege of the fkee university
of brussels

JEAN MASSART

CHARGE DE COURS OF THE FREE
UNIVERSITY OF BRUSSELS

EMILE VANDEEVELDE

PROFESSOR AT THE INSTITUTE OF HAUTES ETUDES OF BRUSSELS

TRANSLATED BY

Mrs CHALMEES MITCHELL

U.B.C. Library

acc. 1)7 7 O

LONDON
KEGAN PAUL, TRENCH, TRUBNER & CO., Ltd.

PATERNOSTER HOUSE, CHARING CROSS ROAD
1899

PREFACE

This treatise, compiled in connection with a scheme
for research work in general sociology, elaborated
in June 1894, was presented to the Institute of
Sociology at Brussels. In drawing up the pro-
gramme of the Institute, the founder, M. Ernest
Solvay, after having mentioned the questions which
especially called for the investigation of his col-
leagues, added the following statement : —

" The Institute of Sociology will take part in the
labours of the modern school of Sociology, the object
of which is to ascertain the normal conditions under
which societies exist, and the laws which govern their
evolution. But the advances of Natural Science in
this century have not yet been sufficiently assimilated
by those Sciences most closely related to it, and it
is from such assimilation that the most important
additions to knowledge may be expected."

In stating that the results of Natural Science
have not been sufficiently assimilated by Sociology,
M. Solvay is only apparently at variance with
those who rightly protest against exaggerated and

6 PREFACE

premature comparisons between social organizations
and animal or vegetable organisms.

The existence of such exaggerations, which have
caused a reaction such as recently induced an emi-
nent American economist to declare the bankruptcy
of biological sociology, is perhaps due to the fact
that, with a few distinguished exceptions, bio-
sociological investigations have hitherto been con-
ducted either by naturalists with a limited know-
ledge of social questions, or by sociologists whose
training in biology was incomplete and superficial.

To prevent this danger, our researches in the
same subject have been made separately from the
social side, and from the biological side, and have
now been co-ordinated and combined.

This work was commenced in May 1893, with
the collaboration of our friend M. Dollo, the curator
of the Natural History Museum at Brussels. In
June 1894, however, M. Dollo's many occupations
no longer permitted of his collaboration. The zoo-
logical part was therefore completed by M. Jean
Demoor, to whom most of the facts quoted in the
first book were given by M. Dollo, whose assistance
we most gratefully acknowledge.

CONTENTS

Preface

PAGE

5

INTRODUCTION

Societies and organisms. 2. Individuals, colonies and
societies. 3. Communities and societies. 4. Distinc-
tive characters of societies of which the members are
united by social contract ....

BOOK I

UNIVERSALITY OF DEGENERATIVE EVOLUTION

Part I. Degeneration in the development of institutions
and organs .....

Cliapter I. In the evolution of organs all modification is
necessarily attended by degeneration .
§ 1. Preliminary considerations

(1) Original formation of organs in ancestral forms

(2) Development of organs in the individual .

(3) Philogenetic evolution of function

(4) Individual adaptation
Section I. Transformation of organs of animals

§ 2. Transformation of homodynamic organs in the indivi

dual (metameric appendages)

The cray-tish .....

§ 3. Transformation of homologous organs in individuals

of different species (limbs) .

Limbs adapted to an aquatic life (Ccralodus, Ortha

canthtts, Protopterus amphibius, Protopterus annec

tens, Lepidosiren ....

Limbs adapted to a terrestrial life

21

22

22
23
23
25
26
30

30
31

41

43
43

CONTENTS

1. Adaptation to walking on two legs (man,
birds). 2. Adaptation to leaping (Dipiis a>gyp-
tius, kangaroo, Tardus spectrum, Rana esculenta.
3. Adaptation to running (horse, ruminants). 4.
Adaptation to flight (birds, Pterosaurians, bats).
5. Adaptation to arboreal life (Arctocelus cala-
barensis, Chanueleo). 6. Adaptation to swimming
(Cetaceans, Sirenia). 7. Adaptation to burrowing
(Talpa curopcea, Heterocephalus) ... 46

Section II. Modification of the organs of plants . 68

§ 4. Modification of homodynamic organs in the indivi-
dual (basilar and apical leaves) ... 68
1. Rosa rugosa. 2. Serratula centaur oides. 3.
Sagittaria sagittifolia. 4. Lathyra Aphaca. 5.
Nymphaea dentata ..... 70

§ 5. Modification of organs which are homologous in

individuals of different species (foliage leaves) . 78

1. Adaptation to climbing (Cobaeca scandens, Vic la
pyrenaica, Cucumis satiinis). 2. Adaptation to
carnivorous nutrition (Utncularia, Nepenthes,
Drosera). 3. Adaptation to an aquatic life (Sagit-
taria, Nymphaea, Vallisneria, Potamogeton, Ranun-
culus, Ouvirandra fenestralis). 4. Adaptation to
defence against ants (Acacia sphaerocephala). 5.
Adaptation to drought (Semper cicum). 6. Adapta-
tion to defence against herbivorous animals
(Caragana, Ilex, Mamillaria, etc.) . . 78

Chapter II. In the evolution of institutions all modification

is necessarily accompanied by degeneration . . 90

§ 1. Modifications of similar institutions in the same

society ...... 91

(1) The communal budgets of Belgium . . 92

(2) Budget of the States of the German Empire . 95

(3) The budgets of Germany, France, and England . 97

§ 2. Modification of similar institutions in different social

groups (the development of landed property) . 98

1. Family property (Montenegro). 2. Village pro-
perty (Russia). 3. Feudal property (England).

CONTENTS

4. Public or collective property (Switzerland).

5. Corporative property (Belgium). 6. Private
property (Switzerland). 7. Summary . . 100

Paut II. Degeneration in the evolution of organisms and

societies ....... 115

Chapter I. All organisms exhibit rudimentary organs . 115

Section I. Rudimentary organs of animals . . 117

§ 1. Rudimentary organs in man .... 117

1. Integumentary system. 2. Skeleton. 3. Mus-
cular system. 4. Nervous system. 5. Digestive
system. 6. Vascular system. 7. Sense organs.
8. Genito-urinary system . . . 117

§ 2. Rudimentary organs in various groups . . 121

1. Ccelenterates. 2. Worms. 3. Echinoderms. 4.

Mollusca. 5. Arthropodes. 6. Vertebrates . 121

Section II. Rudimentary organs in plants . . 145

§ 3. Rudimentary organs in various groups of plants . 145
1. Algae. 2. Mushrooms. 3. Bryophyta. 4. Pteri-

dophyta. 5. Phanerogams . . . . 145

§ 4. Reduced organs in the vegetative apparatus of the

Phanerogams . . . . . . 149

Chapter II. Survivals exist in all kinds of societies . . 151

§ 1 . Instances of survival in various groups . . 153

(1) Instances of survival in the most modern social
groups (the United States) .... 155

(2) Instances of survival in less civilized social groups
(Veddahs, Fuegoes, Australian tribes . . 156

§ 2. Survivals of ancient forms of marriage, and of the

family in Modern Europe .... 161

1. Forms of marriage. — (1) Marriage by capture. (2)
Marriage by purchase. (3) Marriage by consent
of both parties (marriage by simple consent, mar-
riage in facie Ecclesia) .... 161

2. The Family System.— (I) Matriarchy. (2) Patri-
archy ....... 167

Part III. Summary and conclusions .... 170

CONTENTS

- PAGE

BOOK II

THE PATH OF DEGENERATIVE EVOLUTION

Part I. The supposed law that degeneration retraces the

steps of progress ...... 175

Chapter I. The path of degeneration in biology . . 178

Section I. The path of degeneration in animals . 179

1. Morphology and embryology ; the law of recapitu-
lation. 2. Degeneration of the third eye in lizards.
3. Degeneration of the organs of sight in deep-sea
Crustacea. 4. Atrophy of the branchial vessels in
man ....... 179

Section II. The path of degeneration in plants. . 192

1. Rarity of cases of recapitulation in the organ-
ogeny of leaves (Vicia, Acacia with phyllodes).
2. Organogeny of flowers (Brassica oleracea var.
Botrytig). 3. Progressive degeneration of the pro-
thallus in phanerogams .... 192

Chapter II. The path of degeneration in sociology . . 205

§ 1. Investigation of facts ..... 205

1. Tithings, hundreds and counties in England. 2.
Order of elimination of various racial elements in
a country. 3. The degenerative evolution of poli-
tical organizations. 4. Degeneration in monetary
systems. 5. Degenerative adaptation in colonial
legislation. 6. Degenerative evolution of the cor-
porations of Western Flanders
§ 2. A criticism of the supposed inverse path of degenera
tion ......

Part II. The irreversibility of degenerative evolution.
Chapter I. Do institutions or organs which have disappeared
reappear ? .
Section I. Disappeared organs .

1. Plants .....

2. Animals. Teratology of the horse, Hypertrichosis,
etc. Swimming limbs in Stomatopoda and De-
capoda Macroura ..... 223

Section II. Disappeared institutions . . . 227

207

217

221

222
222
222

CONTENTS

PAQK

(1) Apparent revival of bygone institutions . . 227

(2) Apparent disappearance of institutions . . 229

(3) Instances of convergence .... 230
Chapter II. Can rudimentary institutions or organs reassume

their primitive functions ? 232

Section 1. Rudimentary organs . . . 232

1. Animals. (1) Muscles of the ear in man. (2) The
abdomen and appendages in deep-sea hermit crabs 232

2. Plants. (1) Hermaphrodite flowers in Melandryum.
(2) Branches of Colletia cruciata, Crataegus, Yicia

Faha, Sempervivum, Veronica, etc. . . . 233

Section II. Rudimentary institutions . . . 237

(1) Truck system and clearing-house. (2) Corpora-
tions and syndicates. (3) Archaic collectivism and
modern collectivism. (4) The survival of elective
sovereignty in England . . . . 239

Chapter III. Can rudimentary organs or institutions re-
develop and assume new functions ? . . . 242
Section I. Rudimentary organs . . . 243

1. Animals. Respiratory organs in Birgus latro ;
Mesonephric spaces in the higher vertebrates . 243

2. Plants. Staminodes of Pentstemon . . 244
Section II. Rudimentary institutions . . . 245

Levirat ....... 245

Part III. Summary and conclusions . . . 247

BOOK III

CAUSES OF DEGENERATIVE EVOLUTION

Part I. Atrophy Qf organs and institutions . . . 251

Sect-ion I. The factors of atrophy .... 251

(1) Biology (accidental, individual, normal and specific
atrophy). (2) Sociology (accidental and normal

atrophy) ...... 251

Section II. Causes producing atrophy . . . 260

Chapter I. Atrophy of organs ..... 260

§ 1. Atrophy from lack of space .... 261

CONTENTS

(1) Development of the teeth. (2) Atrophy of the
superior glume (Loh'um). (3) Degeneration of
Paleae (composite flowers) and of stamens
(Scrophulariaceae) .... 261

§ 2. Atrophy from lack of use .... 263

1. Functional inutility : (1) Etiolated plants and

immobile limbs. (2) Epicotyl and primary leaf
of Nymphaea. (3) Roots of Utricularia ; coty-
ledons of parasitic plants (Ctiscuta, Orobanche);
leaves transformed to spines in Phyllocactus
crenatus. (4) Atrojjhy of the branchial arches in
mammals. (5) Atrophy of ventral fins ( Pediculati,
Protopttrus). (6) Atrophy of muscles (Cetacea,
Sirenia). (7) Atrophy of the tail in man. (8)
Degeneration of the hyo'id apparatus in man
and birds. ..... 263

2. Transference of function : (1) Atrophy of the tail

in Batrachia anura, and the larval gills in some
insects. (2) Disappearance of limbs (Slow-
worms, Amphisbtena, Snakes, Eels, and Saccu-
lina). (3) Atrophy of the leaf {acacia with phyl-
lodes, A' 'ylophyl 'la). (4) Atrophy of theprotonema
in mosses, and of the leaves in some xerophilus
plants (Muehlenheckia platyvlados, Genista,
Spartium, Alhagi). (5) Disappearance of the
calyx. (6) Atrophy of roots (Pine, Beech,
Corallorhyza, Myrmechis, Tillandsia zisneoides),
or of leaves and stems (Tceniophyllum Zollin-
ger!, Podostemacece) .... 268
§ 3. Atrophy from lack of nutrition . . . 274
1. Parasitic castration (Melandryum album). 2.
Severe or prolonged compression of a limb.

3. Atrophy of the genitalia in neuter bees.

4. Atrophy of the superior flowers in Carex.

5. Atrophy of pistils and stamens (Frltillaria
persica, Viburnam tomentosum, Viburnum Opulm) 274

§ 4. Atrophy without apparent cause . . . 278

Atrophy of perianth (.4 rtemis ia, Poterium). Atrophy
of the'eyes (Myriopodes, Cymothoe). Correlative
atrophy ...... 278

CONTENTS

PAOE

Chapter II. Atrophy of institutions .... 281

§ 1. Atrophy from lack of use .... 282

1. Functional inutility: (1) Offices of the port of

Bruges. (2) The forest-courts of England . 282

2. Transference of function : (1) Republican institu-

tions under the Roman Emjtire. (2) Special
jurisdiction in England .... 284

§ 2. Atrophy from lack of resource . . . 287

1. Local administration at the close of the Roman

Empire. 2. Degeneration of societies in general 287

Part II. Causes of the persistence of organs or institutions

without function . . . . . . • 292

Chapter I. Survival of organs ..... 292

§ 1. Unfunctional organs that are not rudimentary . 292

Absence of variation (flowers of Ficaria ranuncu-
loldes, Lysimachia Nummularia, Elodea canadensis,
stratiotes aloides, cleistogamous flowers, eyes in
the male Machaerites .... 292

§ 2. Unfunctional organs which persist as rudiments . 295

1. Absence of variation. Insignificance of the rudi-
mentary organ (stipules of Tropaeolum majus,
accessory rudiments of enamel organs in the
development of teeth .... 295

Chapter II. Survival of institutions .... 298

§ 1. Integral persistence of an institution . . . 299

(1) Maintenance by compulsion (rotten boroughs in
England ; the States of Normandy and the
Dauphiny in France). (2) Indirect usefulness
(English monarchy). (3) Respect for tradition
(royal prerogatives exercised by the praetor ;
institution of sheriffs in England) . . 299

§ 2. Survival of institutions in a reduced condition . 306

1. Insignificance of the institution (jurisdiction at
da Martinique ; summons <juo warranto ; Diocese
of Cambrai ; marsh-land in Artois ; tribute paid
to Spain by France). Respect for traditions
(survival of the old regime in England ; instances
of survival in law and religion . . . 307

Part III. Summary and conclusions .... 317

General conclusions ...... 320

INTRODUCTION

Ever since the application of theories of evolution
to social phenomena, there has been a constant inter-
change in terminology between biology and sociology ;
societies have been called organisms, and organisms
societies of cells. There is an actual division of
labour among the organs of a living body, while
institutions have been called the organs, or parts
of organs, of Society. The interchange of matter
effected among the organs of an individual has been
called a " physiological contract," while the circula-
tion of money may be compared to the circulation
of blood and lymph. Questions arise as to what
extent such comparisons are legitimate, if they
should be taken in any other than a metaphorical
sense, and if it is possible to set a precise boundary
between the provinces of biology and sociology.

Much has already been written on such problems
as these, and no doubt much more will yet be
written. We shall not attempt either to discuss
or to solve them in these few introductory pages ;
they are merely touched upon here, and will only
be alluded to when absolutely necessary for the
careful investigation of facts bearing upon our work.

I. Societies and organisms. — The analogy exist-

7

8 EVOLUTION BY ATKOPHY

ing, from the point of view of evolution, between
biology and sociology, arises from the fact that the
evolution of societies as well as that of organisms,
is the result of the co-operation of two factors —
similarity and adaptation.

In biology, the similarity between organisms
springing from the same stock, is due to heredity,
while adaptation is the result of individual variation.

In sociology, societies are the descendants of
former societies, in that the new are modelled
upon the old. Similarity is the result of imita-
tion, while adaptation is the result of invention
— i.e. of all improvement and innovation tending
to make a new society different from that which
preceded it.1

These fundamental analogies suffice in themselves
to justify our collaboration, whatever may be the
solution of the question — which is really only a
question of terms — as to whether societies should
be regarded as organisms, or organisms as particular
kinds of societies.2

1 Invention is frequently a combination of several imitations.
When a society is formed, its characters are not necessarily
borrowed from those societies from which it more or less directly
proceeds. It may be modelled upon other social structures with
which it had no hereditary connection. — V. Tarde, Les Lois dc
V imitation.

2 See Les SociiUs animates, p. 128 (Espinas). "Integration,
or grouping together, is a universal law common to all organic
or inorganic existence. Society, properly speaking, is only a
complex and important instance of this universal law."

See La Science Sociale, p. 97 (Fouillee). " All purely physio-

INTRODUCTION 9

In either case, it is certain that organisms and
societies — used in the sociological acceptation of
the term — exhibit some characters in common, and
some distinctive characters.

The common characters may be accounted for
by the co-operation which exists in both cases
between the units of which they are composed
(individuals, or cells).

The dissimilar characters are probably connected
with what constitutes the essential difference be-
tween social aggregates and organic aggregates.
With the former there is a physiological continuity
between the composing units, while with the latter
co-operation is entirely due to mental relations.

II. Individuals, colonies and societies. — Our view
in this matter is considerably at variance with the
current opinion. Many authors, and M. Espinas
among them, regard colonies, whether animal or vege-
table, as societies, even when the members of these
colonies are connected by physiological bonds.1

logical characters of life, viz.: 1. Correlation of parts ; 2. Relation
between structure and function ; 3. Division of all living parts
into other living parts ; 4. Spontaneity of movement ; 5. Partic-
ulate existence ; 6. Development and degeneration, i.e. evolution,
are to be found in a greater extent in animal and human societies."
1 See Lcs Soci&is animalcs, section 2, pp. 207 and foil. —
(Espinas), Cours cle philosophic positive, vol. iv. (A Comte).
• ' These strange societies are to be found among the lower
animals, an involuntary co-operation being the result of an un-
severable organic union, which is either a mere adherence or
actual continuity."

10 EVOLUTION BY ATROPHY

We shall justify the distinction we make by
showing that colonies are entirely distinct from
societies. The two are divergent branches which
spring from the same source, a solitary indi-
vidual. These divergent branches of organic life
may be distinguished in the following manner :
An individual may be either unicellular or mul-
ticellular. Every cell capable of sustaining life
and reproducing its own kind is an individual.1

In the case of multicellular individuals there is
an unbroken physiological continuity, while life
lasts, between the cells of which they are com-
posed. All these units spring from one primary
cell (i.e. a fertilized egg) which sprang itself from
two cells (male and female), proceeding usually
from two separate individuals. From the moment
that this union is effected, the continuity remains
unbroken.2 When such an individual reproduces

1 We regard the cell as the unit of life, although, according to
Altmann, the unit of life is a still simpler structure — the bioblast.
He maintains that the bioblast, or living granule, is the ultimate
element of the cell ; it is born from a pre-existing granule ; it
lives, feeds, reproduces, and is, in fact, an organism (see Die
Elcmcntarorganismen und ihre Beziehungen zu den Zellen ;
Leipzig, 1890. — Studien ilber die Zellen; Leipzig, 1886. — Zur
Geschichte der Zelltheorien ; Leipzig, 1889 (Altmann). The
existence, however, of the bioblast is a pure hypothesis, no
definite proof having been established of its existence ; and to
assume that it is the simplest unit of life is to abandon oneself
to pure imagination.

2 Although this theory is in agreement with the observed facts
of biology, it is necessary to point out the complexity of the
ideas of individual and organism. Boveri (Uber die Befruchtvngs

INTRODUCTION 1 1

asexually, giving rise to offspring which remain
attached to it, the whole forms a colony. All
the cells of this colony spring then from one
single cell (a fertilized egg cell), and an inter-
change of nutriment goes on incessantly among
these cells. This is what Spencer designates as
a " physiological contract." Societies, on the other
hand, consist of units springing from various sources,
whose connection is merely mental. It is only in
the communities which Tonnies calls " Gemein-
schaften," in distinction from societies bound by
mental agreement (Gesellschaften), that all the in-
dividuals spring from one and the same couple.

From our point of view, a colony must not be
regarded as an intermediate condition between an
individual and a society. No known society has
passed through a colonial stage, and the members of
a colony could not, on separating, become a society.
Among the simplest aggregates, all the units are

und Entwicklungsftihigkeit kernloser Seeigel-Eier, und fiber the
Moglichkeit ihrer Bastardirung. Arch. f. Entioicklungsmechanik
dcr Organismen, Bd. ii., pp. 394-443), for instance, has shown
that it is possible to fertilize the eggs of Echinoderms, from
which the nuclei have been removed, by the spermatozoa of
other species of Echinoderms. The egg is a typical cell, an
organism, an individual. All its parts are essential to it, and
are incapable of separate existence, at any rate far any length
of time ; close physiological bonds unite the component parts,
yet it is possible to substitute for an essential part of one
individual a part taken from another individual. It may there-
fore be concluded that the idea of an organism no longer necessarily
implies the idea of continuous functional unity which one was
formerly tempted to ascribe to it.

12 EVOLUTION BY ATROPHY

equivalent, and each retains all the functions of
life {Spirogyra, Choanoflagellates, Hydra). The
colony may disintegrate, but the cells or indi-
viduals set at liberty are capable of living alone,
and do not form a society.1

As the organisation of a colony becomes more
complicated, the units proceed to differentiate. Each
one assumes certain functions in particular, and
becomes less adapted to discharge others. If such
a colony is dispersed, the individuals are incapable
of maintaining a separate existence.2'

1 (a) A thread of Spirogyra consists of cells placed end to end,
physiologically connected with one another. Under certain
conditions, however, quite apart from the phenomenon of cellular
reproduction, all these cells are capable of isolating themselves.
The colony is thus transformed into a set of individuals capable
of maintaining a separate existence, and no longer in connection
with one another.

(b) Among the Choanoflagellates, some consist of free individuals,
while in others all the individuals are united by a common stalk,
and intercommunicate by protoplasmic threads in the stalk. When,
for some cause, these individuals separate, they never form them-
selves into a society.

(c) A colony of Hydra is formed by the budding of a single indi-
vidual. While nourishment is abundant, all the individuals of the
colony retain their connection with each other, and may themselves
give rise to buds. On the other hand, when food is scarce, the
colony disintegrates and each individual lives a free life without
entering into social relations with its neighbours.

2 In some cases the terms individual and colony become ex-
tremely involved. Although one accept our view, it is difficult to
rigorously apply our generalizations to such facts as the following :
Certain male cephalopods, at the period of reproduction, separate
from their bodies a specialized tentacle (hectocotyle) in which is
stored the seminal fluid. This organ, set at liberty, swims in the

INTRODUCTION 1 3

III. Communities and societies. — The bond ex-
isting between all the constituent parts of a society-
is not of the same nature as that which unites the
members of a colony or the cells of an animal or
plant. These physiological bonds are not eompar-

sea, and after living there a certain time, enters the female oepha-
lopod, to effect impregnation.

The individuality of this hectocotyle appears so obvious that
for a long time it was regarded as a distinct species— some kind
of worm. At first sight, then, the hectocotyle seems to be an
individual. This, however, is not the case, as it reproduces, not
a hectocotyle but a cephalopod.

Many Echinoderms, by a spontaneous act of protection, can
separate their arms from their bodies. Such a separated arm
may live, feed and slowly build up again a whole Echinoderm.
Plainly, it would be impossible to indicate the precise point at
which such an organ should be regarded as an individual. The
vegetable kingdom abounds in analogous facts. A strawberry
plant, for instance, gives off runners in the course of the Summer,
which take root, and themselves become strawberry plants. So
long as these young plants are insufficiently developed to main-
tain themselves, the mother-plant continues to supply them with
nourishment. As soon, however, as the young shoot can dispense
with this support, the runner atrophies, and the little plant begins
a separate existence. By prematurely cutting the runner, the
new plant may be compelled to live alone sooner than it naturally
would have done.

With some other plants (Phalangium viviparum) the young
shoots frequently retain their connection with the mother-plant,
although quite capable of maintaining themselves. Under these
circumstances it is obviously impossible to say if these plants
represent colonies or free individuals. Speaking generally, it
may be said that no precise line can be drawn between colonies
and individuals. Many writers on the subject, and Perrier among
the number, consider that every colony where there is a physical
continuity among the members, should be regarded as an in-
dividual.

14 EVOLUTION BY ATROPHY

able, without forcing the analogy, although they have
been so compared by some sociologists, to such means
of communication between individuals and societies
as exchange, traffic, roads, railways, telegrams and
telephones. It is merely a matter of definition, and
if societies are to be termed organisms, they should
be distinguished as organisms by social contract,
organismes contractuels (Fouill^e).

This definition, however, only applies to those
societies which owe their existence to a formal
contract with definite objects in view, and not to
ready-made communities consisting of individuals
already united together without any preliminary
contract. The latter is the case, for instance, in
societies of ants or bees, and in human societies in
those social groups in which the individuals are
united by the bonds of consanguinity. The char-
acters of such communities partly approach the
characters of organic associations, but precisely as
such natural communities approach societies by
social contract, the differences between social groups
and actual organisms become more marked. In the
more complex forms of societies the results of the
characters we have distinguished become most ac-
centuated.

IV. Distinctive characters of societies of which
the members are united by social contract. — (1) A
cell cannot be part of two organisms or of two

INTRODUCTION 1 5

organs at the same time.1 On the other hand,
there is no reason why the members of one social
community should not belong to other communities
at the same time.

2. Speaking generally, the biological conception
of an organism denotes a definite thing — a plant or
an animal in itself, quite distinct from similar
organisms. In sociology, however, there is no
precise line between co-existing social communities.
Are we to regard, for instance, the families, com-
munes and cantons of a state as distinct organisms,
or merely as organs ? Does a free town such as
Hamburg or Frankfort cease to be an organism
when it loses its independence ? Take the various
Swiss cantons, which are now mere organs of
the Helvetic Confederation, like the Provinces
of Belgium, or the Departments of France, would
they become organisms on the rupture of the
Federal bond ? On the other hand, with the
growth of international treaties between the
states of Eastern Europe, will their social indi-
viduality disappear, and will they come to be
regarded as are the United States of America,
as the organs of an organism in process of
formation ? These few examples suffice to show

1 When two organs are united into one whole, the cell exercises
two totally different functions. The liver, as we now know, is a
double organ consisting of a bile-forming liver, and the glycogen -
producing liver, two organs which are embryologically distinct.
The cells of which the liver is composed are both bile-secreting
and glycogen-forming organs.

1 6 EVOLUTION BY ATROPHY

that, so far as social matters are concerned, the
conception of organisms is a pure convention.
In the course of this treatise, we may therefore
regard families, societies and nations as distinct
organisms, or, with regard to their connection with
other and vaster organizations, as organs of the
latter.

3. The structure of an animal or plant depends
upon the physical arrangements of its parts, and on
the physiological links between those parts. The
structure of a society depends upon the links of
social contract existing among its members. We
regard these as two very different things, and we
cannot follow Tarde in pressing the analogy between
them in the following way : " The length, breadth,
and height of an organism are never very much out
of proportion. With snakes and poplars the height
or length preponderates ; among flat fish the thick-
ness is very small compared to the other dimensions,
but in each instance the disproportion exhibited in
extreme cases is not comparable to that shown by
any social aggregate — such as China for instance,
which is 3000 kilometres in length and breadth,
and only one or two yards in average height, for
the Chinese being a short race, build their edifices
correspondingly low." 1

4. Organic modifications are effected more slowly
and with greater difficulty than are social modifica-

1 Les Monades et la Science sociale. (Tarde. Revue de Sociologie,
1893, p. 169.)

INTRODUCTION 1 7

tions.1 The result is an important one from the
point of view of method.

In biology, excepting in the case of individual
adaptation of artificial selection, direct observation
— the historical method, if we may so call it — is
not available for the study of the origin and
modification of organisms. Phylogeny, the science
of organic kinship, resorts to other methods, and
particularly to the comparative method in its
various forms : —

(a) Morphology, the science of determining the
phylogeny of organs by comparing them with the
organs of other creatures belonging to the same
systematic group.

(b) Palaeontology, which determines the direct
ancestors of living creatures.

(c) Embryology, which, so far as it is founded
upon the principle of recapitulation, investigates
the development of organs in the individual, and
draws conclusions therefrom bearing upon its
descent.

1 Among animals there is a special factor which gives a stability
to the specific characters not found elsewhere — this is reproduction.
Specific characters being common to the whole line of descent, arc
very deeply enrooted in the organism. They are not easily modi-
fied by the influence of new environments, but maintain their
likeness to one another in spite of external conditions. They are
regulated by an internal force, notwithstanding the importunities
to variation offered from outside. This force is heredity, and
heredity accounts for the precise way in which specific characters
may be defined. In society this internal force is wanting. (Lcs
/,'< git ••udelamtthode socioloyique. Durckheini, Paris, F. Alcan, 1895.)

B

18 EVOLUTION BY ATROPHY

(d) Teratology, which compares normal with
abnormal forms for the same purpose.

In sociology these various systems of comparison
are only of a secondary value, owing to the great
variability of social forms, while the historical
method of investigation assumes a greater impor-
tance. In sociology, however, there are methods
analogous to the comparative methods of biology :

(a) Archeology corresponds to palaeontology.

(b) Social morphology, by comparison of series
of institutions, makes up to a certain extent for
the absence of direct observation of their origin
and development. Thus, to use the phrase of
Bagehot, by studying the customs and institu-
tions of modern savages, the prehistoric living
may be made to throw light upon the pre-
historic dead. If an institution be found in full
operation among savages, of which a vestige still
exists among more civilized people, it may be
assumed that the vestige was at some time fully
functional among the latter. It must be borne
in mind, however, that in many instances such
vesticres are the result of imitation. Theodore
Eeinach has shown that this applies in the case
of circumcision.1

This gravely weakens the conclusions drawn by
Spencer from the survival of this custom among

1 Dc quclqucs fails relatifs A Vhistoire dc la circoncision chcz les
2>cuplcs dc la Syrie (Th. Reinach, L 'Anthropologic, 1893, vol. iv.,
pp. 28 and following).

INTRODUCTION 1 9

certain Australian tribes. Granting, he says, that
circumcision, the removal of teeth, and other
similar mutilations imply a condition of political
or religious subjection — or both — no longer exist-
ing among these tribes, the custom is obviously the
vestige of a more complex social condition. This
conclusion seems the less reliable, since, according
to the Eev. J. Matthew, the rite of circumcision
was probably introduced into Australia by natives
of Sumatra, and this view is confirmed by the
local distribution of the custom and by other evi-
dences of the same origin, such as the paintings
which have been discovered in certain caves.1

(c) Teratology and social embryology also play a
part in sociology, but it is of less importance than
the others.

Certain customs among criminals show a re-
semblance to the habits of primitive man. On
the other hand, we find cases where the individual
development of an institution or society is a mere
repetition of the development through which similar
institutions and societies of other epochs and places
have passed. Thus, for instance, there still exists
in some parts of modern Russia a voluntary agri-
cultural commune, for the periodical division of the
land, an institution which existed more universally
in the sixteenth and seventeenth centuries during

1 The Cave Paintings of Australia (Journal of the Anthropo-
logical Institute of Great Britain anal Ireland, April 1S93, pp. 51
and following).

20 EVOLUTION BY ATROPHY

the existence and after the disappearance of family
communities.1

In studying the development of these new com-
munities, one can, in a measure, picture to oneself
the development of laws of property, which were
current in other countries at other times, and of
which we possess little or no direct information.
It is, however, hardly necessary to insist upon the
hypothetical quality of such conclusions.

Our methods show, then, that organisms and
societies exhibit considerable differences as well
as analogies, a necessary result of their different
natures. These few remarks must suffice ; to add
to them, we should have to overstep the limits we
have set to this treatise, enter into well-worn con-
troversies, and anticipate our own conclusions.
Having merely explained our terminology, and
indicated our general views, we will proceed to
the subject of our investigations.

1 Tableau dcs original ct de Involution dc la proprUU ct de la
fiimUlc, p. 170 (Kowalevsky).

BOOK I

UNIVERSALITY OF DEGENERATIVE

EVOLUTION

PART I

DEGENERATION IN THE DEVELOPMENT OF INSTITUTIONS
AND OEGANS

The term " Evolution " does not in itself convey
an idea of either progress or degeneration. It
comprises all the changes undergone by an
organism or society independently of the ques-
tion as to whether these changes are favourable
or otherwise. The evolution of an organ, or of
the different parts of an organ, is degenerative if
it tends to the ultimate decay of that organ or of
its parts, and the facts are shewn by means of
arranging series of fossils or living forms, and com-
paring them. Evolution is progressive if it tends
to the development of an organ or to the formation
of a new organ.

These definitions may be applied — mutatis
mutandis — to the changes undergone by societies
and institutions or their constituent parts.

The ideas of progress and of degeneration seem
at first sight to relate to diametrically opposite
phenomena. The term " progressive evolution "

22 UNIVERSALITY OF DEGENERATIVE EVOLUTION

conveys the ideas of progress, development, im-
provement, of increasing differentiation, and of the
progressive co-ordination of the functions or organs
thus differentiated.

Degenerative evolution, on the other hand, con-
veys the ideas of decline, of decay, and of de-
generation, such as the atrophy of the organs of
locomotion in Sacculina, the degeneration of the
leaves of parasitic plants, or the dissolution of
corporate bodies in a declining state.

We propose to show, however, that these two
ideas, which at first sight seem contradictory and
mutually exclusive, are found, on a strict examina-
tion of the facts, to interpenetrate and complete
each other. Degeneration and progress will appear
as the two sides of one whole, or as two aspects of
the same evolution, and it will be seen that all
progress must necessarily be attended by degenera-
tion.

CHAPTER I

IN THE EVOLUTION OF ORGANS ALL MODIFICATION IS
NECESSARILY ATTENDED BY DEGENERATION

Section I. — Preliminary Considerations

In order to effect the demonstration which is the
object of this chapter, only the phylogenetic modifi-
cations of organs will be discussed, setting aside

EVOLUTION AND DEGENERATION OF ORGANS 23

the original formation of these organs in their
ancestral forms, their development in the indivi-
dual, the phylogenetic evolution of function, and the
great variation resulting from individual adaptation.
Before entering upon our immediate subject, it
would be as well to define its limits by saying
a few words concerning the questions not under
immediate consideration.

1. Original formation of organs in ancestral
forms. — Little is known concerning the primary
origin of organs, and their development before their
assumption of the aspect and function by which
we know them. What were the rudimentary
leaves like in the ancestors of flowering plants ?
What were the eyes of the first vertebrates like ?
Did these organs develop from existing organs ful-
filling other functions, or were they formed inde-
pendently ? However that may be, if they arose
from other organs, we know nothing of the modifi-
cations which they underwent ; and if they were
formed independently, we need not discuss the
fact here, for in that case the organ would not
have developed by transformation. Having once
been formed, it would develop and improve, and
this process would not necessarily have been
attended by partial degeneration.

2. Development of organs in the individual. — In
the course of embryological development, organs
do not exactly repeat the successive phases

24 UNIVERSALITY OF DEGENERATIVE EVOLUTION

through which they passed during their ancestral
evolution. Generally speaking, and especially in
the case of plants, the development of organs in
an individual is direct, and gives no clue to its
ancestral history. Moreover, when there is a re-
capitulation of ancestral stages, it often happens
that evolution takes place without leaving traces

of the various
stages. This is
especially the
case in complex
organs which
have been pro-
duced by many
, lines of evolu-
C\- tion converging
* in a single struc-
ture — a struc-
ture which thus
becomes the seat

Fig. 1. — Diagram showing the evolution of pyramidal
cells in the animal series.

The upper series of cells represents the psychic cell in
various vertebrates: A, the frog; B, the lizard;
C, the rat ; D, man. The lower series shows the
progressive stages in the evolution of the pyramidal
cell in the human brain : a, the neuroblast without
protoplasmic processes ; b, the appearance of the
nerve process and of the terminal ramifications ;
c, the nerve more fully developed; d, appearance of
lateral branches of the axis cylinder; e, development
of protoplasmic outgrowths of the protoplasm of
nerve-cell and nerve. (Ramon y Cajal.)

the human brain, passes successively through stages
corresponding to those which are to be found in the
adult fish, frog, bird, and mammal. In this case

of a special func-
tion or set of
functions.

The neuron,
for instance, the
ganglionic cell
of the cortex of

EVOLUTION AND DEGENERATION OF ORGANS 25

the development consists in an increasing com-
plexity of the cell with no formation of unnecessary
rudimentary parts.

3. Phylogcnetic evolution of function. — Evolution
may be regarded from a physiological, as well as
from an anatomical standpoint,1 but, in the former
case, evolution is less a set of changes of function
than an increasing specialization and division of
labour, and under these circumstances it is often diffi-
cult to recognize a degenerative element in the evolu-
tion. A few examples will demonstrate this point :

Self-mutilation is a very common phenomenon
among Echinoderms. Among brittle star-fish this
reaction is controlled by some region of the nervous
system ; in some star-fish the reaction follows more
quickly because the stimulus can act upon the
ganglion at the root of each arm near the circum-
oral nerve-collar. In Asteracanthum rubens, there
is a complete localization of this function, and self-
mutilization only results when an exact region of
the nervous system is stimulated.2 In the Medusa
we find an equally interesting example of functional
evolution (Romanes).3 With some of these (the

1 Evolution fonctionnclle du systeme nerveux. J. Demoor, Revue
iniirrrsitairc, Bruxelles, 1892.

2 Contribution a hi physiologic ncrveuse cles JSchinodermes. J.
Demoor and M. Chapeaux. Tijds. Ned. Dierlc. Vcreen. (2) III.
2 Nov. 1891.

8 Preliminary Observations <>n //<>■ Locomotor System of Medusa ;
Jelly-fish, Star-fish, and Sea-urchins. Romanes, Int. Seient. Series,
1885.

26 UNIVERSALITY OF DEGENERATIVE EVOLUTION

Acraspedote, on the outside of the umbrella being
separated from the central part, the two separate
parts continue to lash the water, the outer part
with even strokes, the central mass more slowly
and feebly. With the Craspedote, on the other
hand, the central part, under the same conditions
seems quite paralyzed and immovable, while the
outer part continues to move in a perfectly normal
manner. The causative function of the movement,
the spontaneity of the movement as it was formerly
called in physiology, is incompletely specialized in
the Acraspedote, whereas in the Craspedote it is
entirely localized.

Individual adaptation. — The individual is by
no means a slave to heredity. It is capable of
certain modifications under the influence of certain
external conditions. These phenomena of individual
adaptation may be arranged in three groups.

(a) When an organism, either animal or vegetable,
is placed under new conditions of existence, when
for instance, it relinquishes a terrestrial for an
aquatic life, light for darkness, or fresh water for
salt or esturine water, its external aspect, and in-
ternal structure, undergo variations of considerable
importance if it succeeds in adapting itself to the
new conditions.1

1 Examples : (a) The leaves of the water Ranunculus with
lacinated leaves (Ranunculus aqua til is fluitans, etc.), are of normal
structure when cultivated on dry land. The epidermis is furnished
with stomata and the constituent cells contain no chlorophyll.

EVOLUTION AND DEGENERATION OF ORGANS 27

The organ does not, however, lose its primitive
and typical characters. Actual organic transforma-
tion cannot, therefore, be said to take place in the
case of individual adaptation.

The same leaves of the same plant when grown in water are
ranch longer than those of the terrestrial type ; the leaves have no
stomata, and the epidermic cells are full of chlorophyll (Askenasy,
Ueber den Einfluss dea Wachstumsincdiums auf die Gestalt der
Pflanzen, Bot. Zeit., 1870, pp. 193 and following). Among the
Stratiotes aloidcs it is not uncommon for the upper part of a leaf to
l'ise above the surface of the water while the base is submerged ; the
epidermis at the base contains chlorophyll, but has no stomata,
while the part of the same leaf which rises out of the water is
furnished with stomata, but has no chlorophyll in the epidermis.

(b) A good example of this individual adaptation may be obtained
by cultivating Cacti alternately in the light and in the dark.
Goebel has shown that when a specimen of Phyllocactus is culti-
vated in the dark, the stems are prismatic and thorny ; if the plant
is afterwards placed in the light, the thorns disappear and the
stems become quite smooth. (K. Goebel, Ueber die Emurirkung des
Lichtcs auf die Gestaltumj dry Kactrcn und aadercr Pflanzen, Flora,
vol. 80, p. 96, 1895.)

(c) The animal kingdom furnishes numerous examples of indi-
vidual adaptation.

The gradual drying up of Lake Aral caused the formation of a
number of basins containing water at various stages of concentration.
The Cardium of this region exhibits a whole series of adaptive varia-
tions. The shells become thinner and horny, their shape elongates,
the openings contract, and their colour becomes duller (Bateson).

Mytilus ediUis (the edible mussel), exhibits three different kinds
of shells. It lives either in salt water, deep water, or shallow
water visited by the tide. In each of these three vicinities the
shells exhibit typical aspects.

The direction of boney lamella? is known to agree with that in
which the greatest strain is habitually applied, and the entire
structure of a bone is dominated by the incidences of the forces
applied to it. When, after a badly-mended fracture, the two broken

28 UNIVERSALITY OF DEGENERATIVE EVOLUTION

(i) Our second group comprises the cases of
cellular adaptation which may be produced in the
course of embryonic life and may result in the for-
mation of an embryo from half or the quarter of
an egg.1

Here again the formation of organs takes place
under new and abnormal conditions, but there is
no " transformation " of these organs into fresh
structures exhibiting other characters than those
of primitive and typical organs.

pieces are joined by an oblique segment, the trabecule in this seg-
ment follow the direction from whence comes the greatest strain, a
different direction to that which they would have taken had the
inserted segment been placed parallel with the two broken pieces.

Without entering upon explanatory theories concerning these
facts, it is important to notice that the adaptations of plants exhibit
different characters to those of animals.

With plants it is only when a young organ is born under new
conditions that it exhibits new characters. This is not necessarily
the case with animals. In an animal organism the separation of
the young tissues from the old is not so noticeable as with plants,
as the organs undergo a continuous renewal and can always adapt
themselves more or less to new conditions.

1 When the two cells resulting from the first division of a fertilized
egg of Amphioxus are artificially separated, each cell may develop
directly into a complete individual. The same happens even when
the first four cells are separated from each other artificially.

When this is effected in the case of Echinus or of an Amphioxus
embryo of eight cells, each cell developes as if it had remained a
part of the whole ; but when the blastular stage is reached, that
stage slowly completes itself.

In the first case the cell adapts itself to the new conditions, in
the second case it is the blastular which does so. (See Analytische
Theorie der organischen Entwickelung. Hans Driesch, Leipzig,
1894.)

EVOLUTION AND DEGENERATION OF ORGANS 29

In an adult cell, adaptation may, in a more or
less normal way, cause the formation of new organs.

Even completely specialized cells of an organism
may, when placed under certain conditions of life,
adapt themselves to those new conditions and give
rise to successive generations of different cells, thus
generating other kinds of organs than those to
which they would have given birth under normal
conditions. This happens, for instance, with all the
cells of Salix, which are capable of reproducing the
whole organism by means of budding. This organic
plasticity is shared with the Begonia Rex, but in that
case it is only thoroughly developed in the epidermic
cells. Analogous examples abound among animals,
especially in the lower zoological groups.1

1 Loeb lias shown that in certain groups, when the organism,
after having been wounded, is subjected to unusual light, position,
or pressure, organs are formed at the wounded part, which are
essentially different from those which would have been formed
in a normal recovery. When this heteromorphosis occurs in
Tubularia mesembrya/nthemtim, Aglaqphenia pluma, AntJiennu-
laria rosa, etc., the positions of the oral and apical poles may
be transposed. When Antcnnularia is placed so that gravity
acts upon it in a contrary direction to the ordinary one, there
are formed oral and apical branches, where they would not have
been formed had the organism been kept in a normal position.
(Jacques Loeb, Unters. zur physio/. Morphologic dcr Thierc : I.
Uebcr Hetcromorphosc , Wiirzburg, 1891 ; II. Orgambildv/ag umd
Wachsthum, Wiirzburg, 1892.)

Similarly there is a true heteromorphosis when in a case of
club-foot the faces and articular surfaces of the bones and
cartilages assume characters adapted to the new work thrown
on them as the abnormality of the joint increases. Another
such case is in the false joints sometimes formed when the two

30 UNIVERSALITY OF DEGENERATIVE EVOLUTION

These various instances of cellular adaptation
called " heteromorphosis " by Loeb, are outside
the limits of our present researches, for reasons
which we have already given.

The law of universal degenerative evolution
applies only to the transformations of organs, and
not to their original formation in either individuals
or species, under normal or abnormal conditions.

Section I.
Transformation of organs of animals.

§ 2. Transformation of homodynamic organs in
the individual.

In order to study the degenerative evolution
exhibited in the specific modifications of the
organs of an individual, it is necessary to choose
homodynamic organs that are numerous, and conse-
quently small enough to undergo different kinds of
transformation fitted to the functions they may
have to serve in the different parts of the body.

The numerous appendages of the cray-fish form
an extremely interesting study of this kind. The

parts of a bone do not unite after fracture. In that case a com-
pleted joint with cartilage ligament and synovial membrane may
be formed in the neighbouring tissues under the influence of the
new stimuli. In extra uterine gestation the placenta is formed
upon some abdominal organ. When this occurs, the unusual
stimulus is sufficient to cause the cells of an abdominal organ
to form a perfectly specialized organ of a nature foreign to their
normal life.

TRANSFORMATION OF ORGANS OF ANIMALS

31

appendages of the different metameres (the somites,
or constituent segments of the body) of the cray-
fish are constructed on the same plan. The
typical organ (fig. 2), when complete, consists of
three parts — the protopoditc (pr.),
which is inserted into the body en _
and carries a gill (br.) ; the cndo-
podite (en.) and the cxopoditc
(ex.), each of which consists of a
series of joints which are attached
to the end of the protopoditc
— the cndopodite to the inner
side, the cxopoditc to the outer.

mw

en

Fig. 2.— Diagram of typical
Let US nOW examine the and complete appendage

. of Astacus fluviatilis : pr.

twenty metameres of the animal protopodite; br, giii;

ex, exopodite; en, endo-

in succession. podite.

In each segment of the body we find a pair of
appendages, the structure of which is
based upon that of a complete ap-
pendage, but in which many different
adaptations have brought about the
pr conservation and increase of some of
the typical parts, and the partial or
fig. 3. — Astacus total atrophy of other parts,
append^ onhe Take first the abdominal segments,
seJmenntbdo0finitDhethey carry appendages (fig. 3) formed
J^protopoditeiof the three fundamental parts: the
In, endSS«te P™topodite (pr.), the exopodite (ex.), and
(Huxiey). tne oulopodite (en.). Each of these

parts is itself divided into different joints, the

32 UNIVERSALITY OF DEGENERATIVE EVOLUTION

description of which is not of immediate impor-
tance. The portion of the appendage which has
degenerated is t\\e podobranch, which has completely
disappeared. The reason is obvious ; these ap-
pendages, having become a support for the eggs,
have lost their respiratory function. The part
specially adapted for that function has been allowed
to atrophy, the more readily because the presence
of gills on the ventral surface of the abdomen
would be incompatible with free movement of that

part of the body when the
cray-fish is swimming.

The appendages of the
sixth abdominal segment
are greatly modified into
a caudal fin (fig. 4).
This also exhibits the three
fundamental parts of a

Fig. 4. — Astacua fluriatilis. — Left ap-
pendage of the fith abdominal seg- complete appendage. Hie
ment. Fins (1,5/1) : pr, protopodite; . . . .

ex.exopodite; en,endopodite( Huxley). protopodite (pr.) IS thlCK

and short and has no gill, degeneration being ex-
hibited in the loss of the gill, and by the reduced
length of the part. The exopodite (ex.), and the
cndopoditc (en.) are modified into two large oval
plates which serve as propellers. The actual de-
velopment of these two parts is accompanied by
degeneration. In a typical appendage, the exopodite
and the cndopoditc terminate in slender parts divided
into several rings by false joints ; in the propellers,
which should offer the maximum resistance to the

TRAXSFOKMATIOX OF ORGANS OF ANIMALS

33

pressure of the water, the segmentation has dis-
appeared, but across the part corresponding to the
exojwdite there still remains a transverse groove.

The second abdominal segment of the female
carries appendages similar to those which have
just been described. In the male, the organs of
this segment, as also those of the segment in front
of it, have become organs used in fertilization.
We must consider what new structures
have appeared here, and to what extent
these new modifications have been
attended by degeneration. The ap-
pendage of the second segment (fig. 5)
is longer than that of the other seg-
ments ; pressed against the ventral
surface of the back part of the thorax,
it stretches out as far as the space be-
tween the second and third walking
legs, the part which corresponds to
the oviduct of the female. It serves as
a channel to conduct the fluid from
the male orifice to that of the female.
It consists of a protopodite (pr.)
and an exopodite (ex.) similar to the corresponding
parts of the appendages we have just described.
The endopodite is profoundly modified. The inner
border of its proximal region, which is not jointed,
is extended into a thin plate rolled into a hollow
horn (a) while the outer border is represented by
an annulated part (h). The part in process of

c

Fig. 5. — Astacus
fluviatilis. Left
appendage of the
2nd abdominal seg-
ment of the male
(fnntview)(l,5/l):
pr, protopodite;
ex, exopodite ; en,
endopodite; a,
rolled plate of the
endopodite; b,
the jointed ex-
tremity of the
same (Huxley).

34 UNIVERSALITY OF DEGENERATIVE EVOLUTION

degeneration is the exopodite. On considering the
development of the various parts of this appendage,
it may be concluded that the exopodite has not
undergone the same modification as the endopodite.
It now remains to conclude this study of the
abdominal appendages by an ex-
amination of the first segment.
Immense variation occurs in the
female. Sometimes there are two

Fig. 6. — Astacus fluvia-

tais. Left appendage appendages, sometimes one or both

of the 4th abdominal . .

segment of the fcmaie are missing ; in any case the

(3/1) : pi; protopodite ; . . ° J

«i,endopo<ute (Huxley), existing organs are very small
(tig. 6). The protopodite (pr.) is small, the exopodite
is missing, and the endopodite (en.) is represented
by an imperfectly jointed thread. The appendages
of this first segment being no longer necessary,
atrophy, and, as has just been
pointed out, there is no regularity
or uniformity in the order of their
disappearance. In the male, the
appendages of this segment (fig. 7)
possess an nnjointed rod (t) cor- jHi

responding to the endopodite of the FlG 7 _ Astacus fiuria.
second segment, and this rod fte JJ SSSSffi
stretches out to a considerable l^.^tJnS
distance. The exopodite is missing, (HllxIey)-
and the articulations between the different parts
have disappeared. This may reasonably be regarded
as the result of degeneration, and confirms what has
already been said concerning the appendage of the

TRANSFORMATION OF ORGANS OF ANIMALS 35

second metamere, where degeneration was exhibited
by the relatively small size of the exopodite.

It may be concluded from this examination that
in the abdominal appendages there is a degenera-
tion common to all — this is the absence of the
podobranch. In accordance with the particular
adaptations of each pair of appendages, special
degeneration accompanies special adaptive develop-
ments as we have shown above.

We now pass to the fourteen anterior metameres
of the cray-lish. The first six of these constitute
the head, and carry on the first segment the stalked
eyes ; on the second, the antennules ; on the third,
the antennae ; on the fourth, the mandibles ; on the
fifth and sixth, the two pairs of maxilla;. The
eight metameres of the middle of the body form
the thorax, and carry on the seventh, eighth, and
ninth, the maxillipedes ; on the tenth, the claws ;
and on the eleventh, twelfth, thirteenth, and four-
teenth, the walking legs.

The structure of all these appendages may be
referred to that of the typical primitive appendage.
First take the thoracic appendages, commencing
with the third maxillipede (fig. 8) which is the
most complete appendage. It consists of a proto-
podite formed of two parts (coxopodite and basipodite)
and carries a podobranch (br.), a well-developed
endopodite {en.) consisting of five jointed parts, and
of a small exopodite segmented like those of the
abdominal appendages.

c 2

1G UNIVERSALITY OF DEGENERATIVE EVOLUTION

The third maxillipede may be regarded as the
mean type of the average thoracic appendages.
Compared with it the other appendages exhibit a
true alternative development of their parts. When
the exopodite is large the endopoditc is small, and
vice versd. Besides these general modifications in
which the degenerative evolution of the entire
appendages is exhibited, each one of the segments
undergoes certain modifications of its own.

Fig. 8. — Astacus fluviatilis. Third left maxillipede
(1,5/1): co, coxopodite,and ba, basipodite, forming
pr, protopodite ; br, gill ; ex, exopodite ; en, en-
dopodite (Huxley).

Fig. 9. — Astacus fluviatilis. Second
left maxillipede (1,5/1) : co, coxo-
podite, and 6a, basipodite, forming
pr, protopodite ; en, endopodite,
ex, exopodite ; br, gill (Huxley).

In the second maxillipede (fig. 9), which much
resembles the third, the exopodite (ex.) is large, and
the endopodite (en.) is small, the prodopod,ite (pr.) is
better developed, and the podobranch (br.) has
begun to atrophy.

The same evolution is exhibited by the first

TRANSFORMATION OF ORGANS OF ANIMALS 37

maxillipede (fig. 10), but here it is more striking,
and considerably modifies
the general appearance of
the appendage. The exopo-
dite (ex.) is well developed,
especially at its base ; the
endopoditc (en.) is small and
consists of only two joints ;
in the prodopodite (pr.) the „

1 " vr ' Fig. 10.— Astaeusfluviatihs. First

two component segments leftmaxiiiipedca.s/i):™ coxo-

L ° podite, and 6a, basipodite, form-

n big pr, protopodite ; en, endo-

.ca podite; ex, exopodite ; ep, epi-

podite (Huxley).

are transformed into
two long thin plates,
and the podobranch
is replaced by a mem-
braneous plate (the
epipodite) (ep). Behind
the maxillipedes are
the claws and the four
walking legs. In these
five pairs of appen-
dages the exopodite
shows most signs of
degeneration. We will

Fig. 11. — Astaeus fluviatilis. Second left .

walking lee (1,5/1): eo, coxopodite and HOW examine the COlll-
ba, basipodite, forming pr, protopodite;

br, gill; is, ischiopodite ; me, meropodite; poildlt parts 01 these
ea, carpopodite; p, propodite; da, dacty-

lopodite (Huxiey). appendages (fig. 11).

The, protopodite (pr.) consists of two parts (ba. and
co.), and carries a gill (br.) except in the last walking

38 UNIVERSALITY OF DEGENERATIVE EVOLUTION

leg. The endopodite consists of five primitive joints,
which are well developed and form the ordinary
claw ; these five parts are the ischiopoditc (is.), the
mesopodite (me.), the carpopodite (ca.), the propodite
(pr.), and the dactylopodite (da.). The exopodite is
missing. No vestige or rudiment of it is to be
found in any phase of the development of the cray-
fish. In the lobster, however, which is closely
allied to the cray-fish, the exopodite is still to be
found during the larval period. The third funda-
mental part of the primitive member persists also
in prawns throughout the entire period of life, but
the organ is very small. At the extremity of the
first and second pairs of walking legs there is an
apparatus consisting of a fixed part — an elongation
of the protopodite — and of a moveable part — the
dactylopodite. This furnishes the walking leg with
a prehensile organ which is well developed in the
first pair of walking legs, and which is enormously
increased in the true claws. In this evolution
degeneration is exhibited by the disappearance of a
joint, for in these appendages the basipodite and the
ischiopoditc are immovably united. This morpho-
logical degeneration corresponds to a functional
change in the appendage. So long as the claw was
used for locomotion a joint at this point was in-
dispensable for progression. It is this joint which,
in six or eight-footed beasts allows of the horizontal
motion of the member necessary for locomotion,
which in six-footed beasts results from the general

TRANSFORMATION OF ORGANS OF ANIMALS 39

structure of the parts and their auricular combina-
tions. On the claw becoming prehensile, the joint
hinge consolidates, the lever
thus becoming much stronger
and permitting the claw to be prj'
used to greater advantage.1

It now remains to examine
the appendages situated in
front of the maxillipedes, i.e.
the appendages of the head. Fl0. U.-Astacus fluviatuu.
In the second maxilla (fig. 1 2) ^XoZt^lS^
a special transformation may t^'SokZ' K°SJ&
be observed. The coxopodite nathide (Huxley)-
(co.), and the bctsijiodite (ba.), are flat plates ; the
endopodite (en.), which is small and undivided, ex-
hibits signs of degeneration in its
size, and in the absence of all
articulations. The exopodite, accord-
ing to some authorities on the
via.is.—AtocusAuvia- subject, no longer exists, while

tilt*. First left max- , , . .

iiia (l.s/i): co, coxo- according to others it constitutes

podite, and 6a, basi- ,

podite, forming Pr, with the empodite (the analogous

protopodite; en, en- . °

dopodite (Huxiey). part to that which we regard as
representing the gill in the maxillipede), a large
peculiarly-shaped blade, the scaphognatlride (sc).

In the first maxilla (fig. 13), a partial degenera-
tion of the organ is very marked : the exopoditc and

1 See J. Demoor, Bccherches sur la Marche des Crustace's (Arch,
de Zool. cxp. et gen., 2" serie, t. iv. , 1891).

T. List, BncryuiKjsapparat des Arthropoden, 1. Theil, Astucus
Fluviatilis. Morphol. Jahrhuch., xxii. Bd. 3. Heft, 1895.

W_£^

40 UNIVERSALITY OF DEGENERATIVE EVOLUTION

the epipodite are missing ; the endopodite is reduced
to a mere unjointed stem, and only the protopodite
retains its two normal component parts.

The mandibles (fig. 14), the appendages of the
fourth segment, are modified entirely for mastica-
tion. They consist of a strong transverse piece
(pr.) provided at the extremity with an inner
surface (st.) for grinding and sawing, and of a
three-jointed piece (m.) with bristles which point
outwards. The first piece is the result of the

Fig. 14. — Astacus fl.uvia-
tilis. Left mundible
(1,5/1): pr, protopodite;
en, endopodite ; st, rasp-
ing surface of the proto-
podite (Huxley).

Ftg. 15. — Attacui fluvia-
tilis. Left antenna
(1,5/1) : pr, protopodite ;
ex, exopodite; en, endo-
poditc (FTuxley).

Fig. 16. — Aslacus fluvia-
tilis. Left antennule
(1,5/1) : pr, protopodite;
ex, exopodite; en, endo-
podite (Huxley).

modification of the two parts of the protopodite,
which have united to form the organ of mastica-
tion ; the second, which represents the endopoditc,
is the feeler, an organ of sensation. All the other
parts of the appendage have disappeared.

The antenna (fig. 15), which is a tectile ap-
paratus, is formed of two parts representing the
segments of the protopodite (pr.). The long-ringed
process is the endopodite, while the lateral scale of
the antenna represents a much reduced exopodite (ex.).

TRANSFORMATION OF ORGANS OF ANIMALS 41

The antennule (fig. 1G) consists of a protopodite
(pr.), furnished with an annulated endopodite (en.)
and exopodite (ex.).

The eye-stalk (fig. 17) consists of a two-jointed
protopodite (pr.). This is all that remains, the
endopodite and the exopodite being m,

absent altogether. KiyPr

This examination of the appen- Fig 17_Aslacus fluvia_
dages of the cray-fish clearly shows gj^ . £« ^gj||*
that all fresh adaptation in the (Huxiey).
appendage entails the modification of some parts
and the degeneration of others. In each case
evolution is accompanied by degeneration.

§ 3. Transformation of homologous organs in
individuals of different species.

The limitations of our present knowledge make
it difficult to determine definitely the origin of
limbs among vertebrates, but they are universally
supposed to have developed from the lateral folds
which still persist in Amphioxus, and which pro-
bably existed in the ancestors of vertebrates.1

1 A. Morphological proofs :

(a) Lateral folds of amphioxus.

(b) Identity of the skeletons of paired and unpaired fins.

(c) The number of spinal nerves passing to the fins.

(d) The mode of entrance of these nerves into the fins.
B. Embryological proofs :

(a) Continuous lateral folds in the embryoes of fish.

(b) The formation of metameric pouches (coelomic in-

vaginations not only at the point of origin of the
limbs, but between them).

42 UNIVERSALITY OF DEGENERATIVE EVOLUTION

At first the only skeleton of these lateral rods
consisted of parallel rods of a tough material. By
the transformation of these rods, the skeleton of
the limbs of vertebrates was ultimately formed.

The skeleton of limbs then consisted originally
of a certain number of parallel rods. One of these
rods, lying in the long axis of the future limb,
became longer than the others, while the neigh-
bouring rods began to slant, so that those nearest
to the elongated rod spread out like a fan, and
gradually moved outwards along the principal rod.
This phenomenon was repeated several times, so
that eventually those rods nearest to the principal
rod passed towards the free end of it, and as the
others followed in the same direction, the fin finally
acquired a feather-like structure.

This transformation of the continuous folds into
limbs — a progressive transformation, since new and
more perfectly adapted organs were formed — was
accompanied by degeneration, for a considerable
part of the folds disappeared. In the same way,
although the transformation of the parallel rods
into bipinnated fins constituted a development,

C. Palcvontological proofs :

(a) The series of paired spines in Diplacanthus and in

Climatius between the pectoral and ventral spines.

(b) The skeleton of the fins of Cladoselache.

D. Philogenctic proof: The necessity of the bifurcation of the

unpaired folds at the anal region (Stemarchus).

E. Physiological proof: The lateral folds are the undifferenti-

ated condition of organs of equilibrium in modern fish,

TRANSFORMATION OF ORGANS OF ANIMALS 43

the rods nearest to the free extremity of the
principal rod were considerably reduced in size.

Two series of transformations should now be
followed — that of limbs which have not yet
ceased to be aquatic, and that of limbs adapted
to terrestrial existence.

Among aquatic creatures, the Pleuracanthides
and the Dipneusti (Xcnacanthus and Ceratodus)
have best preserved the bipinnated fin. In
Ceratodus especially it is exhibited in almost the
primitive condition. Gradually, however (as in
Orthacantlius), the rods situated along one edge
of the principal rod disappeared, and the fin, no
longer bipinnated, became unilateral. This pro-
gressive transformation entailed the disappearance
of nearly half of the fin.

With creatures which have become adapted to a
terrestrial life, the limbs — so far as can be judged
from what is known at present — appear to have
undergone the following transformations : The bi-
pinnated fin (such as that of Ceratodus) (fig. 18)
is always the starting-point. Then the lateral rods
of one half almost completely disappear (as in
{Protopterus Amphibius). Next, the other half
follows (as in P. annectcus) (fig. 19), and finally
only the principal rod remains (as in Lcpidosiren)
(fig. 20).

At this point the limbs are reduced to mere
lopped stems ; they have not, however, atrophied ;
the degeneration which has accompanied this de-

Fig. 20. — Lepidosiren paradoxa, F

■genie des Dipneusles, Bull. Soc, beige <le Geol., de Paleont., et d'Hydrol., t. IX., 1895.

ter T. E. Gray). (See Dollo.)

ter Kay Lankcster). (See Doilo.)

46 UNIVERSALITY OF DEGENERATIVE EVOLUTION

velopment has reached its ultimate limit ; the
limb has now to disappear altogether, or else to
provide itself with new elements.

Judging from the embryology of the Urodele
Amphibia, the primitive terrestrial limb had only
one toe at first, and then a second appeared, then
a third, a fourth, and a fifth. In this way was
formed the primitive five-toed limb.

Let us now trace the adaptation of this five-toed
limb to the principal functions it fulfils in the
animal kingdom — walking upon two legs, running,
jumping, climbing, burrowing, swimming, and flying.
We shall find that the transformations it has under-
gone for all these functions have been accompanied
by degeneration.

1. Adaptation to walking on two legs. — Adapta-
tion to the upright position may be effected in two
different ways, by walking or jumping. We will
first investigate the former.

This adaptation to bipedal progression may be
effected with or without the intervention of arboreal
life, as may be seen by the following examination,
the adaptations of man and of birds.

(a) Man. — The first sign, among mammals, of a
definite adaptation to arboreal life, is the oppos-
ability of the great toe. The great toes of human
beings are not opposable, the cases of " prehension "
exhibited among savage races being due to lateral
movements ; we shall see presently, however, that
this opposability of the great toe which is exhibited

TRANSFORMATION OF ORGANS OF ANIMALS 47

by all modern monkeys, probably existed at some
time in man or his immediate ancestors.

The terrestrial five-toed foot must have acquired
an opposable great toe in the course of adaptation
to arboreal life. Without attempting to enumerate
all the attendant phenomena of degeneration en-,
tailed by this modification, we will merely examine
those exhibited by the nails of the hind foot.
Comparison with the supposed ancestors of the
Primates (Insectivores) shows that in the primitive
terrestrial foot the nails completely covered the
extremities of the terminal phalanges. With the
lowest Lemur (Tarsius) the nails of the second and
third toes are in the form of claws, while all the
others are degenerating ; they cover only the
anterior face of the terminal phalanges, and are,
in fact, flat nails. In all the other Lemurs, only
the second toe has a claw, whereas in true monkeys
all the nails are flat. Finally, in the Orang the
great toe often has no nail at all.

We then come to limbs adapted to arboreal life,
such as those of the Gorilla. With some anthropoids,
however, and especially the Orang, the modification
has gone further, and is accompanied by phenomena
of degeneration in other parts of the limb. While,
in the case of the Gorilla (fig. 21) the great toe
has a true articulation between the metatarsal and
the first phalanx, and also between the first and
second phalanges, with the Orang (fig. 22) the great
toe is much reduced so that it has no articulationrv

?/£13RAV

48 UNIVERSALITY OF DEGENERATIVE EVOLUTION

between the metatarsals and phalanges, and fre-
quently the last phalange is absent altogether.

The development of another foot, that of man,
adds confirmation to the heading of this chapter.
The great toes of man or his immediate ancestors
.must have been opposable. This may clearly be
seen by a careful study of the muscles. We know
what a resemblance there is between the muscles

,7YiUa£o«>e.

Figs. 21 and 22. — Skeletons of the feet of the Gorilla and Orang-outang.
ca calcaneum; as, astragulus; sc, scaphoid; Ire cu, first cuneiform; 2e cu,
second cuneiform ; 3e cu, third cuneiform ; c, cuboid ; if, metatarsals ; phal,
phalanges; ph and p, distal and terminal joints (after Waterhouse and Hawkins).
•See Huxley, Man's Place in Nature,

of the soles of the feet and those of the palmar
region. With the exception of man, all mammals
which have opposable thumbs, have also opposable
great toes. On the other hand, there are various
mammals (prehensile marsupials, rodents, bats, mar-
mosettes, etc.) which have opposable great toes but

TRANSFORMATION OF ORGANS OF ANIMALS 49

not opposable thumbs. "We may presume then
that the hind limbs were the first to become
adapted to arboreal life, and, since the great toe
of man is not opposable, it may be concluded that
it ceased to be so when terrestrial life was assumed.
This modification is not wholly degenerative, for
the functions of the lower limbs are no less im-
portant in man than in apes. There are, however,
undoubted signs of degeneration in the changes
entailed. The muscles which rendered the great
toe opposable were probably the first to degenerate,
but by referring to a skeleton it may be seen that
whereas the great toe and the metatarsals have
gained in length, the first phalanges, and still more
the other phalanges, are reduced. In some cases
the last phalange of the little toe has even dis-
appeared altogether.

(b) Birds. — First take the reptile-footed Dino-
saurians. These are four-footed beasts, and the
hind limbs are plantigrade and five-toed. Counting
from the inner side the phalanges of the toes are :
2, 3, 4, 5, 4 in number.

In passing on to the bird-footed Dinosaurians
we find that with Hypsilophodon, the foot is func-
tionally four-toed. From the first to the fourth
toe the phalanges are the same in number as with
the reptile-footed Dinosaurians, but the fifth toe is
greatly reduced, being merely represented by a
pointed metatarsal. In Camptosaurus, the fifth toe
has disappeared, the second, third, and fourth are

D

50 UNIVERSALITY OF DEGENERATIVE EVOLUTION

fully functional, but the first, although retaining the
usual number of phalanges, is greatly reduced in
size. Finally, in Iguanodon,1 which is fully adapted
to the upright position, the second, third, and fourth
toes are still fully functional, but not only has the
fifth toe disappeared, but the first is merely repre-
sented by a pointed metatarsal.

Thus it is apparent that each new phase in the
transformation from quadruped to biped is attended
by partial degeneration, although the hind limbs,
having alone to fulfil the function of locomotion,
necessarily acquire a fuller development. The
Iguanodon is too specialized in its development
to have been the ancestor of birds ; but this is
not so with other bird-footed Dinosaurians, such
as Camptosaurus and Hypsilopliodon, where the
hind foot is sufficiently primitive to have been
the forerunner of the bird's claw.

Most birds have four functional toes, which have
retained the primitive number of phalanges, but the
legs of running birds exhibit more or less important
variations. The modifications they have undergone
have obviously not diminished their functional im-
portance, but quite the contrary. These modifications
have, however, been accompanied by degeneration
with running birds, and the first toe disappeared
first, as in the Cassowary and the American Ostrich,

1 Dollo : First, second, third, fourth, and fifth Notes sur les
Dinosauriens clc Bernissart. Bull. Mus. royal d'Hist. nat. de Belg.
1882, 1883, 1884.

TRANSFORMATION OF ORGANS OF ANIMALS 51

to be followed in the second stage by the second
toe. Then the fourth began to atrophy until the
whole weight of the body was supported by the
third toe alone, as in the ostrich.

2. Adaptation to leaping — We now come to the
various instances of adaptation to leaping. This
adaptation is one necessarily undergone either by an
animal which originally lived a terrestrial life, such
as the Jerboa, or by one which had ceased to be
arboreal in adapting itself to leaping, such as
the kangaroo ; or, again, by an animal which
formerly lived an arboreal life, and which con-
tinued to lead such a life with the additional
adaptation to leaping, such as the tarsius ; and,
finally, by an animal which first lived and swam
in the water, and which continued to do so after
becoming adapted to leaping, such as the frog.

In each of these cases adaptation to leaping has
resulted in the fuller development of the hind limbs,
the size of which, as well as the functional impor-
tance, has greatly increased, which shows that the
modification is of a progressive nature. All these
modifications are, however, attended by degeneration.

(a) The Jerboa (Dijms). — Some rodents, such as
rats, porcupines and squirrels, are five-toed. This
five-toed foot represents the primitive one from
which, after passing through various still existing
phases, was derived the three-toed foot of the jerboa.

With the hare there is no first toe, so that the
foot is functionally four-toed.

52 UNIVERSALITY OF DEGENERATIVE EVOLUTION

The fifth toe next disappeared, leaving the foot
three-toed, as in the viscacha, the
capybara and agouti. In the foot
of the agouti the metatarsals have
considerably gained in length, and
are closely pressed together.

In the jerboa (fig. 23) the three
metatarsals (M.) are united, and
the intervening tissues have dis-
appeared. Besides this modifica-
tion, the hind limbs of the animal,
being used for leaping only, have
become thinner, while gaining in
fig. 23. —Hind nmb of length by the elongation of the

Dipus (Jerboa). I, first •'. .

(h) The Kangaroo. — Although
now inhabitants of the plain,
kangaroos are descended from
arboreal marsupials, which them-

rudimentary digit; II, metatarsals.1

second digit; III, third

digit; IV, fourth digit;

M, bone formed by

fusion of 2nd, 3rd, and

4th metatarsals ; ca,

caleaneum; as, astra-

gulus; n, navicular;

c3, third cuneiform ; c-,

second cuneiform ; cb,

cuboid. (After Flower, selves have gradually ceased to be

Introduction to the Oste- ° '

oiogy of the Mammalia.) arboreal. All the intervening
phases of this transition from type to type are known.

1 The last segment of the leg of the jerboa — the foot being
formed by the union of the second, third and fourth metatarsals
— corresponds more or less to the formation of the bird's foot.
This, however, is a case of convergent modification more apparent
than real, for in the cannon bone of birds, besides the second, third
and fourth metatarsals, there is the whole set of distal tarsals,
whereas in the jerboa the distal tarsals remain free and isolated
bones.

2 L. Dollo, autographs of the course given at the Institute of
Solvay, 1891-1892, 5th lecture, The Origin of Kangaroos.

TRANSFORMATION OF ORGANS OF ANIMALS

13

<-C^>i

In the Opossum (Didelphys) we find a normal
five-toed foot, in which the third toe predominates,
bnt which is fully adapted to arboreal life, the first
toe being opposable and provided with a fiat nail.

The foot of the Phalanger
(Plialangista) (fig. 2-4) is
also five-toed, and the great
toe is opposable, but here it
is the fourth toe (iv.) which
predominates owing to the
degeneration of the second
(ii.) and third (iii.)

The Kaola (Phascolardos)
exhibits the same type of
foot, but with an even more
marked diminution of the
second and third toes.

In Hi/psiprijiniwdon the
first toe has become rudi-
mentary.

In Peramdes the first toe
is only represented by a
metatarsal with phalanges.

In the kangaroo (fig. 25)
it has completely disappeared.

Finally, in Clueropus (fig. 26) the fifth toe is
reduced to a mere thread, as are also the second
(ii.) and third (iii.) toes.

This last stage, which represents the present and
most perfect adaptation of marsupials to leaping,

Fig. 24.— Right fore-paw of
Phalanger.
c, calcaneum; a, astragulus; n.
os eentrale ; cb, cuboid ; t-1, c-, c:;,
firstjSei'ond. and third cuneiforms ;
I, II, III, IV, V,— 1st, 2nd, 3rd,
•ftli and 5th d'gifs, of which the
second and third are degenerating.
(After Flower, Introduction to t/ie
Osteoloijy of the Mammalia.)

54 UNIVERSALITY OF DEGENERATIVE EVOLUTION

was attained by the predominance of the fourth toe,
accompanied by the almost total extinction of the
other toes. The persistence of the fourth toe

Fig. 25. — Right fore-paw
of Kangaroo.

r, calcaneuni; a, astragu-
lus; cb, cuboid ; n, os
centrale; on.third cunei-
form. II, III, IV, V —
second, third, fourth
and fifth digits, the
second and third being
degenerate. (After

Flower's Introduction
to the Osteology of the
Mammalia.)

Fig. 26. — Choeroput.
Right fore-paw.
c, ealcaneum ; a, astraga-
lus ; n, os centrale; cb,
cuboid; c3, 3e, third
cuneiform; II, III. IV,
V, — second, third,
fourth and fifth digits,
of which the second,
third and fifth are de-
generating. (After
Flower's Introduction
tn the Osteology of the
Mammalia.)

Fig. 27. —Right hind-paw
of Tarsius Spectrum.

c, ealcaneum ; a, astragu-
1ns; r, navicular; c1,^-,
c3, first, second, and
third cuneiforms ; eb,
cuboid ; I, II, III, IV,
V,— first, second, third
and fourth digits. (After
Flower's Introduction
to the Osteology of the
Mammalia.)

instead of the third is accounted for by the
previous arboreal existence of marsupials before
they became adapted to leaping.

(c) The Tarsius (Tardus spectrum). — The tarsius,
which is a small lemur, is to be found in Celebes.

TRANSFORMATION OF ORGANS OF ANIMALS 55

It is so-called owing to the remarkable length of
its tarsus (fig. 27), a very exceptional formation,
for the elongation of an animal's foot is usually
effected by means of a lengthened metatarsal.
This abnormal formation in the foot of the
tarsius can, however, be accounted for. Like
most of the Primates, the tarsius is adapted to
an arboreal life, and, among other characters, it
exhibits an opposable great toe. When on the
ground, however, instead of running on two or
four legs, it progresses by leaping ; and this
may also be noticed with some of the lemurs
of Senegal (Galago). The tarsius has become
adapted to leaping by the lengthening of the
hind limbs, the thigh, leg and foot being equally
elongated. The foot, however, being of fan-like
structure, it is clear that if all the toes were
elongated, the feet would overlap one another
in the middle line, and therefore with most
animals that leap the elongation of the foot
involves a reduction in size of the outer and
inner toes to prevent overlapping. The tarsius,
however, has remained arboreal, although it has
taken to leaping, and it therefore requires a well-
developed foot with an opposable great toe in order
to obtain a firm grip on branches. To avoid over-
lapping of the feet, the length has been augmented
at the ankle, which, not being shaped like a fan,
does not consequently gain in length.

A more detailed examination would doubtless

56 UNIVERSALITY OF DEGENERATIVE EVOLUTION

show that this modification of the foot has been
attended by the degeneration of some parts of it,
but, as we have to restrict ourselves here to the
data with which we are already furnished, it is the
degenerative phenomena exhibited by other parts of
the limb consequent to adaptation to leaping, which
we shall proceed to enumerate.

There are two leg bones, the fibula, and a larger
one, the tibia. These two bones are usually held
together by muscles, or by an interosseous membrane,
the latter being merely degenerated muscular tissue.
As a rule, the tibia and the fibula reach the entire
length of the leg and are quite separate. In the
process of adaptation to arboreal life, these bones
have remained apart, as may be observed in most
arboreal animals of the present day. Moreover, in
order to promote prehension by the foot, the tibia and
fibula in prehensile-footed marsupials may turn upon
one another, as is the case with our radius and ulna.
With the tarsius it is quite otherwise. Here the
lower halves of the tibia and fibula are united, and
the muscles which formerly connected the two bones
have consequently disappeared. There is further
evidence of degeneration in the upper half of the
fibula which has become reduced to a mere thread.
There is no question that these phenomena are the
result of adaptation to leaping, for they are not
exhibited among exclusively arboreal animals, while
many parallel cases of modification are known to
exist amongst other species.

TRANSFORMATION OF ORGANS OF ANIMALS

57

(d) The Frog (Rana Esculenta). — Adaptation to
leaping entails, as a rule, the elongation of the hind
limbs. When a large blade-like swimming foot is
required, the number of toes cannot be diminished
and length must be augmented, as in the tarsius, in
the region of the ankle (fig. 28).
But, in the frog, it is the
astragulus (a.) and the calcan-
eum (c.) which have elongated,
whereas in the tarsius it was
the calcaneum and navicular
bones.

Besides the union of the
calcaneum and the astragulus
bones — which may be regarded
as a phenomenon of degenera-
tion, as it entails the disappear-
ance of the intervening tissues
— the adaptation of the frog to
leaping has been attended by a Ro. at-Rjghtfootof Sana
yet more characteristic sign of a< Mtn»gn™s'"'"; oaican
degeneration : the tarsus, which
is a very complicated structure
in tailed amphibia, is reduced,
especially in the distal row, to a few minute
bones.

3. Adaptation to running, — The ruminants and
the horse furnish us with two excellent examples
among mammals of adaptation to running.

Their common ancestor is known to have been a

i um ; 1-3, n, h, A1, tarsals ;
m, metatarsals; I. II, III,
IV, V, digits. (After
Ecker, Anatomy of the
Frog.)

58 UNIVERSALITY OF DEGENERATIVE EVOLUTION

clumsy five-toed animal which lived in swamps. As
the horse type was evolved, the legs, which originally
only served to support the weight of the body in a
slow progress, were gradually adapted to running.
This modification was certainly advantageous to the
animal, as it enabled it to escape more easily from
its enemies. It was, however, accompanied by de-
generation, for the ancestor of the horse lost first the
great toe, then the fifth, and next the second and
third toes became so reduced that eventually only
one toe remained functional.

Adaptation to running was effected differently
with the ruminants, but it was equally accompanied
by degeneration. With them, first the great toe
disappeared, then the second and fifth became simul-
taneously much smaller, while the third and fourth
continued to increase equally in length. In process
of time the second and third toes entirely disap-
peared, and the metatarsals of the third and fourth
joined together to form the cannon bone. In this
way mammals were evolved with cloven hoofs and
adapted to running.

4. Adaptation to flying. — There are three types
of animals among the vertebrates : birds, ptero-
saurians, and flying bats, by which is meant, not
those bats which are merely able to float in the
air like a parachute, but those which can both raise
and guide themselves during flight.

(a) Birds. — We have seen that the ancestor of
all terrestrial vertebrates was a five-toed animal.

TRANSFORMATION OF ORGANS OF ANIMALS 59

The hand of the oldest known bird, however, —
Archceopteryx — had only three digits, and the
question arises as to whether this atrophy of the
two outer digits was effected in the process of
the animal's gradual adaptation to flight or if it
dates from a still earlier period.

Archceopteryx had four toes. The fore legs were
smaller than the hind legs. Now, with bipedal or
quadrupedal animals which make no special use
of their fore legs, the contrary may invariably be
noticed. It is of small importance, therefore,
whether Archceopteryx was quadrupedal or bipedal
before it took to flying, for neither of these modes
of progression would have entailed the loss of two of
its digits, and the disappearance may reasonablybe at-
tributed as due to the process of adaptation to flight.

The three remaining digits of Archceopteryx were
complete in their structure. The phalanges, which
were of different lengths, are furnished with claws,
and were of the same number as is normal among
reptiles, i.e. 2, 3, 4, counted from the inner side.
The wing of Archceoptcryx had not therefore under-
gone much modification beyond the toes of the two
outer digits, but it must be remembered that the
bird was a weak flyer ; instead of having a large
boney sternum necessary for powerful flight, the
sternum of Archceopteryx was formed of cartilage,
and the wings were short and rounded in shape.

With good flyers, on the other hand — such as
pigeons or sea-birds — the adaptation to flight is

60 UNIVERSALITY OF DEGENERATIVE EVOLUTION

more perfect, the modifications entailed are much

greater and have been accompanied by further signs
of degeneration. In the fowl
(fig. 29) all the claws have
disappeared {degeneration No. 1) ;
the thumb (i.) and the index
finger (ii.) have each lost a
phalanx ; the middle finger (iii.)
has lost three {degeneration No.
2) ; finally, the metacarpals of
the index and middle fingers
have fused (a.) thus entailing
the degeneration of the muscles
between those parts {degeneration
No. 3).

(6) The Pterosaurians. — These
reptiles, which are now extinct,
were able to fly, like bats, by
means of a membrane. There
is no connection between them

f,g. 29.-skeieton of wing and birds which fly in quite a

i, First digit orW thumb ; ii, different way.
mtLoV'St digit; The membrane of the Ptero-
th/TLranTthira saurians (fig. 30) was supported

digit,. (After Huxley.) by ft framework of which the

most important part was the fifth finger (v.), the
digit corresponding to our little finger, which was
greatly elongated.

The patagium (the flight membrane) consisted of
two kinds of membrane, the antebrachial membrane

TRANSFORMATION OF ORGANS OF ANIMALS

61

which extended from the neck to the wrist, and the
true wing membrane, which reached from the free
end of the little finger to the ankle.

There are three essential types of Pterosaurians.
Pterodactylus, which had a short tail and teeth all
along the gape;
Rhamphory -
hnchus, which
had a long tail
and a beak set
in the front
part of the jaw
with teeth be-
hind it ; and
Pteranodon,
which had
neither teeth
nor tail. This
last type was
the most spe-
cialized of the
Pterosaurians,
and the strongest flyer of the whole group. They
attained to a huge size, the skull measurement averag-
ing 1 metre 20, and the wing extension 8 metres 30.
The modification which the fore-limbs of the Ptero-
saurians underwent in the course of their adapta-
tion to flight, was accompanied by degenerative
phenomena in proportion to the extent of the
specialization (see Pterodactylus spectabilis) (fig. 30).

Fig. 30. — Pterodactylus spectabilis (after Huxley,
Anatomy of V^rlebrated Animals).

62 UNIVERSALITY OF DEGENERATIVE EVOLUTION

In the process of adaptation to flight the thumb (i.),
and the little finger (v.) first lost their nails and
became useless. Next, the metacarpal and the
proximal phalange of the thumb became much
smaller, and united with the bones of the wrist,
while the distal phalange degenerated to a mere
thread. Fresh evidences of degeneration presented
themselves as the specialization became more ad-
vanced. In Ptera-
nodon, for instance,
the metacarpals of
the clawed digits
were considerably
reduced in size, the
arms having almost
ceased to exercise
any function but
that of flight. There
was a special pro-
vision for the support
of these great wings,
the development of which was only equalled among
vertebrates by that of the pectoral fins of the
skate. Whereas the legs were upheld by the pelvis,
which was in its turn supported by the sacrum, the
arms of Ptcranodon were attached to the middle of
the shoulder blade, the latter being supported by
the vertebral column — an absolutely unique con-
dition. This kind of pectoral sacrum implied the
fusion of a number of vertebrae, and involved a

Fig. 31. — Pteropus (a bat). Skeleton of fore-
limb: ;•, radius; c, degenerating ulna ; I, II,
1II,IV, V, digits. (After Huxley, Anatomy
of Vertebrated Animals.)

TRANSFORMATION OF ORGANS OF ANIMALS 63

consequent degeneration of the muscles which in
earlier Pterosaurians caused the movement of one
vertebra upon another.

Two phases may thus be observed in the evolu-
tion of the Pterosaurians : first, Ptcrodeictylus and
Bhamphorynchus, and next Pteranodon, while de-
generation attended the modifications of both.

(c) Bats. — Bats, like the Pterosaurians, fly by
means of a membrane, only that the membrane,
instead of principally extending from the little
finger to the body, is equally developed between
all the fingers.

Let us consider what degeneration is involved by
this modification.

Firstly, all the digits have lost their nails, except-
ing the thumb and the index finger in fruit-eating
bats, and the thumb alone in insect-eating bats,
while some of the phalanges of the digits are
missing, the usual number being two instead of
three. Further, instead of the ulna and radius
being equally developed in the fore-arm, the ulna
is greatly reduced in size.

5. Adaptation to arboreal life. — We have seen
that, as a rule, the first evidence of adaptation to
arboreal life is the opposability of the great toe.
This modification is accompanied by the degeneration
of the nail, for, instead of consisting of a claw cover-
ing the entire extremity of the last phalange, it
becomes a small nail covering only the upper side.

Another modification is exhibited at a slightly

64 UNIVERSALITY OF DEGENERATIVE EVOLUTION

later stage. In the primitive foot of the aminota,
the third toe is the longest, but among animals
which have become adapted to an arboreal life (fig.
32) the fourth toe preponderates, thus allowing of
a wider grasp, and the second (ii.) and the second
and third toes gradually degenerate (Ardocebus
calabar ensis, Potto). A clear proof that this modi-
fication is due to adaptation to
arboreal life, lies in this same
tendency being exhibited by two
groups not closely allied to one
another — the marsupials and the
lemurs.

In a still more advanced stage
the phalanges of the second digit
fig. z%— Ardocebus mia- disappear almost completely, and,

barensis. Right hand. ri r * '

i ii, in, iv, y=the Ave for .d\\ practical purposes, no

fingers, of which II and r r r >

III are degenerating, longer exist.
(After Huxley, Anatomy °

of vertebrated Animals.) Among reptiles, chameleons
furnish a striking example of adaptation to climbing.
With them three digits are opposable to the other
two (figs. 33 and 34), instead of one being oppos-
able to the other four. To further promote this
opposability, there is a fusion of digits (syndactilism),
i.e. each set of digits is enclosed up to the nails
within a common integument. The result of this is
that the enclosed digits are no longer capable of
lateral movement, and the muscles have degenerated
in consequence. The wrist and tarsus also show
signs of partial degeneration.

TRANSFORMATION OF ORGANS OF ANIMALS

05

6. Adaptation to swimming. — Among mammals,
adaptation to swimming entails the functional dis-
appearance of the lower limbs, and the modification
of the fore-limbs into fins.

This modification involves considerable degener-
ation. The alteration in the number of digits
varies ; the whale-bone whales have kept all five ;
in rorquals only four remain ; and, after a series

cd/e

Fig. 33.— Skeleton of hand of
Chameleon. (After Cuxier.)

Fig. 34.— Chameleon. Skele-
ton of foot. (After Cuvier.)

of intermediate conditions, the dolphin is evolved,
the fins of which are functionally two-toed. If the
Cetaceans vary as to the number of digits they ex-
hibit, they all have this character in common — the
joints have disappeared. The arm no longer articu-
lates at the elbow, and neither the wrist nor the
phalanges are jointed. The simple stratum of
cartilage which takes the place of the true joints
allows of a slight movement of the various
segments, but not of true articular movement.

E

66 UNIVERSALITY OF DEGENERATIVE EVOLUTION

This results in the degeneration of all the

muscles which caused this mobility in their

terrestrial ancestors.

A similar degeneration may be observed among

Plesiosaurians, Ichthyosaurians, and Mosasaurians.1
With regard to Sirenia, the Dugong is unable to
move the arm at the elbow
joint, but the Manatee can
grasp things with its fins.
Besides many other modifi-
cations, the adaptation of
these animals to an aquatic
life has entailed the fusion
of the two bones of the fore-
arm, thus involving the de-
generation of those muscles
which formerly prevailed
over the movement of the
two bones when separate.
7. Adaptation to burrow-

ce, os centrale; rs, falciform %n0- There are tWO types

XSjoi^digt^T/ft^ of adaptation to burrowing
flower-) —that of the Mole and

that of the Hetcrocephalus.

In the mole (fig. 35), the fore-limb, although it

has become shorter in order that it may better fulfil

Fig. 35. — Fore-limb of Talpa

europaea (the Mole).

R, radius ; s, scaphoid ; e, semi

lunar ; c, cuneiform ; p, pisiform ;

n, unciform ; m, os magnum ;

1 L. Dollo, Premiere note sur les Mosasauricns de Mesvin {Bull.
Soc. belg. Giol. PaUont. Hydrol., vol. iii., 1889) ; Nouvellc note
sur Vosteologic de Mosasauriens {Bull. Soc. belg. Giol. PaUont.
Hydrol., vol. v., 1892).

TRANSFORMATION OF ORGANS OF ANIMALS 67

its function, is obviously in process of development,
for it has not only retained all five digits, but, what
is functionally a sixth digit, has made its appear-
ance (rs.)- The phalanges are very well developed
(phal.), but, on the other hand, the distal phalanges
(ph.) have degenerated and are very short.

Heterocephalus (fig. 36) is a burrowing rodent,
only, instead of being talpoide as is the Bathyergus,
i.e. instead of exhibiting a body furnished with

Fig. 36. — Heterocephalus PHlippi. (After Oldfield Thomas.)

strong short legs, Heterocephalus looks more like
an ordinary quadruped. In process of adaptation
to an underground life, the hair, especially that on
the legs, has disappeared. This is the more re-
markable, as in no other case does it occur among
mammals unless as an adaptation to aquatic life.
Here it is in the atrophy of the roots of the hair
that degeneration is manifested.

One form of hair, however, is still to be found
on Heterocephalus, for there are a few bristles on
the outer sides of the feet. These serve as brushes
to sweep away the sand while burrowing.

68 UNIVERSALITY OF DEGENERATIVE EVOLUTION

Section II.
Modification of the organs of plants.

§ 4. Modification of homodynamic organs in the
Individual.

For the same reasons as we have already
pointed in the case of vertebrates, those leaves

which grow out of
different parts of the
main stem, and have
undergone many
modifications in
order to adapt them-
selves to their vari-
ous functions, will
best serve as ex-
amples for our pre-
sent demonstration.
In descriptive
botanical treatises

Fig. 37. — Growing point of \icia vana,

showing the origin of leaves the term "leaf" is

1, 2, 3, 4, leaves respectively older; leaf 1 is
still in .the primordial condition ; leaf 2 is applied without dis-
divided into epipodium and hypopodium (h). L r

In leaves 3 and 4 the epipod is divided into crimination. As a

petiole and leaflets, and the hypopod forms

8tiPu,es- rule, however, this

term should only be applied to the true leaves, i.e.
those of which the parts are fully developed
throughout.1

1 The embryological development of the leaf among the Angio-
spernis (fig. 37) exhibits the following phases : — The leaf arises
from the growing point in the form of a small bud, which grows

MODIFICATION OF THE ORGANS OF PLANTS 69

These foliage leaves, which are solely assimilative,
are philogenetically the oldest, for the assimilative
function of leaves was certainly an earlier function
than any they may now exercise ; but it is not un-
common to find both basilar (Nicdcrbl&tter) and
apical leaves (Hochbldttcr) as well as foliage leaves
(Laubbldttcr) in the same plant, and even growing
from the same branch. These leaves, which arise
directly from the foliage leaves, have more or less
completely lost their primitive function, and have
assumed others. Their structure has undergone
corresponding modifications which are all attended

for some time without any evidences of differentiation, and con-
stitutes what is called the primordial leaf. Next the leaf is
differentiated into a proximal part, which almost surrounds the
stem (the hypopod), a distal part (the epipod), and an inter-
mediate part (the mesopod). From this point the various parts
of the leaf begin to develop individually.

Little foliacious lamina (the stipules) frequently arise laterally
from the hypopod, and, while still in the bud these stipules, being
often much larger than the rest of the leaf, serve as a very effective
protection to the young organs.

The way in which the epipod differentiates varies very much in
different cases. The lamina is developed from it, and little buds
are gradually formed around the point until the whole is ready to
branch out. At this stage the young leaf is still rolled up within
itself, and protected by stipules, when there are any.

Next, the mesopod grows into the petiole, and the growth of the
petiole causes the separation of the different parts of the leaf.

The leaves of many plants are much less complicated than
these. Sometimes the mesopod is missing, and the hypopod and
the epipod are left in contact. Sometimes there is practically no
hypopod, and in some cases the primordial leaf develops with-
out any differentiation into hypopod and epipod. See fig. 49
( Sempervi vum arach noideu m).

70 UNIVERSALITY OF DEGENERATIVE EVOLUTION

by degeneration. A good demonstration of this will
be to examine the leaves of the following types :
Rosa rugosa, Serratula ccntauroides, Sagittaria sagitti-
folia, Lathyrus Aphaca, and Nymphaca dentata.1

1. Rosa rugosa. — In the foliage leaves (fig. 38, f)
of a rose-branch, such as of Rosa rugosa, there are two

Fig. 38. — Leaves from the same branch of Rosa rugosa.
A — E, successive basilar leaves ; F, foliage leaf; G — I, successive apical leaves.

lateral stipules which arise from the hypopod and
are fused with the base of the petiole. The petiole,
which arises from the mesopod, has from two to six
pairs of leaflets, and also a terminal leaflet — all of
which are derived from the epipod.

1 In studying these types we will confine our attention to the
leaves arising directly from the foliage leaves, to the exclusion of
the floral leaves.

MODIFICATION OF THE ORGANS OF PLANTS

71

The basilar leaves (fig. 38, a to e), which are
usually withered in an adult branch, have more or
less completely lost their assimilative function, and
merely serve as a protection to
the bud. The hypopod plays an
important part in this modifica-
tion, but there are evidences of a
partial degeneration, the epipod
being reduced in size in propor-
tion as the assimilating function
is lost.1

The apical leaves also have
partially lost their assimilative
function, and have assumed, like
the basilar leaves, a protective
function which they exercise on
the floral buds. This modification
is effected differently to that of
the basilar leaves, but is equally ^25^j&<a8£:
attended with evidences of degener- F'bract of the involucre-
ation. The hypopod continues to gain, as the epipod
loses, in importance, but the leaflets, instead of de-

Fig. 39. — Leaves from the
same branch of Serra-
tula centauroides.

1 On examining the foliage leaves of a plant from above down-
wards, one finds successively leaves with large stipules and small
leaflets (fig 38, e), then leaves with very small leaflets crowded at
the end of the hypopod and which have only the basilar part of the
petiole (fig. 38, d), and then leaves of which the epipod and the
free part of the petiole have disappeared (fig. 38, c). Finally, at
the base of the plant there are leaves in which the hypopod is
markedly reduced, and which carry nothing but small stipules
(fig. 38, a and b).

72 UNIVERSALITY OF DEGENERATIVE EVOLUTION

creasing in size, decrease in number. At first there
may be three leaflets, and then only a single large
leaflet (fig. 38, g).
Finally, towards the
floral end of the
branch the leaflet be-
comes smaller and
smaller until it dis-
appears altogether
(fig. 38, H and i).
The apical leaves, like
the basilar leaves, are
eventually reduced
simply to the hypopod.

2. Serratula ccn-
tauroides. — The foli-
age leaves of this
species are close to
the ground, their bases
are ensheathed, and
the long petioles ter-
minate in rhomboidal
blades.

In passing on from
these assimilative
leaves to the basilar
and apical leaves, a
double adaptation becomes apparent, as in Rosa
rugosa. In Rosa rugosa, however, both adaptations
were protective, while in Serratula ccntauroidcs this

Fig. 40. — Sagittaria sagittifolia.
A, winter bud with membraneous leaves en-
closing the swollen portion and ro led leaves
protecting the bud ; B,C, submerged leaves;
D,E, floating leaves ; F, aerial foliage leaf;
G, leaf protecting the flower-bud ; H,
seedling grown from a seed planted in the
water on the surface of the mud. I, seed-
ling from seed planted in ihe water under
a layer of mud. In figures H and I, c is
the cotyledon, I the first leaves.

MODIFICATION OF THE OBGANS 0F PLANTS 73

is only the case with the basilar leaves, which sur-
round the winter buds.

The modifications of the basilar and apical leaves
of Serratula centauroides'axe similar to those of Bosa
rugom, and it is therefore unnecessary to repeat the
description.

Those of the apical leaves tend towards their
protection from rain. When rain falls upon the
radical leaves, they are flattened out upon the
ground, which prevents them from being torn.
This is not so with the apical leaves, and had
their primitive structure remained unchanged, they
would have been constantly exposed to destruction.
M. Stahl l has shown how these leaves have pro-
tected themselves by shortening their petioles and
fringing their edges. In this way the leaf has
acquired greater resistance, and the water is better
enabled to run off it. This modification, however,
although one advantageous to the plant, has in-
volved the partial degeneration of the petiole and
the blade.2

3. Sagittaria Sagittifolia. — The Sagittaria is an

1 E. Stall], Reycnfall unci Blattgestalt (Ann. du Jard. hot. dc
BuUcnzorg, vol. xi., 1893).

2 In the leaves (fig. 39, u and c), immediately above the foliage
leaves, the petiole gradually disappears altogether, while the blade
has become deeply serrated, thus providing for the water drainage
while retaining an assimilative surface.

Further up the stem the leaf blades are more and more reduced,
until eventually they are no longer differentiated into hypopod and
epipod, but arise directly from the growth of the primordial leaf
(fig. 39, i) to f).

74 UNIVERSALITY OF DEGENERATIVE EVOLUTION

aquatic plant, the foliage leaves of which emerge
from the water. The blade of these leaves (fig.
40, f) is shaped like an arrow-head, and the barbs
facilitate rain-water drainage. The petiole is very
long, and terminates in a sheathed base derived
from the hypopod.

The apical leaves (fig. 4, g) are represented by
transparent membraneous scales, which serve as a
protection to the floral buds.

The basilar leaves, which either float on the
surface of the water or else are entirely sub-
merged, have undergone both development and
degeneration. Degeneration is more noticeable on
glancing downwards from the foliage leaves to
those which protect the winter buds.1

1 (a) The leaves nearest to the foliage leaves float upon the
surface of the water (fig. 40, e). The lateral barbs,
being no longer required, have begun to degenerate.
{b) The floating leaves immediately beneath exhibit barbs
which are still more reduced (fig. 40, d), and there are
no stomata except those upon the upper surface.

(c) The next leaves are completely submerged and lengthened

out into long ribbons which broaden into flat blades
at their upper ends, which are not separated from the
hypopod by petioles (fig. 40, c). In this case the blade
is very much smaller, and the petiole has disappeared.

(d) The next leaves to these are submerged, and are ribbon-

like in shape. They spring immediately from the
primordial leaf without differentiating into hypopod,
mesopod, and epipod.
(c) After these leaves follow leaves consisting solely of a hypo-
pod, which is much reduced in size (fig. 40, a). These
leaves are folded round, and serve as a protection to the
bud.

MODIFICATION OF THE ORGANS OF PLANTS 75

Lathyrus Aphaca. — The foliage leaf (fig. 4, leaves
4 and 5), which most nearly resembles the primi-
tive type,1 consists of two stipules and a petiole
which terminates in a point and is provided with a
pair of lateral leaflets.

Each individual plant is furnished with from one
to three of these foliage leaves.

Above them are leaves (fig. 41, leaf 6) consisting
of only a hypopod which forms two large stipules
that partially enclose a small point arising from
the epipod. The entire function of assimilation is
exercised by these stipules, and consequently both

(/) Finally, there are some scaly leaves which also consist

merely of a hypopod. These protect the tubers (fig.

40, a), and are practically devoid of chlorophyll. These

various leaves follow one another in the course of the

growth of the plant, in the inverse order to that in

which they are here described. As winter approaches,

some upright stems appear, the inflated ends of which

are stored with reserve nutrition (fig. 40, A). It is

round these tubers that the scaly leaves are to be found.

The bud which does not develop until the following

Spring is at the top, surrounded by folded leaves. The

ribbon-like leaves which grow under water and also those

broadened out at the ends make their appearance in the

course of the Summer. Next come the floating leaves

with slightly developed barbs, then the leaves which rise

well above the surface of the water, and finally the scaly

leaves which grow out of the flowering stalk.

1 The foliage leaves of Lathyrus Aphaca may be regarded as the

best representatives of the primitive leaf, for- in most species of

the genus Lathyrus, each leaf is furnished with leaves and with

one or more tendrils. L. Aphaca is probably specialized from

other species, for the majority of the leaves have lost their lateral

leaflets.

76

UNIVERSALITY OF DEGENERATIVE EVOLUTION

\

v_Qy

the petiole and the lateral leaflets have atrophied.

Higher up on the stem
(fig. 41, leaves 7 and 8)
are some leaves consisting
of two stipules and a non-
branching tendril. The
latter represents the meso-
pod and the epipod from
which the lateral leaflets
have disappeared. Below
the foliage leaves, there
are usually three leaves
which have considerably
degenerated (fig. 41, leaves
1 to 3). The hypopod
is represented by two
very small stipules, and
the epipod by the tiny
point in the centre. The
portion of the stem out
of which the basilar leaves
would grow does not,
under normal conditions,
emerge from the soil,
which accounts for the
way in which these leaves
have degenerated.

5. Nymphaea dentata.

foliage ieaves; 6, leaf composed Pprp rhp fnliacrp lpqvp<?

only of a pair of stipules. 7, 8, — nere une ionage ledVeb

terminal leaves turned into tendrils. are ^^ floating, fleshy,

Fig. 41. — Seedling Of Lalhynu

Aphaca.

1, 2, 3, very rudimentary leaves ; 4, 5

MODIFICATION OF THE ORGANS OF PLANTS

77

and toothed at the edges. During germination the
seed-leaves remain within the seed as with Lathyrus
Aphaca.

The first leaf is upright and reed-like in shape
(fig. 42, a 1). Its principal function is to pierce
the layer of soil which
covers the grain, thus
allowing the little ter-
minal bud to emerge
into the light. Com-
pared with the other
leaves, this needle-like
leaf seems to have con-
siderably degenerated.
The plant next pro-
duces a series of little
slender short-stemmed
leaves which grow
under the water. The
first of these leaves
are narrow and ribbon-
like (fig. 42, A 2); the
next are broader (fig.

4,9 T^ thpn fnllmv Vig. 42.— Vymphma denlata.

tL, HJ, Uieil lOllOW a, seedling with two leaves (1, 2); B, C,

more slender sub- leaves taktn fro,n 0,ae'' plants-
merged leaves, but with no lateral barbs (fig. 42, c) ;
then floating leaves, the edges of which are not
dentated, and last of all there are some leaves with
dentated edges. Owing to their submergence, the
basilar leaves of Nymphaea have greatly degener-

78 UNIVERSALITY OF DEGENERATIVE EVOLUTION

ated ; the blade is very thin, being composed of a
few layers of cells, and there are no stomata.

The basilar leaves and the apical leaves fre-
quently lose either partially or entirely the as-
similative function which they previously exercised.
Some of these leaves serve as a protection to the
buds of both leaves and flowers. Others are
adapted as protection from rain, and others have
undergone considerable modifications owing to their
existence either under water or under ground.

In all these instances of modification it can be
seen that some degeneration has invariably attended
each change which has taken place.

§ 5. — Modification of organs luhich are homologous
in individuals of different species.

Having investigated the modifications of leaves
in the individual, we will now give our exclusive
attention to the various modifications undergone
by the foliage leaves ; we shall see among the
various species we examine that degeneration has
played a part in each instance of modification.

Of these adaptations the following are the most
characteristic : adaptation to climbing, to a carni-
vorous diet, to aquatic life, to defence against ants,
against drought, and against herbivorous animals.

1. Adaptation to climbing. — Climbing plants
attach themselves by means of tendrils to the
nearest support within their reach. These tendrils,

MODIFICATION OF THE ORGANS OF PLANTS 79

which are thread-like and sensitive to contact, are

Fig. 43. — Cobaea scandens.

A, B, seedlings ; the leaves of the second end in tendrils ; C, young leaf of

adult plant.

modifications of either stems, leaves, or roots, the
latter being the most rare.
We will confine our atten-
tion at present to those
adapted from leaves.

In Cobaea scandens the
first two leaves of the
seedling usually terminate
in a leaflet (fig. 43, A) ;
the leaves which come
after are much finer and
are modified into tendrils
(fig. 43, c).

This modification of
leaflets into tendrils, which
entails the almost com-
plete disappearance of the original assimilative func-

o. 44. — Basal portion of a young
plant of Vicia Pyrenaica.

80 UNIVERSALITY OF DEGENERATIVE EVOLUTION

tion of the leaflet, is by no means uncommon. Vicia
PyiTimica (fig. 44) furnishes a good example of
this change, of which the degeneration of the blade
is the necessary consequence.

In Cucumis sativus (the cucumber) some of the
leaves are the shape of ordinary assimilative leaves,
while others are entirely modified into tendrils ; in
the case of the latter* the blade has completely
degenerated.

2. Adaptation to carnivorous nutrition. — Some
plants, instead of obtaining nutriment exclusively
from minerals and carbon dioxide, are capable of
assimilating animal matter from insects and other
small organisms, which they capture by means of a
special function exercised by the leaves.

(a) Utricularia. — In the aquatic Utricularia the
leaves are minutely sub-divided like many other
submerged plants. The leaves of Utricularia
vulgaris exhibit leaves of this type. Some of
their ramifications carry pouches, which serve for
the capture of minute organisms. These pouches
are formed by the modification of part of the
blade, and have almost completely lost their
chlorophyll.

Utricularia intermedia (fig. 45) exhibits two
kinds of branches ; the one kind stretches out
horizontally and carries green leaves ; the others
are devoid of chlorophyll, and the leaves are
purely carnivorous.

(b) Nepenthe. — The distal extremity of the leaf

MODIFICATION OF THE ORGANS OF PLANTS

terminates in a receptacle of complicated structure
which is adapted for the capture of carnivorous
nutriment. Degeneration has attended this change,
for the modified portion of the leaf is almost

ij/u.jlfn

Fig. 45. — Utricularia intermedia (after Goebel, Pflanzenbiologische Schilder-
ungen, vol. ii., p. 135).
The plant has grown from a winter bud of which the remains are visible to the
left. The horizontal, assimilating branch bears two pendulous twigs with
carnivorous urns.

entirely without chlorophyll. A carnivorous diet,
however, for these plants, is only supplementary to
the typical plant nutrition, and therefore the assimi-
lative function remains active.

F

82 UNIVERSALITY OF DEGENERATIVE EVOLUTION

Iii Utricalaria this function is exercised by the
leaves (as in U. intermedia), or by certain parts
of the leaves (U. vulgaris). In Nepenthe, however,
another and different assimilative organ is exhibited.
This consists of two lateral herbaceous growths, which
are not formed from the blade, but arise from the
petiole. In between the receptacle and the en-
larged portion of the petiole is a part in which
the petiole has assumed the function of a tendril.
While the receptacle is in process of formation, this
tendril twines itself round a support in order to
obtain the additional strength that will be required
later on to uphold the receptacle when full of diges-
tive juice.

(c) Drosera. — The leaves of the sundews {Drosera
rotundifolia, D. longifolia, etc.) are furnished with
a great number of emergences, each of which ter-
minates in a digestive gland. There is a large
drop of sticky fluid at the end of each gland,
and these drops, which sparkle in the sunlight,
have given the plant its name (Bos solis = sundew)
(fig. 46).

These emergences contain very little chlorophyll.

" A plant of Drosera, with the edges of its leaves curled
inwards so as to form a temporary stomach, with the glands
of the closely-inflected tentacles pouring forth their acid
secretion, which dissolves animal matter afterwards to be
absorbed, may be said to feed like an animal."

— Darwin, Insectivorous Plants, p. 18.

This adaptation to a carnivorous diet has not

MODIFICATION OF THE ORGANS OF PLANTS

83

come about without a certain amount of conse-
quent degeneration. The chlorophyll has dis-
appeared, excepting a small quantity exhibited in
the upper and under
surfaces of the blades
of the leaves, in the
flower stalks, in the
central tentacles, and
in the petioles.

3. Adaptation to
an aquatic life, — The
adaptation of leaves,
particularly of sub-
merged leaves, to an
aquatic life entails
important changes,
and these changes
are invariably at-
tended by degenera-
tion.

1. Owing to the
buoyancy of water,
submerged leaves are

not- rlpnpnrlpnt nn Fir" 4r'-— Leaf vi Drosem t,m<ji/olia.

nuu ueptuiueu u un Neai. the t0P) an insect is in the act of being

nnv hio-hlv flpvplrmPfl captured and the suckers are bending over
•my Illgiliy ueveiuptU jt Lower down is an insect reduced to a

organic support, and chitonous ske,eton-
the fibres and other thick-walled cells have con-
sequently degenerated to a great extent.

2. There being no transpiration in submerged
leaves and very little in floating leaves, the

84 UNIVERSALITY OF DEGENERATIVE EVOLUTION

former are very seldom provided with stomata,
and the latter only exhibit stomata on their
upper surfaces (see page 26, note on Ranunculus
and Stratiotcs). The conducting apparatus too, i.e.
the wood and the roots, has greatly diminished in
size owing to the diminution of transpiration.

3. Very little light penetrates to submerged
leaves, and oxygen is only partially soluble in
water. The blade has therefore become much
smaller. Very little importance, however, is at-
tached to this last modification, as the size of the
blade varies greatly in different specimens, whereas
the other modifications are similar in all.

(a) In the species already mentioned — Sagittaria
(p. 72) and Nymphaea (p. 76) — the first leaves
which are formed are submerged, with petioles either
short or entirely absent, and very slender blades.
In Vallisneria and many species of Potamogcton there
are only slender submerged leaves.

All these leaves augment their surface contact
with the water by ridding themselves of the deeper
layers of assimilative cells. Very little light could
penetrate to these cells, and any oxygen formed
would be difficult to get rid of.

(b) In other plants, especially in the Dicotyledons,
another kind of modification takes place for the
same end ; the blade is divided into very thin seg-
ments which in some cases present a hair-like
appearance.

The Ranunculus is a good example of this.

MODIFICATION OF THE ORGANS OF PLANTS 85

Besides the many species found on moors and culti-
vated land and in woods, there are other kinds
of Ranunculus — R. sceleratus, for instance — which
grow by the water-side. In this species it is not
uncommon to find floating leaves, and leaves which
wholly emerge from the water, growing on the same
stem. In other species, such as Ranunculus
hederaceus, there are seldom any but floating leaves.
Among the numerous varieties of Ranunculus aqua-
tilis there are some which, in addition to the floating
leaves, have also some leaves which are completely
submerged and deeply dentated ; in other varieties
of the same species there are only slightly dentated
leaves, while in Ranunculus fluitans the leaves are
all fringed.1

(c) In a submerged plant which is to be found in
Madagascar, Ouvirandra fenestralis (fig. 47), the
diminution in volume, compared to the surface, is
effected in a different manner. After the leaf has
developed in the ordinary way, all the parts of the
leaf in between the veins of the blade disappear.
This results in the blade being reduced to a delicate
piece of network, consisting entirely of communicat-
ing veins accompanied by adjacent assimilative cells.

4. Adaptation to defence against ants. — In the

1 Ranunculus sceleratus grows by the water-side ; R. hederaceus
is to be found on marshy ground, the internodes and petioles being
quite short ; B. aquatilis grows in shallow water, and the internodes
are of sufficient length to raise the flowers out of the water ; R.
fluitans grows in strongly-running streams, so that floating leaves
would be useless.

MODIFICATION OF THE ORGANS OF PLANTS 87

tropics, especially in South America, plants are
much exposed to the ravages of leaf-eating ants
(Atta). In order to protect themselves, they provide
other kinds of ants with shelter and nourishment,
and this necessarily entails several modifications.

Fig. 48. — Part of a leaf of Acacia sphaerocephala (after Schimper, Die Wechsel-
beziehungen zicischen Pflanztn nnd Ameiscn im (ropischen Amerika. Fase. 1
of Schimper' sbotanische Mittheilungenaus den Tropen. Jena, 1888).

In Acacia sphaerocephala (fig. 48), for instance, the
blades of the leaves are bipinnated and the proximal
leaflets terminate in small cavities filled with a nutri-
tive secretion. The stipules, which no longer contain
chlorophyll, are modified into hollow thorns in which

88 UNIVERSALITY OF DEGENERATIVE EVOLUTION

the ants live and obtain nutriment, not only at the

ends of the leaflets, but also in a thorny gland which

is situated upon the petiole.

5. Adaptation to drought.
— In places where rain
seldom falls, plants are
provided with natural
reservoirs of water. These
reservoirs are situated
either in the roots or the
stems and occasionally in
the leaves. Where the
leaves are fleshy, these
reservoir leaves are gener-
ally very simple in forma-
tion. Those of Sempervi-
vum and of several other

genera arise directly from the primordial, non-
differentiated leaf,

and there is no

differentiation into

hypopod, epipod

and petiole (fig.

49). See further

on (fig. 73), p. 236

(Sempervivum tec-

tOTllffi) ^\g. 50. — Branch of Caragana.

6. Adaptation to defence against herbivorous
animals. — Thorns are an adaptation which serve as
a protection against herbivorous animals. They

Fig. 49. — Growing point of Semper
vivum arachnoideum.

MODIFICATION OF THE ORGANS OF PLANTS

89

are derived from the modifications of various parts
of the plant, either the roots, stems, leaves, or
even of the floral stems.

This modification is always accompanied by some
degeneration ; the thorn, which is quite hard, ex-

TlQ.bX.—Mamillaria elephanlidens (after Lemaire? The figure is taken from
Goebel, Pflanzenbiologuche Schilderungen, vol. i., p. 71).

cepting at the point, is made up of thick-walled
cells from which the protoplasm has disappeared.

Leaf thorns arise from the modification of various
parts of the leaves — either of the stipules (as in
Caragana), the petioles (id.) (fig. 50), or of the

90 UNIVERSALITY OF DEGENERATIVE EVOLUTION

blade (as in Ilex). In each case the leaf par-
tially retains its assimilating function and its
chlorophyll.

In other plants, however, and especially in Cacti,
and the fleshy-leaved Euphorbias, the leaves exhibit
further evidences of degeneration, their function
being exclusively one of defence (fig. 51).

CHAPTER II

IN THE EVOLUTION OF INSTITUTIONS ALL MODIFICA-
TION IS NECESSARILY ACCOMPANIED BY
DEGENERATION

The distinction we have drawn between the
homodynamic organs of an organism, and the homo-
logous organs of organisms belonging to other
species, is not applicable in sociology, as we have
already pointed out in the introduction. Institu-
tions, however, may be regarded from two distinct
standpoints — from a statical point of view, as they
exist in the same society, and from the dynamical
point of view, as existing from epoch to epoch, and
from society to society. In both cases, we shall
arrive at the same conclusion as in biology, that
all modification entails degeneration.

In order to demonstrate this we will examine
in succession the modifications undergone by the
principal types of financial organizations now exist-

EVOLUTION ANT) DEGENERATION OF INSTITUTIONS 91

ing in Europe, and the most important stages in
the evolution of landed property amongst various
peoples.

§ 1. — Modifications of similar institutions in the
same society.

The financial organization of European towns
and states has undergone very important changes
since the middle ages. Taxes and duties have
attained a now universal importance, as substitution
for the revenue from crown lands, which constituted
the principal, if not the only, resources of the
sovereigns of the feudal ages.1

1 There are three stages in the evolution of financial systems in
countries (such as England, for instance) where the question of
finance has been most successfully dealt with.

1. The Feudal System, wherein the king had no separate revenue

apart from the nation, and wherein the revenue of the
sovereign was principally derived from crown land, the
cultivation and administration of which was carried on as
a source of private income to the king.

2. The Co-existence of the Feudal System and the Modern System,

viz., the disappearance of personally held land and its
attendant institutions, the development of the public pro-
perty of the State or Township, and the imposition of
duties and taxes.

3. The Modern System, viz., the complete separation of the

personal property of the sovereign from the property of
the nation, the increasing importance of taxes and duties,
and the almost complete disappearance of State and town
lands. Industries taken over by the State — such as rail-
ways, postal and telegraphic arrangements, etc. — and by
the towns — such as gas, water, etc. — constitute monopolies,
and are no longer subjected to the law of competition which
is always active in private business. —WAGNER.

92 UNIVERSALITY OF DEGENERATIVE EVOLUTION

Here, too, we find that all modification is
attended by degeneration. As a demonstration
of this, we will examine in succession — firstly,
the Communal budgets of Belgium ; secondly, the
State budgets of the States which compose the
German Empire ; thirdly, the budgets of Germany,
England, and France, regarded as belonging to one
group — the Western Kepublic of Auguste Comte.

I. The Communal Budgets of Belgium.

Collective property in its archaic form still exists,
in spite of considerable modifications in certain
parts of the Ardennes, of the Fagne, and of
Lower Luxembourg.1

At Wanlin, for instance, the collective property
constitutes nearly one-half of the land — 220 hec-
tares, of which 130 is arable land, and 90 is wood-
land. The arable land is divided into allotments,
apportioned among the heads of the various families
every eighteen years, for which they pay an annual
rent of 1 0 francs. There are no taxes or communal
duties, the revenue derived from this public property
being sufficient without them.

This economic and fiscal system, suitable to a
scattered and agricultural people, existed formerly
throughout the whole country, but the increase in
population and cultivation of the land have almost

1 Paul Errera, Les Masuirs. Historical and judicial researches
in the vestiges of the old territorial system of Belgium. Brussels,
Weissenbruch, 1891.

EVOLUTION AND DEGENERATION OF INSTITUTIONS 93

put an end to this archaic condition of things.
Private property has encroached upon collective
property, and taxes and duties are now substi-
tuted for the revenue which was formerly derived
from land alone.

This modification was of a progressive nature,
the extended requirements of the population de-
manding a corresponding augmentation of the com-
munal revenue. Degeneration, however, is exhibited
in the disappearance of the collective land of the
communes and of the attendant administrative
institutions.

The monographs of Paul Errera and the docu-
ments collected in 1875 by the Statistical Commis-
sion relating to the expenditure and receipts of the
communes, demonstrate the various stages of this
degenerative evolution : —

1. The vestiges of the old system may be

observed to decrease in number and im-
portance on approaching the North-West
from the South. The proportion of re-
ceipts from land compared with those
derived from taxes and duties is greatest
in Upper Belgium ; then follow those of
Condroz, Hainault, Brabant, Campine, and
finally, of the two Flanders.

2. The constitution of the two Flanders passed

through the following stages : —
(a) In certain localities round about Bruges the
old and the new system co-exist. Side by side, for

94 UNIVERSALITY OF DEGENERATIVE EVOLUTION

instance, with the communes of Oedelem, Beernem
and Oostcamp we find, though in a degenerate con-
dition, the old collective property of the aenborgers
of Beverhoutsveld, with its administrative college of
Veldheeren}

(b) In Sysseele, and in several other communes,
the assembly of Veldheeren has been abolished, and
the property which .they administrated has become
the property of the commune by whom the rents
are collected.

(c) At Gand, the grazing land of the Heemisse of
Saint-Bavon, which was declared collective property
by a decree of November 16, 1887, was afterwards

1 Beverhoutsveld is a vast domain situated outside the gates of
Bruges, in the territory of the commune of Oedelem. According to
the terms of a decision of the 13th August 1859, it constitutes a
collective property belonging to three sections of the communes of
Oedelem, of Beernem, and of Oostcamp. The family representatives
of these three sections are alone privileged to rent these lands,
and the revenue accruing from them is divided among the three
sections.

The old administrative authority, the college of Veldheeren,
which was in existence in the thirteenth century, still exists,
though in a new form. According to the terms of the decision
of 1859, the police are provided for from the taxes of Oedelem, as
the property is within that district ; "but in other matters, and
in the administration of the rights and revenues accruing from
it, the property shall be under the control of a commissioner
appointed, so far as possible, by the interested parties — i.e. the
aenborgers — in conformity with the laws and regulations relating
to the matter."

The Vrij-Geweid, also situated in the environs of Bruges, is
similarly administrated. (See Errera, des Masuirs, chaps, xix.,
xx. and following.)

EVOLUTION AND DEGENERATION OF INSTITUTIONS 95

transformed into private estates, and has been so
built over as to become almost unrecognizable.

(c/) In some towns the receipts from land only
represent a very small part of the communal revenue,
3 per cent, at Eoulers, O'l at Saint-Nicolas.1

It is thus evident that the evolution of com-
munal finance, which has been especially char-
acterized by the development of duties and taxes,
has been accompanied by the degeneration of the
old system of collective property.

II. Budget of the States of the German Empire.

The feudal system of finance has left its mark
upon Germany, especially in the eighteen subsidiary
states where archaic institutions have been best
preserved.'2 The principal stages in the progressive
evolution of the modern system, and the corre-
sponding degeneration of the feudal system may be
enumerated as follows : —

1. Mecklembourg-Strelitz still exhibits the primi-
tive system in a comparatively little altered
form. The Grand Duke is the sole adminis-
trator ; the budget is not separated from the
civil list; the revenue amounts to an average

1 Hector Denis, Vlmpdl, pp. 55. Brussels, Ve. Monnom, 1889.

2 Leroy-Beaulieu, TraiU de la science des Finances, I, 21, and
following.

Adolf Wagner, Lchr-und Handbuch der Politischcn Ockonomic ;
Finanzwisscnschaft. Erster Theil, sees. 214-216.

96 UNIVERSALITY OF DEGENERATIVE EVOLUTION

of seven millions, of which five are derived
from land.

2. The Duchy of Mecklembourg-Schwerin is ad-

ministered partly on the modern and partly
on the feudal system,' but the former pre-
dominates. There are two public budgets.
That of the Duke is of the greater im-
portance, and amounted to 14,500,000
marks in 1887, of which 7,000,000 were
derived from landed property.1 The second
budget is that of the municipal expenditure
(Gemeinsamen Finanzverwaltung), which is
defrayed by means of rates and taxes, the
receipts of which amounted in 1887 to
4,175,000 marks.

3. In the larger States of the Empire the feudal
system has completely disappeared, but the revenue
obtained from collective property forms an im-

portant part of the
Bavaria

public

income.

■ 1^3 %

Wur tern berg

. 13-2 %

Saxony
Baden

. 9-7%

Prussia

• 8-4 V

4. In the budget of the Empire, which is of
recent construction, naturally no traces of the feudal
system remain.

1 These lands comprise more than 99 German square miles with
over 206,000 inhabitants — about ,37 per cent, of the population.
L. Beaulieu, i., p. 43.

EVOLUTION AND DEGENERATION OF INSTITUTIONS 97

III. The Budgets of Germany, France, and England.

(a) Germany. — We have just seen that in the
principal States of the Empire there is still an
extensive amount of collective property. In
Prussia, for instance, besides forest land, there
are nearly 1,500,000 acres of arable land, an area
equal in extent to one of the smaller departments
of France.

(b) France. — Of the old collective property of
France only forest land remains, all grazing and
arable land having long since been alienated.

The budgets of modern times, however, still
exhibit traces of the old feudal system. The
State, for instance, up to the last few years,
continued to receive quit-rents for properties
under the old system.1 The budget shows an
annual decrease in these receipts, having fallen
from 100,000 francs in 1857 to 32 francs in
1869. In 1876, however, there was a rise to
2000 francs.

(c) England. — Here the decline of collective
property is still further exhibited. In the
Statistical Abstract published in 1877, the net
revenue from Crown lands for Great Britain and
Ireland figures at £40,000 net. Among the " mis-
cellaneous receipts" we find only £200,000 which
can be regarded as revenue derived from collec-
tive property, i.e. 15 millions in a budget

1 Leroy-Beaulieu, Traite de la science des finances, i., p. 35.
G

98 UNIVERSALITY OF DEGENERATIVE EVOLUTION

receipt of 1,956,000,000 francs. Later on, in
1884-1885, these revenues became rather larger
(principally owing to the purchase of the shares
in the Suez Canal which had belonged to the
Viceroy of Egypt), but the proportion remains
the same — 22 to 23 millions in a budget of
2 milliards 300 millions. We see, then, that in
all the financial systems of modern Europe the
progressive evolution of the modern system is
taking the place of the more or less rapidly-
degenerating feudal system.

§ 2. Modification of similar institutions in different
social groups.

According to the primitive constitution of things,
the land occupied by a tribe or clan was regarded
as res nullius, and consequently at the free disposal
of all the members of the community (the Feld-
Walt- und Weidegcmeinschaft of V. Maurer).

With the increase of population, the value of
land rose, and the state of things became modified,
the rights of groups and individuals becoming con-
solidated and at the same time limited. Then arose
gradually or simultaneously the following various
forms of landed property : — (1) Land held by
families ; (2) by villages ; (3) feudal property ;
(4) communal or public property ; (5) property
belonging to corporations ; (6) private property.

Family, village, and feudal property represent,

EVOLUTION AND DEGENERATION OF INSTITUTIONS 99

aniong certain peoples, three successive stages in
the evolution of property. When the old system
of land tenure was abolished, private and communal
or public property began to develop simultaneously.

While certain lands which were free to all the
inhabitants became transformed into collective
property, other such lands lost their public char-
acter and became private property. In the first
case, the communes, on being called upon to fulfil
functions of increasing complexity, proceeded to
transform all or part of the properties concerned
into patrimonial property or property for the use of
the people {communaux, allmcndwi)}

In the second case the property of the old
community became the joint but undivided property
of the members of the corporation ; when, however,
for purposes of cultivation it became necessary to
divide it, the corporative property became trans-
formed into private property.

1 Girou, Le droit administratif de la Bchjiquc, No. 688. "There
were three kinds of communal property :

"(«) Property directly appropriated to the use of the public such
as public squares, streets, churches, &c.

" (b) Communal property properly speaking — i.e. the real estate
and rights belonging to the tribune and to which the people
were entitled to a personal share. These consisted of the
forest land, rights of appanage, waste land, moorland, and
the rights of pasturing.

"(c) Patrimonial property, i.e. that held by the commune,
the revenue from which went to the commune to defray the
expenses of administration. It included timber land, arable
land, house property, market places, &c,"

100 UNIVERSALITY OF DEGENERATIVE EVOLUTION

On reviewing in succession these various phases
in the evolution of landed property, it will clearly
be seen that modification has everywhere been
attended by degeneration.

1. Family property {Montenegro). — Of all the
Balkan States, Montenegro — owing to the natural
barriers which separate it from the rest of Europe — -
has best preserved its archaic institutions. Here,
side by side with modern institutions, may be found
the old system of division into forty-two tribes
(pleme) which are sub-divided into clans or confrater-
nities (brastvo) and into patriarchal families (zad-
rugas and inokosnas).1 The development of modern
political and judicial institutions has, however, con-
siderably lessened the importance of the plemes
and the brastvos, so that progression in this direc-
tion has not been effected without accompanying
degeneration.

With regard to property, the two different forms
of family tenure have been substituted for what
was formerly the tribal or clan system. Of the
former collective property of the clan, there only
remain the following traces : —

1. Property rights held over certain portions of

land — generally forest or waste land.

2. The right of pre-emption in favour of mem-

bers of the brastvo or of those related to

1 For information about the common or differential characters
of the zadruga and the inokosna see Ardent, La Famille zougo-
slave cm MonMntyro. {R4forvie sociale, 17th October 1888.)

EVOLUTION AND DEGENERATION OF INSTITUTIONS 101

a member within the first six degrees of

lineal descent.1

3. The right of allotting to relatives their share

in the duties of helping widows and paupers

in their work. The workers in this case

receive no payment, neither have they any

right to demand maintenance.

Still rarer are vestiges of the collective property

of the pleme. A few portions of land, however, still

belong to that body, and it is probably a survival of

this ancient condition of things that foreigners are

1 Article 48 of the Civil Code of 1888, drawn up by Bogisic in
all possible accordance with "the excellent customs" of Monte-
negro, begins with the statement that "the right of pre-emption,
a privilege which has so long been enjoyed by the members of the
brastvo, by persons whose lands adjoin, and by the members of the
village and pleme, still flourishes, and will probably continue to
do so."

Bogisic adds that, in accordance with this right, "any person
desiring to sell his land, or any kind of real estate belonging to
him, is constrained, according to the established custom in such
cases, to first offer it in legal order to those persons who enjoy the
right of pre-emption, in order to give them an opportunity of pur-
chasing it at the price at which it is to be offered to the public.

Article 49, sec. 1, gives a list setting out the order of precedence
of those who enjoy the right of pre-emption.

1. Members of the brastvo within the first six lineal degrees of

descent.

„ t, . ,. . . , , 'N Transference, of recent

I. rersons owning adjoining lands. | . . ...

o rp> ,, , ,., .,, I origin, to neighbours

3. lhe other members of the village. J- „ . , . . . °

. ™ ,, , r , i i I of rights originally con-

4. I he other members of the pleme. J .. j , , °.

1 ' fined to relatives.

If none of those entitled to the first offer desire to purchase, the
owner may then sell his property to any other Montenegrin.

102 UNIVERSALITY OF DEGENERATIVE EVOLUTION

not permitted to acquire landed property in
Montenegro,1 and that the public are unrestricted
in the right to hunt over any ground they
choose.2

2. Village 'property {Russia). — Village communes
and the periodical division of land — the mcr of
Eussia or the clessah of Java, for instance — do
not represent types of a primitive system, but
are the outcome of a whole series of modifica-
tions. Kowalevsky traces the evolution of the
present system in Eussia through the following
principal stages : —

1. The joint use of land by the members of one

family group {rpechischc), corresponding to
the zadruga of the Southern Servians, and
sometimes comprised of more than forty
persons.

2. The division of the mother-family into

separate households, thus forming a village
community, and the temporary allotment
of the land of the community among the
separate families.

1 The old Montenegrin law relating to landed property, which
prescribes the purchase of land in Montenegro by any but
Montenegrins, is still in full force. No transaction in violation
of it is legally binding (Dickel, Etude sur le nouveau Code civil
tnonUnigrin).

2 Throughout the Southern Slavonic countries hunting is the
free right of all. Anyone may hunt where and how they please,
not only on public ground, among the mountains and forests, but
upon private property, whether cultivated or not (Dickel, p. 36).

EVOLUTION AND DEGENERATION OF INSTITUTIONS 103

3. The alienation of all or part of the allot-

ments assigned to the families and the
constitution of agricultural communes no
longer necessarily consisting exclusively
of persons akin.

4. The periodical division of land, which, as the

population increased, was instituted with a
view to re-establishing an equal distribu-
tion. This system of division, which was
established gradually, only dealt with the
more valuable sort of land, such as meadows
and arable land. The forest land and pas-
turage, that at least which was not already
annexed by the Lords of the Manor, was
free to all.
This transformation of family communities into
village communities was not effected without
accompanying degeneration. The administrative
institutions of the family group disappeared, and
the rights of pre-emption in favour of blood rela-
tions were gradually replaced by village rights.
The importance of the family, regarded as an
economic group, decreased in proportion with the
increase of the importance of the village.

In some places, however, and especially among
the Ossetes who inhabit the valleys of the
Caucasus, the old system may still be found.
There, at any rate up to within the last few
years, the aouls (villages) are principally com-
prised of families holding land in joint tenure,

104 UNIVERSALITY OF DEGENERATIVE EVOLUTION

frequently sharing all tilings in common. These
aouls are very rarely met with nowadays.1

Besides these family communities, there yet
remain among the Ossetes, as in Montenegro,
numerous vestiges of the primitive system of
clan property, i.e. the appropriation of certain
portions of land by the members of the clan,
the common use of pasturage and forest land, the
enforced participation in certain public works, and
the rights of heritage over unclaimed land, or un-
appropriated property which had become so owing
to the lapse of some " feu " or by the extinction of
a family community.2

3. Feudal property {England). — The introduction
of the feudal system into England resulted in the
substitution of a new method of grouping, in place
of the old agrarian communities. These new
groups, like the townships of earlier times, con-
sisted of a complete organization occupying definite
boundaries. Instead, however, of being a family
group administrated by a democratic organization
and government, this new system, which Sumncr-
Maine calls a " manorial group," consisted of
a tenantry autocratically grouped together and
governed by a feudal chief — the Lord or Seignior.3

1 See Kowalevsky, Coutume contemporaine ct loi ancienne, p. 42.
Paris, Larose, 1893.

2 See Kowalevsky, Coutume contemporaine, et loi ancienne, pp. 68
and following. Paris, Larose, 1893.

3 See Sumner-Maine, Les Communautds de village; 1 D., litudes
sur VHistoire du Droit; 1 D., VAncicn Droit ct la coutume ancienne.

EVOLUTION AND DEGENERATION OF INSTITUTIONS 105

Although it cannot definitely be asserted that
each one of these manorial groups was developed
from what had formerly been a village community,
yet it is evident that such was frequently the case,
and that the transformation was accompanied by
partial degeneration. For instance : —

1. The assembly of the inhabitants of the town-

ship, which formerly exercised complete con-
trol over all legal and administrative affairs,
disappeared, and in its place sprang up the
Manorial Court which was comprised of a
limited number of tenants and presided over
by the Lord of the Manor or his representa-
tive.

2. The collective property became absorbed, or

was at least considerably reduced by the
acquisitions of the Lord of the Manor, or by
divisions effected by members of the com-
munes. The old system of collective pro-
perty held by townships did not, however,
wholly disappear.
(a) The " rights of usage " in regard to waste
land, forest and moors (such as the use of unclaimed
pasturage, the cutting of timber, etc.), were still
enjoyed by the old inhabitants, and even in some
instances by other persons upon whom these "rights"
had been conferred.

Eniile Laveleyc, Lcs Communautes de famillc ct de village
{Eevuc a" economic politique, 1888, pp. 350 and following).
Vinogradolf, Villainage in England, Oxford and London, 1892.

106 UNIVERSALITY OF DEGENERATIVE EVOLUTION

(6) With regard to meadow land, sometimes the
Lord of the Manor put up enclosures for his own
benefit from Candlemas till Midsummer, the rights
of the community being established during the
remainder of the year only ; sometimes it was the
community who put up the enclosures, when the
Lord of the Manor was only entitled to the use of
the land during the intervals ; sometimes pasturage
was held as the joint property of the old community,
or rather of their descendants the tenants ; but as a
rule it was regarded as more or less common pro-
perty. The best meadow land was divided up into
what were termed " deals " and apportioned by
drawing lots.

(c) With regard to arable land, the method of
appropriating and cultivating the land occupied by
the tenants retained many traces of the village
system of collective property. For instance : the
enforced rotation of crops ; the periodical division
of land in certain parts of the country ; the
division of land into three breaks in other places ;
and the destruction after the harvest of the en-
closures surrounding the crops, after which the land
was used for the herding of cattle.

These survivals may yet be found in some districts
of modern England, in spite of all the great changes
in the English system of property ; changes such as
the disappearance of the serf and the appearance of
yeomen in the course of the thirteenth to the six-
teenth century, and the dispossession of the yeoman

EVOLUTION AND DEGENERATION OF INSTITUTIONS 107

in favour of the growth of large properties in the
course of the sixteenth to the eighteenth century.

4. Public or collective property (Switzerland). —
It is easy in Switzerland to follow the course of
the transformation into political communes of the
old system of economic communes, whether village
or manorial.

In the mountainous parts of Switzerland this
transformation is still incomplete, and side by
side with the modern commune may be seen
the old form of collective property, though in a
more or less advanced stage of degeneration.

The successive stages of this evolution may be
enumerated as follows : —

1 . The village communities (the Feld- Walt-und-

Wcidegcmeinschaft of von Maurer).

2. The collective property of the inhabitants,

whether feudal, free, or partly both
(Gemischte Gemeinde).
The Feldgemeinschaft completely disappeared
after the Eeformation, the collective land
of the community, the joint use of which
was the right of all the inhabitants, being
restricted to mere waste land, forest land,
and pasturage (Allmend).1

1 The Allmoui, in the primitive sense of the word, meant that
part of the old collective property held in joint tenancy by a
community of inhabitants or any other public body, the use of
which was limited to those who had a personal title to it. This
primitive meaning has changed in Switzerland — excepting in the

108 UNIVERSALITY OF DEGENERATIVE EVOLUTION

3. The institution of Biirgergemeinde — public

corporations — the members of which
enjoyed the sole use of the Allmend,
and upon whom fell the costs of local
administration ; as the population increased,
the conditions of admission into the Biirger-
gemeinde became more and more strict.

4. The creation of Einwolincrgemeinde, or political

communes, established side by side with the
old system, and in many localities eventu-
ally taking its place. In these the responsi-
bilities of the former system were assumed,
and part or the whole of the collective
property was appropriated. Where the
old communities still survive, they have,
as a rule, ceased to exercise the greater
part of their original functions. In the
Cantons of Berne and Saint-Gall, for
instance, the old communities have de-
livered up the greater part of their
possessions to the political communes to
provide for the expenses of general ad-
ministration. Their only now remaining
function is the administration of the small
remainder of their patrimony and the main-
tenance of the indigent members of the
community.

Canton of Schwyz — owing to the changes in the institution itself
(see Miaskowski, Die Schweitzer -ische Allmend in Hirer geschicht-
lichen Entiviekelung, von xiii. Jahrhundert bis zum Gegenwart).

EVOLUTION AND DEGENERATION OF INSTITUTIONS 109

Independently of this decay of the old system
of communities, the formation of political com-
munes was attended by other phenomena of
degeneration : —

1. The suppression of all or part of the " rights
of usage" enjoyed by the inhabitants.

The Einwohnergcmeindc, being called upon to dis-
charge more and more onerous and complex func-
tions, were obliged to either partially or wholly
transform the communal possessions, to the per-
sonal use of which the people were entitled, into
property appropriated to the use of the public,
either directly {i.e. into churches, teaching insti-
tutions, etc.) or indirectly, as a means of obtaining
a revenue {Brwerbsquelle).

2. The decrease in collective property.

Many of the Biii'gergemeinde, although no longer
discharging public functions, retained part of their
estates, which were held by the members in joint
tenancy. On the other hand, as the increased
population necessitated the cultivation of the
Allmcnd, the original " right of usage " re-
sulted in many instances in a transformation of
the land into private property.

This transformation, however, was not always
complete, and all the intermediate stages may be
traced between the old collective tenure and the
appropriation by individuals.

5. Corporative property {Belgium). — Here we will
limit the sphere of our observations to Belgium, in

110 UNIVERSALITY OF DEGENERATIVE EVOLUTION

order to avoid repetition, similar examples being
almost universally exhibited throughout Europe.1

Our information is obtained from the work of
Paul Errera, entitled Les Masuirs, recherchcs his-
toriqucs ct juridiqucs sur quelques vestiges des formes
ancienncs de la propria en Belgique.

The masuirs (the amborgcrs of Flanders) were
the mansuarii or mansoarii of the Merovingian
period, originally serfs, afterwards tenants and
copy-holders, and finally freemen.

Their history exhibits the following stages : —

1. The feudal epoch in which the masuirs — i.e.

all the members of the manorial group —
enjoyed " rights of usage " over all waste
lands, forests and pasturage adjoining their
holdings. These rights appear to have been
conferred by the Lord of the Manor, but
they really dated from a much earlier
• period.

2. As the increasing population necessitated the

regulation and limitation of these rights,
certain conditions of property and residence
were stipulated for in those seeking admis-
sion to the rights of the masuirs, and these
privileged persons organized themselves into
corporations which were more or less ex-
clusive and separate from the general
community.

1 With regard to corporative property in Switzerland, see von
Miaskowsjd, Die schiveizerische AUmencl, pp. 37 and following.

EVOLUTION AND DEGENERATION OF INSTITUTIONS 111

3. By degrees — by means of cantonments, pur-

chases, prescriptive claims, &c. — these cor-
porations absorbed the best part of the
land, and became almost independent of
the Lord of the Manor; as a rule, the latter
gave up half of the common territory to
them, and freed the surplus from all rights
of usage. In the corporations of masuirs,
however, there were still a few remaining
vestiges of some of the institutions of the
old manorial group from which they had
gradually developed. The Lord of the
Manor, for instance, himself being an in-
habitant and a masuir, had a right to a
share in the property of the community,
and further, in his seigniorial capacity,
certain privileges accrued to him such as
" la haute fleur des bois," i.e. tithes and
pannage (crops of acorns).

4. The Revolution put an end to all feudal

rights, and removed the last remaining
traces of the origin of the masuirs.
Throughout this long series of transformations, it
is evident that degeneration has followed in the
track of progress. Besides the disappearance of
the manorial group and its attendant institutions,
the rights of the masuirs may be said to have
become more restricted as they became more defined
and secure. In the early days, all the inhabitants
enjoyed joint rights over a vast common territory,

112 UNIVERSALITY OF DEGENERATIVE EVOLUTION

at the close of the old system this territory had
become much reduced in extent, and had become
the property of a more or less large group of
privileged persons.

6. Private property {Switzerland). — After the
Kevolution, the communities of masuirs and other
similar corporations ceased to have any recognized
legal existence. Those which still survived in
spite of the irregularity of their legal position,
owed their existence to their insignificance. The
others dispersed themselves, or were dispersed, and
the property which had belonged to them was
either incorporated with the communal estate, or
divided up among the members of the old com-
munity.

In each of these cases the transformation was
attended by degeneration, for the archaic adminis-
trative organization disappeared.

We saw in the allmend of Switzerland, this
same divergent evolution of public and private pro-
perty, part of the common land being transformed
into communal property, while the use of the sur-
plus ended in some instances in the land becoming
ultimately the private property of individuals.
This frequently occurred where land was cultivated
as orchards. In early times both fruit and fruit-
trees belonged, like the land itself, to the community,
and in certain parts of the Cantons of Uri and
Schwyz this is still the case. By degrees, however,
individual rights over fruit-trees planted on the

EVOLUTION AND DEGENERATION OF INSTITUTIONS 113

allmend came to be recognized. These rights,
whether temporary or held for a life-time, eventually
became perpetual, and finally this right to the
private acquisition of trees led to a right to acquire
the land itself. This last transformation was not
effected without a struggle and occasionally the
land was reclaimed by the community, the pro-
prietor of the trees receiving compensation. Now-
a-days the possession of trees and land usually go
together. Duality of this kind, however, is still to
be met with in certain localities. In the Sernfthal
(in the Canton of Glaris) a still stranger custom
prevails with regard to the maple forests. There,
the soil, the trees, and the fallen leaves (the latter
being used as litter for cattle) all belong to different
persons.1 With regard to house property there are
more intermediary conditions between use and
possession. In some villages, the chalets as well
as the ground upon which they are built, belong to
the whole community ; in other villages, both are
part of the collective property. Sometimes private
possession is restricted to the house or chalet, the
right to the ground upon which it is built lapsing
with the existence of the house. In order to limit
the number and durability of these buildings, many
restrictions are imposed, such as the prohibition to
build houses of stones, or chalets of wood cut from
trees not belonging to the builder himself or to the
corporation to which he belongs, etc.

1 Miaskowski, Die schweizcrischc Allmend, pp, 18 and following.
H

114 UNIVERSALITY OF DEGENERATIVE EVOLUTION

7. Summary. — This long series of modifications,
the result of which was the transformation of
primitive communities of goods into the modern
forms of public and private property, was ac-
companied throughout by degenerative changes.
The establishment of family property entailed
the curtailment of tribal and clan rights. Family
property passed into property held by the village ;
next the development of feudal tenure involved the
degeneration of the old agrarian communities ;
finally, the primitive organization of property with
the administrative and political institution depen-
dent on it, disintegrated and disappeared as the
primitive community of goods lapsed into the
personal enjoyment of these by individuals, and
as the primitive method of land tenure passed
into the rights of private property.

We see then that degeneration has always
accompanied evolution : the destruction of old
institutions is involved in the formation of new
institutions.

PART II

DEGENERATION IN THE EVOLUTION OF ORGANISMS
AND SOCIETIES

We have seen that modification of organs and of
institutions is always associated with partial de-
generation. We have now to show that, similarly,
when organisms and societies become modified, de-
generation is shown in some of their organs or
institutions. This shows again the universality of
degenerative evolution.

CHAPTEE I

ALL ORGANISMS EXHIBIT RUDIMENTARY ORGANS

All existing organisms have lost some organs in
the course of their phylogenetic development.

This may be proved in two ways : either there
are remaining vestiges of these organs, or else they
are to be found in other creatures which may be
regarded as ancestors.

116 UNIVERSALITY OF DEGENERATIVE EVOLUTION

1. Rudimentary organs, signs of a degenerative

transformation in the organism itself, are
either organs which have ceased to be
functional, or which have so diminished in
importance that their total disappearance
would be unattended by any appreciable
loss to the organism. In the majority of
cases this cessation of function is attended
by a corresponding structural decay.

2. The system of comparing living organisms

with their presumptive ancestors equally

demonstrates the retrogression of certain

organs.

Among the Orobanchacea? for instance, parasitic

plants derived from normal green plants, no trace

of cotyledons is observable from the period of

germination.1

Among animals, taking the horse as an example,
several organs have wholly disappeared. In the
genealogy of the horse, which is well known, the
earliest ancestor Eohippus, possessed five functional
fingers on the fore-feet, and four toes on the hind-
feet. The horse still possesses one functional finger
and one functional toe, two rudimentary fingers and
two rudimentary toes. Two fingers and two toes
have entirely disappeared.

It is hardly necessary to point out that this
system of comparison does not demonstrate the

1 L. Koch, Die EntioickelurigsgeschicMe dcr Orobanchen. Heidel-
berg, C. Winter, 1887.

RUDIMENTARY ORGANS OF ANIMALS 117

degenerative changes attending the phylogenetic
development of an organism with such incontestable
certainty as does the existence of vestiges of rudi-
mentary organs.

It is our belief that all organisms contain
vestiges of organs, either more or less apparent.
In our present condition of knowledge, however,
it is quite impossible — particularly as regards
plants — to prove this theory universally. It is
to be hoped, however, that future researches will
ultimately succeed in establishing it.

In the meantime, we will point out the most
typical among the cases known to us.

With animals, as with plants, our investigations
have extended not only to every kind of group, but"
to the most varied systems of organs, thus giving
our theory an extremely wide application.

Section I.

Rudimentary Organs of Animals.
§ 1. Rudimentary Organs in If an.

Throughout the whole human organic systems
signs of degeneration abound.

1. The Integumentary System. — In the ancestors
of man, the entire surface of the skin was covered
with hairs. Man's clothing of hair is far from
perfect, the hairs of which it is composed being
rudimentary.

1 1 8 UNIVERSALITY OF DEGENERATIVE EVOLUTION

According to Hertwig, the teeth should be
regarded as part of the tegumentary system, as
they really represent the scales of the skate,
situated within the buccal cavity.

In man, the last molar, or wisdom tooth, is a
rudimentary tooth. The small-sized shallow crown,
the diminished number of tubercles, the fusion of
the roots, the tardy appearance and occasional
absence altogether, are all indications of a rudi-
mentary condition.

2. The Skeleton. — With few exceptions, the
articular surfaces of the bodies of mammalian
vertebrates are covered in youth with bony
plates. These sometimes become very thick, and
are called terminal epiphyses. In some mammals
— the Sirenians, for instance — the terminal epi-
physes have disappeared. In man they still
exist, but in an advanced stage of degeneration.
In the lower vertebrates, such as the crocodile,
many more ribs are functional than in man.
In the crocodile all the ribs connected with the
cervical vertebrae are functional, whereas in man
they have degenerated. Of one entire section of
the human vertebral column — the tail — so fully
developed in the majority of other vertebrates,
only a vestige now remains.

Other rudimentary skeletal pieces are the lesser
horn of the hyoid bone, the stylo-hyoidean liga-
ment, the coracoid process, and the interclavicular
ligament.

RUDIMENTARY ORGANS OF ANIMALS 119

3. The Muscular System. — The cutaneous muscles,
those of the shell of the ear, and those that move
the tail, which in most mammals are well developed,
are still present in man, but have degenerated.

Further, there is to be found in man the
intra-acetabular part (the round ligament) of the
deep flexor muscle of the toes which is functional
in some animals — in young ostriches, for instance.
In the adult ostrich the intra-acetabular part is
separated from the rest of the muscle, which is
attached to the pelvis. Traces remain in the
horse of a connection between the intra- and the
extra-acetabular parts ; the muscle itself is divided
into two parts, the pectineal muscle in the thigh,
and the deep flexor muscle of the toe situated in
the leg. In the orang-outang, this degeneration
has made further advances than in man, the intra-
acetabular part of the muscle having entirely
disappeared.1

4. The Nervous System. — Here we find numerous
signs of degeneration, of which the following are
a few examples :

In the brain the pineal gland, the last remaining
vestige of what was formerly a functional eye, is
present.

In the spinal cord the filum terminate still
exists. We know that the spinal cord in man does
not retain its normal thickness to the extremity of

1 See Sutton, Ligaments, their nature and morphology. London,
1887.

120 UNIVERSALITY OF DEGENERATIVE EVOLUTION

the vertebral column but is arrested at the first
lumbar vertebra. There a considerable number of
special nerves leave it, forming a mass of branches
like a horse's tail. Along the centre of these
nerves, in the middle line, a slender filament
represents the spinal cord to the extremity of the
coccyx. This filament is the filum terminate, the
spinal cord in a condition of degeneration.

5. The Digestive System. — The csecum and its
vermiform appendage, are well known to be organs
which have degenerated.

6. The Vascular System.- — In quadrupeds the
intercostal veins are vertical, the blood consequently
flowing against gravity. These veins contain valves
which indirectly facilitate the upward and onward
flow of the blood by preventing it from running
back. Man, being a biped with a vertical thorax,
is provided with intercostal veins that are almost
horizontal. The ancestral valves being no longer
indispensable are in a condition of degeneration.

7. Sense Organs. — In the olfactory organ there
remains a degenerate Jacobson's organ. In the
organ of sight there is a third eyelid in a state
of degeneration. In the organ of hearing there
remains on the shell of the ear a kind of point
(Darwin's point) which is the last remaining vestige
of the ancestral elongated and pointed ear.

8. Genito-urinary System. — There is a whole
series of rudimentary organs in the genito-urinary
system of the higher animals. As is well known,

RUDIMENTARY ORGANS OF ANIMALS 121

the Wolffian body plays a considerable part in the
formation of the system. This body, the primitive
kidney, loses its urinary function at a certain
stage of embryonic development, and the permanent
kidney which gradually develops alongside, assumes
the urinary function. Later on, the Wolffian body
assumes new functions, becoming an important part
of the genital apparatus.

In this transformation partial degeneration occurs,
resulting in such reduced structures as the epi-
didymis, the organ of Rosemnuller, the vas dbevrans,
etc. (see fig. 57).

§ 2. Rudimentary organs in various groups.

1. Ccelenterates. — The Coelenterates comprise three
great groups.1 The Anthozoa, of which the coral is
a type, the Hydrazoa, which include fresh water
Hydra and the common jelly-fish of our seas, and
the Ctenophora, of which the chief representative
in our seas is Cydippe, a globular transparent animal
frequently to be found floating in large numbers on
the surface of the water.

The colonies of Anthozoa are usually composed
of individuals all exactly alike. In some species,
however, in the Pennatulida.' and the Alcyonaria
for instance, there is a distinct differentiation
amongst the numerous individuals composing the
colony. Side by side with sexual individuals

1 See C. Vogt and Emile Yung, T rait 4 d' anatomic compare, vol. i.

122 UNIVERSALITY OF DEGENERATIVE EVOLUTION

provided with tentacles and the eight mesenteric
folds, are other far simpler individuals : the zooides,
the function of which is respiratory and of which
the greater part of the organs have degenerated ;
the generative organs are lacking, the tentacles are
very small, and the mesenteric folds only number
two instead of eight. Degeneration, then, is

© O 7 3

exhibited side by side with specialization.

Among the Hydromedusce similar examples
abound. It is known that the polyp-like or
medusa-like forms of this group which may live
independently, frequently associate themselves
together to form colonies, sometimes predominantly
polypoid, sometimes completely medusoi'd, and occa-
sionally a mixture of the two.

In these cases a marked polymorphism is often
apparent. The different individuals become adapted
to definite functions, and the corresponding organs
undergo special development ; the other parts of
the body having become either unnecessary or
merely accessory, begin to degenerate and finally
disappear. Thus we see in Hydro'id colonies, not
only the hydra-like members, nutritive, fixed and
sterile, and the medusa-like members which are
reproductive and become free from the colony, but
also certain individuals which are termed gono-
phores. These gonophores are really medusa-like
members which have lost their independent move-
ment, and have consequently more or less lost
both their tentacles and their umbrella-like discs,

RUDIMENTARY ORGANS OF ANIMALS

123

i.e. their organs of locomotion. In some colonies
of Hydroi'ds, polymorphism has made such advances
that there are tactile individuals of which the
digestive tube lacks both mouth and tentacles, and
other purely defensive individuals of which the
internal organs are almost all in a state of atrophy.

Opinions differ regarding
the complicated question of
the structure of the Siphon-
ophora. The organism (fig.
52) may be regarded as a
simple medusa of which
the different appendages —
the pneumatophore (pn.),
the swimming bells (cl.),
the siphons (s.), the shield
(&.), the tentacle (t.), the
palp (pa.), the gonophores
(go.), etc. — constitute the
organs, or as a colony each
part of which is represented
by an individual polypoid
adapted to fulfil a special
function. Whatever theory
is accepted, it is clear
that a whole series of parts of the creature must lie
regarded as organs in a condition of degeneration.

We accept the second of these theories,1 and

Fio. 62. — Diagram of the structure
of one of the Siphonophora.

pn, pneumatophore or float ; cl,
swimming bell; b, protective
polyp ; t, tentacle ; pa, palp; g,
Ronophore; p, peduncle bearing
the mouth; s, individuals for nu-
trition.

1 Haeckel, System dec Meduscn: Jena, 1880-1881.
TraiU d'anatomic comparer.

A. Lantr.

124 UNIVERSALITY OF DEGENERATIVE EVOLUTION

regard the whole creature as formed from a
Craspedote Medusa which has become mother of a
colony and of which the umbrella, developed into
the pneumatophore (jpn.) has had its radial canals
greatly simplified and its tentacles reduced to one
during the growth of the colony. The stalk-like
stomach (p.) of this medusa has increased in length,
but this development is attended by corresponding-
degeneration, the buccal aperture, which is situated
at the free end of the peduncle in normal
Craspedote Medusae, having entirely disappeared.
The stalk, formed in this way, serves as a support
to the number of other individuals of which the
colony is constituted, and which are remarkable
for the great morphological variation they exhibit.
Among these individuals, those at the top, i.e.
those nearest to the air-sac, fulfil the function of
locomotion. They become transformed into swim-
ming bells (d.) and contain no organs whatever.
Below these locomotory organs are the sexual
individuals, or gonophores (#.), and the sterile
individuals (s.). The former are of meduso'id
structure, the umbrella is more or less perfect, and
they are sometimes provided with tentacles, and
possess a peduncle or manubrium which some-
times has a buccal aperture. The sterile individuals
provide nutrition for the others. The organs no
longer essential to them atrophy in a variable
degree. In the case of some the umbrella is
present, in others it is absent, and between these

RUDIMENTARY ORGANS OF ANIMALS

125

two extreme types come intermediate types which
exhibit every possible stage of degeneration.

In the Ctenophore group development and de-
generation are exhibited simultaneously in the
organs of locomotion. The fundamental and typical
shape of the Ctenophore is round or oval, and the
eight sides are provided with swimming plates,
originally uniform — as in Bcro'e.
The individuals belonging to
this group exhibit important
evidences of modification in
their external morphology. The
body being sometimes com-
pressed in various directions,
the shape is altered from the
original, and assumes a more
or less irregular appearance.
The organs of locomotion
undergo a corresponding
change. Take for example
an adult specimen of Bolina
norvegica (fig. 53); the body is lobate, although it
was round during the larval period ; the swimming
plates are not uniform, four being long and reaching
along the whole length of the body, the other four
being only developed in the upper half of the body
as far as where the lobes are inserted, where they
end as degenerate hair-like processes. By referring
to the Cestidte, which are ribbon-like in shape, it
will be seen that by means of compression the body

Fig. 53. — Bolina Norvegica,

seen from the broad side.
C, short rows of swimming
plates; A, long rows; L,
lobes. (After Vogt and
Yung, Traite d'anatomie
comparie.)

126 UNIVERSALITY OF DEGENERATIVE EVOLUTION

of the animal is lengthened out into the shape of a
narrow ribbon. Of the primitive lateral row of
plates, four are represented by mere vestiges, and
the other four, which continue to be functional, are
situated, closely coupled together, on the two edges
of the animal.

2. Worms. — We will next take in succession the
Plathelminthes or flat worms, the Rotifers, the
Nemathelminthes, or round worms, the Annelids
and the Gephyreans.

Among the Flathclminthes, the group of Cestodes
contains the common tape-worm of man (Tccnia
solium). In the course of its parasitic existence
this worm has undergone considerable morphological
changes. The digestive tube is lacking, and the
whole nervous system has become greatly simplified.
The degeneration of the nervous system is not,
however, complete, for important vestiges still per-
sist. The degeneration of the digestive tube is
much more thorough. In Tmnia solium it is alto-
gether absent. In species closely allied to the Taenia
some slight vestiges of the digestive apparatus yet
remain. The head or scolex of some species of
Tetrarhynchus contain glandular cells which have
been homologized with the salivary glands of other
flat worms (Trematodcs). In other species of Tetra-
rhynchus there is a rudimentary organ which repre-
sents the oval sucker of the Trematodcs, and in
Anthroccphalus clongatus the orifices of the salivary
glands are in the region of this vestige of the
digestive tube.

RUDIMENTARY ORGANS OF ANIMALS 127

Many facts in support of our argument may be
drawn from a study of the excretory organs of the
Cestodes.

The condition of these organs in Caryophylleus
mutabilis best represents the primitive condition.
Within the body of this worm are a large number
of narrow ducts with ciliated funnels communicating
with the spaces in the parenchyma (Fraipont).
These organs communicate with canals which
gradually reunite and anastomose to find a vent in
one single aperture, the foramen caudate, which is
situated in the posterior part of the body where
there is a bladder. In the Cestodes, however, where
the body is very long, the action of the bladder
is insufficient to secure a complete evacuation.
Secondary apertures are therefore formed at in-
tervals along the main ducts. This new structure
entails the degeneration of the terminal bladder
which has become superfluous. In Botryoccphalus
punctatus, which possesses a great number of
excretory apertures, the primitive evacuatory appa-
ratus— i.e. the contractile cavity — has completely
disappeared.

Rotifers are minute animals, usually living in
fresh water, a few being marine. One of them, an
inhabitant of damp earth or moss, has been supposed
to possess the power of revivifying after complete
dessication. At the anterior end of the body, a
rotifer possesses a complicated ciliary apparatus
which fulfils the function of locomotion, and from

128 UNIVERSALITY OF DEGENERATIVE EVOLUTION

the rotatory movements of which the name of the
group have been given. This organ is well developed
in those types which lead an independent existence,
but in those where movement is more restricted or
where the character of the organ has changed, it is
considerably modified and reduced in size. In
Philodince (crawling Eotifers) the organ of rota-
tion has lost the central part, and in its place is
substituted a very complicated organ of prehension.

In sessile forms such as Floscularia, the organ is
modified and only the primitive character of the
inner ring is retained, while the outer ring is
segmented and becomes a series of arms or lobes,
furnished with stiff' bristles. In Apsilus, another
sessile form, the organ of locomotion has disap-
peared ; this is obviously an instance of true retro-
gression, for in young specimens a vibratory crown
still persists.

The Nemathelminthcs contain such round worms
as these common intestinal parasites : Ascaris,
Oxyuris, Strongylus, etc. These all belong to
the Nematode group, and possess a complete
digestive tube. Gordius, however, exhibits organs
which are reduced in a marked degree ; in the
adult animal the buccal orifice of the digestive
tube is closed, and the posterior part of the
intestine has disappeared, although in the young
worm the alimentary canal is complete. This
modification, though incomplete and appearing
only in the adult life of Gordius, is complete

RUDIMENTARY ORGANS OF ANIMALS 129

and permanent in some other Nematodes. In
Echinorhynchus, for instance, the digestive tube
is absent, and nourishment is obtained by means
of osmotic soaking through the body walls.

The Annelids comprise the annulated sea-worms
and forms like the common earth-worm (Lumbricus
terricola). In these creatures we will take, first,
the development of the eyes. In Oligochcctes, which
for the most part live in soil or mud, the organs of
sight are greatly reduced. The Naulomorplice alone
have eyes. The Archiannelida — Histvior (a para-
site), for instance — possess eyes when young, but
in the adult state the eyes have greatly degener-
ated. As a rule, the eyes of the Polychcetcs are
well developed, and in some of them quite re-
markably so. In species, however, which do not
move about much, the eyes are merely represented
by small pigmented spots.

We may mention, too, the Gephyreans, without
pledging ourselves as to their exact relationship.
Bonellia viridis, the history of which is well known
and of great interest, belongs to this group. The
male Bonellia lives as a parasite on the proboscis, or
in the gullet or the nephridium of the female. It
is flat and small, and has neither mouth, arms, nor
circulatory system. All the organs remain as in
the larval condition, with the exception of the
genital organs, which are fully developed.

Bonellia and Dinophilus, a rotifer, of which the
male is degenerate, exhibit a progressive degenera-

I

130 UNIVERSALITY OF DEGENERATIVE EVOLUTION

tion of all the organs not connected with reproduc-
tion. Degeneration has made furthest advances in
Bonellia, which affords a striking example of retro-
gression side by side with development.

Investigations of animal series such as these
might well be continued throughout the various
classes and groups, showing the existence of rudi-
mentary organs in all. We will restrict ourselves
here, however, to mentioning the larger subdivisions
only, taking one example from each group.

3. The Echinoderms. — This order comprises
star-fish or asterids, sea-urchins, Crinoi'ds and
Holothuria or trepangs.

Of these we will take the star-fish, and proceed
to examine its digestive tube. Under normal
circumstances, the intestine terminates in a dorsal
anus, centrally, or slightly excentrically placed, pre-
ceded by a very short but well-developed rectum.
Asteracanthion and Solaster furnish good examples
of this. In some kinds of Asterids — in Astropecten
aurantiacus, for instance — the anus no longer exists,
and the rectum, having become useless, is greatly
reduced, though still exhibiting signs of its original
condition.

4. Mollasca. — Of the group of Mollusca we
will take the Gastropods and the Cephalopods as
examples.

There are two kinds of Gastropods, straight
and twisted, the former representing the primitive
type. The straight types — such as Chiton, Patella,

RUDIMENTARY ORGANS OF ANIMALS

131

Haliotes, and Fissurella — are bilaterally symmetri-
cal, while in other Gastropods the spiral twisting
of the body causes a progressive diminution of the
organs situated on the side towards which the
twisting occurs ; the organs of the left side may
therefore become smaller, and finally atrophy
almost completely.

The internal shell of the Cephalopods furnishes
a striking example of
a rudimentary organ.1
The Nautilus (fig. 54)
has a shell, the spiral
coils of which are
pressed tightly against
one another. The spiral
is divided into a series
of chambers by means
of partitions, each par-
tition being provided
with an aperture for

t-ViP irlrm'sQi'nn nf thp '. telmi»al chamber of the shell ; a, body
Lilt! clUllllbMUIl Ul Uie of the animal ;/, siphon; r, mantle fold.

siphon (I.). The shell W*0—*
of Spirilla (fig. 55 and 56, a), a creature still existing,
is only partially curled round ; the last chamber of
the shell is very small, and only encloses a part of
the animal, most of which remains outside the shell,
and partially covers it by the mantle (p.), the
shell being therefore partly external and partly
internal. On examining the fossil species : Spiru-

1 W. Boas, Lchrbuch der Zuolvgie. Jena, 1894.

132 UNIVERSALITY OF DEGENERATIVE EVOLUTION

lirostra (fig. 56, b), Belemnites (fig. 56, c), and
Conoteuthis (fig. 56, d), a progressive simplifi-
cation of the shell may be observed, the latter
becoming less and less coiled, until finally the
original shell is transformed into a straight
chambered portion surmounted by a stiliform
process. Degeneration is principally exhibited in

Fig. 55. — Spirula
prototypos.
a, body of the animal;
p, mantle fold ; c,
shell, partly internal,
partly external.
(After Owen.)

Fig. 56 — Shel's of various Cephalopoda.
A, Spirula; B, Spirulirostra ; C, Belemnites ; D, Cono-
teuthis; E, Ommatostrephes; F, Loligopsis. (After
Boas.)

the segmented part, which becomes more and more
reduced. In Ommatostrephes (fig. 56, e), and in
all existing species (such as Loligopsis, fig. 56, f),
with the exception of those just mentioned above,
degeneration has become complete. The shell no
longer contains a cavity for the reception of the
animal, and the phenomenon already mentioned
with regard to Spirula — the development of the

RUDIMENTARY ORGANS OF ANIMALS 133

organism round about the shell — has become
more marked in character. The shell has
become internal instead of external, and forms
the so-called cuttle-bone. This structure, being
only the vestige of what was originally an
external shell, must be regarded as a reduced
organ.

5. Arthropoda. — The group of Arthropods com-
prises the Myriopoda, the Crustacea, the Arachnida,
and the Insecta.

Instances of rudimentary organs are very common
among the Crustacea, but our investigations with
regard to the appendages of the cray-fish were so
thorough that we will give examples from another
group, that of the Insecta, instead.

Insects are characterized by the possession of
three pairs of legs and two pairs of wings. The
organs of flight exhibit a multitude of special
adaptations, and numerous instances of degenera-
tion are exhibited. In the Ncuroptera (dragon
flies), Ifymenoptera (saw flies and bees), and Lepi-
doptera (moths and butterflies), the four wings are
generally all alike and fully developed. In the
Diptcra (flies), the posterior wings have atrophied
and are represented by two reduced organs, the
" balancers," these being absent in certain types.
The Strcpsiptera form a group of which compara-
tively little is yet known; the larvae live in the nests
of bees ; the females have no wings ; in the males
the anterior wings are rudimentary and the pos-

134 UNIVERSALITY OF DEGENERATIVE EVOLUTION

terior wings are fully developed. In the group of
Coleoptera (scarabs, cockchafers, longicorns, and
beetles), the anterior wings (" wing covers ") have
become very resistant, and constitute a perfect
cuirass covering and protecting the abdomen, and
the posterior wings, the organs of flight, are folded
beneath the wing covers when in repose. In the
Scaraban and the Calosoma the posterior wings
are much reduced. In other Coleoptera the wing
covers are united by their inner edges, thus render-
ing all movement of the underlying wings ineffectual
and useless. The result may, as in Gibbium,
be the total atrophy of these under wings. In
the Staphylinidre, the upper wall of the abdomen
is so strong that the protection of wing covers is
unnecessary ; consequently the wing covers have
degenerated into little lamella which cover only
the anterior quarter of the body. In the female
Lampyris (glow-worm), the wings have totally dis-
appeared. The Orthoptera (cockroaches and ear-
wigs) exhibit a great number of variations in the
organs of flight. In cockroaches all four wings
are usually fully developed ; in some specimens,
however, and these principally female, a very pro-
nounced degeneration of all the wings may be
observed. The anterior wings of the Forficulides
are reduced. In the group of Phasmidae we find
side by side with species of the genus Bacillus,
which have no wings and look like dried twigs,
Phyllium siccifolium to which the large green and

RUDIMENTARY ORGANS OF ANIMALS 135

yellow wings give the appearance of a leaf. The
neuters of Termites have no wings.

6. Vertebrates. — Among Vertebrates the genito-
urinary system contains a great number of rudi-
mentary organs. In order to fully understand the
nature of these organs, it will be necessary to
glance through the ontogenetic and phylogenetic
development of the system.

The first stage in the formation of the kidney-
system is the 'pronephros (fig. 57, a). This primitive
organ consists of intricate canals (a, a, a), opening
into the body cavity at the point where the glomeruli
are formed on the sub-intestinal vein. All these
canals originally had apertures to the exterior.
Later on, however, these uriniferous tubules became
connected with one single excretory canal (c.) opening
into the cloaca (ce.). The primitive genital gland was
situated close to the pronephros. In process of time
the mesonephros replaced the pronephros (fig. 57,b);
in origin it was quite distinct from the pronephros,
its appearance being that of a secretory urinary
gland (g.) and its secretory canal (c.) (segmental
duct), was the same as that of the pronephros.
The urinary system thus formed, became and still
continues to be, closely connected with the genital
gland, the discharging canals of which passed
through the mesonephric kidney in order to find
a passage to the exterior through the segmental
duct. During the mesonephric stage, another canal
was formed which started from the cloaca and

136 UNIVERSALITY OF DEGENERATIVE EVOLUTION

opened out into the general body cavity, this was
Mailer's duct (m.).

Fig. 57. — Development of the Urino-genital system in higher Vertebrates.

A. Pronephric stage, a, tubules; c, excreting duct; ce, cloaca.

B. Mesonephric stage. G, mesonephros; r, remains of pronephros ; c, excreling
canal; tc, neutral genital gland ; M, M uller's duct ; ce, cloaca,

C. Metanephric or adult stige, in the male. R, permanent kidney; U, ureter;
V, bladder; T, testis; e and v, epididymis and vas deferens ; s,vas deferens;
hm, hydatid of Morgagni; h, hydatid;/), paradidymis; um, uterus masculinus.

D. Metanephric or adult stage in the female. R, permanent kidney; U, ureter ;
V, bladder; Ov ovary; p and p', parovarium and paraophoron: w, Weber's
organ ; V, vagina; u, uterus; l, aperture of Fallopian tube; A, hydatid.

The mesonephros, however, was not the perma-
nent kidney. In the course of time the metanephros,
the permanent excretory gland, was developed (fig.

RUDIMENTARY ORGANS OF ANIMALS 137

57, c and d). This development was attended by
important modifications, further instances of de-
generation taking place, and fresh organic connec-
tions being established. An examination of the
male and female sexes is necessary in order to
explain what really took place.

In the male (fig. 57, c) the mesonephros began
to atrophy ; that part which was connected with
the testes was transformed into the epididymis and
the vas deferens (e. and v.) of the true male genital
organs; the remaining part atrophied, and when the
permanent organization was attained, only persisted
in the form of a paradidymis (p.) and a hydatid
{h.), organs which are quite without function in the
adult state.

The discharging canal which, during the mesone-
phric stage was common to both urinary and genital
glands, remained simply in connection with the
testes, and then became the vas deferens (s.) of
which the cloaca having disappeared, the terminal
extremity became gradually individualized. The
permanent kidney (r.) became connected with a
fresh canal — the ureter (u.) which was formed by
degrees at the expense of the primitive discharging
canal, and subsequently became separated from the
latter in order to empty itself into the bladder (v.).

These changes were attended by a remarkable
evolution of Midler's canal, which first increased
in size, and then at a certain point proceeded to
atrophy until all that remained were the distal and

138 UNIVERSALITY OF DEGENERATIVE EVOLUTION

proximal extremities in the shape of reduced organs,
the hydatid of Morgagni (hm), and the uterus
masculinus (uni), neither of these being functional.
The intervening part of the canal remained, and
formed a canal which has been described by Gasser.

We see then, that in the genito-urinary apparatus
of an adult male there are :

(1) Organs which have come into existence at
different times, but which have retained their
original functions, viz. : the testes, the kidney
(metanephros), and the ureter; (2) Organs which
are functional, but of which the ultimate function
differs from the original, viz. : the epididymis and
the vas deferens ; (3) Reduced organs, vestiges of
what were formerly active organs, viz. : the hydatid
and the paradidymis ; (4) Reduced organs, vestiges
of Miillerian canal which only became active in
the female, viz. : the hydatid of Morgagni and the
male uterus.

In the female (fig. 5 7, d), the development of
the renal part is similar to that we have just
described in the male. Taking first that part of
the mesonephros which became connected with the
genital gland and the corresponding discharging
canal, we find that the canal disappeared with but
rare exceptions in which it formed Gartner's duct (#.),
the lower part persisting in the form of a rudiment
(Weber's organ (w.)) ; the upper part became reduced
to a small tissue which surrounded the paraovarium
(p.), and the paraophoron (p.) vestiges of what was

RUDIMENTARY ORGANS OF ANIMALS 139

formerly the mesonephros. Miiller's canal became
considerably enlarged ; it formed the vagina, the
uterus, and the Fallopian tubes ; at the upper end
it was connected with the hydatid, a vestige of the
mesonephros.

It is plain then that the genital-urinary apparatus
of the female comprises some organs of which the
functions remain unchanged : the ovaries, the per-
manent kidney, the Fallopian tubes, the uterus, the
vagina, and the ureter, and some rudimentary organs,
vestiges of what were once active organs ; the
paraovarium, the paraophoron, hydatid and Weber's
organ.

The complicated development of this system
becomes clear if a careful study is made of the
history of the genito-urinary apparatus of the entire
series of vertebrates.

It appears that the various phases through which
the embryos of the higher vertebrates pass are
stages similar to those which may be observed in
the adult lower vertebrates.

The principle of recapitulation, that the embry-
onic stages of higher animals recapitulate successive
stages attained by the adults of lower animals,
receives a full corroboration from the facts we have
been displaying.

Amphioxus, for instance, remains still at the
pronephric stage : fish as a rule have a mesonephric
or permanent kidney. Some lizards (Lacerta) up
to the age of two years make use of the mesone-

140 UNIVERSALITY OF DEGENERATIVE EVOLUTION

phros as the organ for eliminating urine, but, at
the same time they make use of the metanephros
which is also functional.

In Chamccles the mesonephros remains partially
active throughout life. Both birds and mammals
completely lose the mesonephros, and in the adult
stage the metanephros is the only active kidney.

This is not the place in which to complete our
study of the recapitulation theory, and we shall
have to recur to it later on ; but it was impossible
to describe the numerous rudimentary organs of
this system without taking a comprehensive glance
at the individual and specific development of the
whole. This investigation, moreover, raises another
question.

It has just been sbown that the epididymis is
only a vestige of the mesonephros, but in this case
it cannot lie said that there has been degeneration ;
what has happened is that an organ has been trans-
formed, and that one function has been replaced by
another. According to some authorities the supra-
renal capsule, an organ of unknown, but doubtless
essential function, is the result of the transformation
of the pronephros. If this theory be ultimately
established, it will furnish a second example of what
we have stated above. The thyroid gland may
and ought to be investigated from this standpoint.
The various component parts of this organ had
no original connection. In higher vertebrates only
the central part of the organ appears to be similar

RUDIMENTARY ORGANS OF ANIMALS 141

to the same organ in the whole series of vertebrates.
In man the mesial part of the thyroid gland is
reduced, but it cannot correctly be said that the
organ itself is degenerating. The contrary may
even be asserted, for all we know upon the subject
goes to prove that in mammals the thyroid gland
is formed and established at the expense of the
primitive rudimentary organ of which all the
morphological and embryological connections are
changed. This secondary development, entailing
the loss of a reduced ancestral organ, is attended
by a functional modification of great importance.
The primitive function probably discharged by the
gland has given place to another and rather vague
function, but one which is connected with the
breaking down of toxic matter formed by living
tissues.

An examination of the vertebrae of vertebrates
will show the existence of rudimentary organs
throughout the whole group, each type and each
individual among the vertebrates exhibiting special
degeneration. In order to briefly demonstrate this
point, take two quite different types of which all
the vertebrae are well known and can therefore
be examined without difficulty : man and the
frog.

We know that the construction of the primitive
bony vertebra was as follows : — a centrum, carrying
neurapophyses, an intercentrum, carrying hsema-
pophyses, and a pair of unforked ribs.

142 UNIVERSALITY OF DEGENERATIVE EVOLUTION

Each phase in the evolution of this primitive
vertebra has been attended by degeneration.
In man the vertebral column consists of : —

(a) Seven cervical vertebrae.

(b) Three dorsal vertebrae.

(c) Five lumbar vertebrae.

(d) Three sacral vertebrae.
(<?) Six coccygeal vertebrae.

Each one of these vertebrae exhibits important
modifications, and shows signs of degeneration.
The proatlas is represented by its intercentrum only.
The atlas consists of a centrum and neurapophyses,
but there is no zygapophyses ; it possesses one pair
of ribs and hamiapophyses in a reduced condition.
The axis consists of the same elements but carries
postzygapophyses.

The four following cervical vertebrae consist of
the same elements, but carry both zygapophyses
and postzygapophyses. In man all the cervical
vertebrae, with the exception of the first (the
proatlas) have lost the intercentrum. The next
vertebrae which is generally regarded as the seventh
cervical vertebra, consists of the same elements, but
it ought to be regarded as the first dorsal vertebra.
The vertebral artery and the sympathetic nerve
trunks accompanying it, do not pass through the
vertebral canals ; in some cases these do not exist.
The so-called seventh cervical vertebra has occasion-
ally one fully developed rib which articulates with
the sternum like the ribs of true dorsal vertebrae.

RUDIMENTARY ORGANS OF ANIMALS 143

There are thirteen dorsal vertebrae. With the
exception of that which has just been described,
they consist of : — a centrum, neurapophyses, and a
pair of fully developed ribs ; the intercentra has
completely disappeared, and the haemapophyses,
which form the head and neck of the rib, are in a
reduced condition. In the five lumbar vertebrae
which follow, the rib disappears, or to speak more
accurately, the transverse processes are all that
remain of what were the ribs, and have ossified
with the vertebra.

The sacrum is a region profoundly modified to
support the pelvic basin. It is formed by the
fusion of five vertebrae, each consisting of a
centrum, neurapophyses, and short bicipital ribs.
The first three vertebrae are the true sacrals, as
these alone support the basin ; the two following are
really caudal vertebrae in process of fusion with the
sacrum. In monkeys other than anthropoids there
are really only three sacral vertebrae and these are
at once succeeded by the tail. In man, the tail1
consists of six vertebrae of which the two first —
which consist of a centrum and neurapophyses and
of bicipital ribs — have fused with the sacrum,
while the four remaining lower vertebrae, which
consist of only the centra (the first still exhibits
rudimentary neurapophyses) have fused, and form
the coccyx.

1 See Albrecht, La queue chez Vhomme (Bull. soc. Anthvop.
Brux., vol. iii., 1884-1885,

144 UNIVERSALITY OF DEGENERATIVE EVOLUTION

2. The Frog. — The vertebral column in the
frog consists of nine vertebrae and the coccyx
(urostyle).

The first vertebra (the proatlas), which is fully-
functional, retains a centrum and two well-developed
neurapophyses, but the transverse processes, the
intercentrum, the haemapophyses and the ribs have
all disappeared.

The eight following vertebrae each have a
centrum, neurapophyses and transverse processes
which at least partially represent the ribs.

The coccyx, whether it be formed by the
lengthening out of the last caudal vertebra or by
the fusion of several, is undoubtedly part of the
vertebral column which has been transformed.
The coccyx of the frog is equal in length to the
whole of the remaining part of the vertebral
column, and is fully functional ; it serves as a
support to the pelvic region and fulfils the part of
a sacrum from the physiological point of view.
The coccyx, excepting at its commencement, con-
sists of only one centrum, or of several fused
centra, all other elements of the vertebrae have
disappeared. Here then is an animal in which the
modifications of the vertebral column have been
attended by the following retrogressive phenomena :
in the upper part of the vertebral column from
three to five parts of the vertebrae have disappeared ;
in the lower half, all the parts, excepting one, are
gone.

RUDIMENTARY ORGANS IN PLANTS 145

Section II.
Rudimentary Organs in Plants.

We have just glanced through a series of
rudimentary organs in animals, and many more
examples might easily have been furnished, but,
when dealing with rudimentary organs in the
vegetable world, much greater difficulty is met
with. In plants, the elimination of non-functional
organs is usually complete, and the vestiges left are
insignificant and hard to recognize. We can find,
however, amongst the various groups of the vege-
table world, and especially among the Phanerogams,
some instances of reduced organs.

§ 3. Rudimentary organs in various groups of
plants.

1. Algae. — On the surface of sea-wrack (Fucus)
may be found, distributed in large numbers, little
crypts (conceptacles) with hairs growing out of
them. On certain parts of the plant, these crypts
represent the organs of reproduction, producing
eggs and spermatozoa ; in other parts they fulfil
no known function and may be regarded as con-
ceptacles arrested in the course of development.
The fact that in other Fucacise (Splachnidium),
fertile conceptacles are distributed over the entire
surface of the plant adds support to this theory.

K

146 UNIVERSALITY OF DEGENERATIVE EVOLUTION

2. Mushrooms. — Among the Peronospora and
the Saprolegnese there originally existed, besides
the various sexual means of propagation, a typical
reproductive process, including eggs, and antheridia,
consisting of male branches separated by a cell-
wall from the rest of the organism.

In Pythium, for instance, in which this primitive
stage may be 'observed, an actual fecundation takes
place, the protoplasmic contents of the antheridia
being injected into the ova.

In other species, the organs of reproduction have
undergone a more or less complete degeneration.
In Phytoplitliora, a small part only of the male
protoplasm passes into the ova. In some species
of Saprolcgnia and Achlya, the male branch con-
tinues to attach itself to the ova, but the membrane
between them remains intact, and consequently
protoplasmic communication is not established.

In other species, the antheridia are very short,
and do not even touch the female cells.

In Leptomitus, which exhibits an advanced stage
of atrophy, the female organs are not discernible,
and reproduction is carried on completely asexu-

ally-1

3. Bryophyta. — In the germination of a certain
number of Hepatica?, belonging to such widely

1 For further details see A. de Bary, Vcrglciclicmlc Morphologic
unci Biologic dcr Pike, Leipzig, Engelman, 1884. W. Zopf, Die
Pilze. In Schenk's Handbuch dcr Botanik, 4. Bd., Breslau,
Trewemlt, 1890.

RUDIMENTARY ORGANS IN PLANTS 147

separated genera as Blasia, Eadula, and Prcissia,
four cells of equal size are formed, arranged round a
centre. One only of these cells proceeds to develop
into a plant, and the others simply atrophy. In
all probability the Hepaticaa have sprung from
some ancestor, in which each spore gave rise to
four individuals.

4. Pteridophyta. — According to Farlow,1 in-
stances of apogamy — the loss of sex — such as have
just been described as existing among mushrooms,
are also exhibited in certain ferns. In some
species,2 the eggs are not fertilized, but the organs
of reproduction still persist in a reduced condition ;
in other species there are no spores, and the
prothalli spring directly from the leaves (apospory).

5. Phanerogams. — Some of the Phanerogams —
Silene (fig. 58), Mclandryum, (fig. 59), Aspara-
gus, etc. — exhibit unisexual flowers, but have
obviously sprung from species of which the
flowers were hermaphrodite.

In Silene maritima (fig. 58) there are herma-
phrodite flowers (fig. 58, b), and also unisexual
flowers. The female flowers (fig. 58, a) still possess
some tiny stamens, each of which is provided with
filaments and anthers in a state of degeneration.
The male flowers have non - functional pistils,

1 W. Farlow, Ueber unycschlcclitlichc Erzcuyuny von Kcimpfltinz-
chen an Farnprothallien. Bot. Zeit., 1874, p. 180.

- A. de Bary, Ueber apogamc Fame u.s.w. Bot. Zeit., 187S,
p. 449.

148 UNIVERSALITY OF DEGENERATIVE EVOLUTION

Fig. 58. — Flowers of Silene maritima.

A, female flower with rudimentary stamens;

B, hermaphrodite flower.

consisting of an ovary stylus and stigma in reduced
conditions.

In Melandryum (fig. 59, a) the female flowers

retain only
vestiges of
stamens, and
the pistils of
the male flowers
are reduced to
mere filaments.
In Asparagus
officinalis all the
transition stages
between herma-
phrodite and unisexual flowers may be observed :
in the unisexual ,,

flowers, the organs r-^
of the opposite sex
still exist, though
in various stages of
degeneration. In
many unisexual
flowers which have
sprung from herma-
phrodite flowers —
Valeriana dioica,
for instance — no
traces of the non-
functional organs remain.

Among the Phanerogams, rudimentary organs

Fig. .19. — Flowers of Melandryum album.
A (to the left), a female flower; to the right, a
male flower ; e, rudimentary stamens form-
ing a circle at the base of the ovary.

RUDIMENTARY ORGANS IN PLANTS 149

appear not only in the reproductive organs, but in the
accessory organs of the flower — the calix and corolla.
Many of the Umbelliferae exhibit a reduced calyx.
The corolla persists, though in a very reduced
state in cleistogamous flowers (i.e. flowers which
never open, and which are self-fertilizing) — such as
the Oxalis, Impatiens, Violet, etc. The corollas of
the winter-opening flowers of Stellaria media are
much reduced, and for a very obvious reason —
the corolla exists only for the attraction of insects,
and there are no insects at that time of the year.

§ 4. Reduced organs in the vegetative apparatus
of the Phanerogams.

We have seen that reduced sexual organs are
exhibited among the various groups of plants, and
we will now mention a few instances of reduced
organs in the vegetative apparatus of the Phan-
erogams.

1. The embryo within the ripe seed of Phanerogams
contains a rudimentary root which develops during
germination. In certain Nympheaceaj — Nelumbium
Euryale and Victoria — this root never properly
develops.

In other aquatic plants degeneration has gone
further ; in the embryo of Utricularia, for instance,
the root has entirely disappeared.1

1 For further details relating to the roots of the Nympheaceaj and
the Utricularia see Goehel in Pflanzcnbiologische Sohilderungen,
vol. ii., Marburg, Elwert, 1891-1893.

150 UNIVERSALITY OF DEGENERATIVE EVOLUTION

2. As a general rule, the cotyledons, which are
the two first leaves to appear after germination, are
formed within the embryo. The ripe seeds of some
Anemones, however, contain no traces of cotyledons.1
These are formed, nevertheless,
after germination, sometimes sprout-
ing up out of the ground and
becoming functional, and occasion-
ally remaining underground, in
which case they are quite small,
without chlorophyll and non-
functional {Anemone nemorosa) ;
these underground leaves may
fairly be regarded as rudimentary
organs.

3. In Lathy rus Nissolia (fig. 60)
there are some very small stipules
of unimportant function, at the
base of the simplified leaves ; these
reduced stipules are occasionally
absent altogether.

4. The foliage organs in the
adult Oxalis oulpeurifolia are
merely represented by enlarged

leaflets. These phyllodes bear reduced leaflets
which rapidly disappear. In an adult specimen of
the Acacia which has phyllodes, these reduced
leaves are absent.

1 E. de Janczewsky, ittudes morphologiques sur Ic genre Anemone.
(Revue g&airale de botanique, t. iv., p. 241.)

Fig. 60.— Seedling of
Lathyrus Nissolia.

SURVIVALS EXIST IN ALL KINDS OF SOCIETIES 151

5. In several plants the assimilative function
of the leaves is lost, either because the plant is
parasitic or saprophytic, such as Corallorhiza,
Rafflesia, Cuscuta and Orobanche, or because the
assimilative function is relegated to the stem alone
as in the Euphorbia, of the desert, Ruscus, Mamil-
laria} Phyllocactus, Phyllantlius, and Miihlenbeckia,
or to the roots as in Tctmiophyllum?

In each of these cases the leaves are greatly
reduced, and only serve as a means of protection to
the functional organs, principally to the flowers
and buds, but though very minute they may often
be discerned quite easily on the young shoots.3

CHAPTER II

SURVIVALS EXIST IN ALL KINDS OF SOCIETIES

It may be said as certainly of societies as of other
organisms that certain modifications have taken
place, and that no society actually represents a
primitive social organization. All have been
submitted to more or less important modifications
and have lost some of their early institutions in

1 See fig. 51.

2 See further on the figs, of Phyllocactus (fig. 78), Phyllanfhus
(fig. 84), M'dhlenbeckia (fig. 80), and Tccnioph ylhnn (fig. 81).

3 Goebel, Pflanznibioloyischc Schilderv ngi m , Bd. i.

152 UNIVERSALITY OF DEGENERATIVE EVOLUTION

process of their development. In many cases this
can be historically demonstrated.

It may fairly be asserted that in all societies
there are instances of survival, i.e. survival of
customs, beliefs • and institutions, the original
character of which has so completely disappeared
that they might well be dispensed with altogether.

We shall deal only with such survivals as
correspond — mutatis mutandis — to the rudimentary
organs of animals and plants.

These survivals are of two kinds, the institu-
tion itself, such as the various corporations of the
city of London, which may still persist, though in
a modified condition, or there may remain only
traces of the institution in forms, ceremonies,
symbols, public games and fetes, customs and
legislative formula.

In order to demonstrate this point, it will not
be necessary to make a complete enumeration,
furnishing examples from all countries of the
world, or to draw up a complete list of survivals in
any given country. It will be enough to establish
two points :

1. That instances of survival shall be shown to

exist in all societies, even where they are
least likely to be found.

2. That, in any institution — that of the family,

for instance— survivals may be found of
all the former stages through which it
passed into its present condition.

SURVIVALS EXIST IN ALL KINDS .OF SOCIETIES 153

These two points being established our conclu-
sions drawn from them may be given a very wide
application.

§ I. Instances of survived in various groups.

"It is a well-known fact," says Kowalevsky,
" that as the past gives place to the present it
leaves traces which vary in number and impor-
tance." x

This is obviously the case with regard to most-
customs, but it is unnecessary to point out all the
instances of survival which abound among the
peoples of the countries round about us. They are
naturally most common in barbarous societies
where the servile imitation of the ancestor plays a
much greater part than with us. This is strongly
urged by Bagehot in the following passage : —

" Man," he says, " may be defined as a creature
of habit. As he has done a thing once, so he will
probably do it again, and the oftener he has done
a thing the more likely he will be to repeat it in
the same way, and, what is more, to insist upon
others doing the same.

" By means of counsel and example he transmits
to his offspring the customs he himself originated.
This is true of the human beings to-day and will
doubtless hold good for all time. It is character-

1 Kowalevsky, Tableau des origines de la propriiti et de la
fa at ill e, p. 7.

154 UNIVERSALITY OF DEGENERATIVE EVOLUTION

istic of primitive societies that sooner or later most
of these customs come to be regarded as having a
supernatural sanction, and the whole community is
impressed with the belief that if the old tribal
customs are violated, incalculable misfortune will
follow." l

Social modifications are therefore effected very
slowly and with great difficulty — stagnation is the
rule, and progress but a rare exception, innovators
being forced to retain the greater part of the old
institutions, introducing only an inevitable minimum
of change. A course of history, or a careful study
of the conditions of social institutions at an earlier
stage of development than our own, will furnish
numerous instances of survivals. It now remains
to be seen if there are no rudimentary social groups
wherein all the primitive institutions have been
retained, and which, having undergone no modifica-
tions, exhibit no traces of degeneration. We shall
further see if no form of society exists uninfluenced
by the spirit of tradition, and where institutions
which have come to be regarded as no longer useful,
disappear suddenly and entirely either by voluntary
dissolution or by legislative measures. Only in these
two extreme cases can the existence of survivals
be questioned. We have then to establish two
points:

(a) That all societies, even those to be regarded as

1 Bagehot, Lois scicntifiques du dtrcloppcment dcs nations, p. 154,
Bibl. scient. intern., Paris, F. Alcan, 1885.

SURVIVALS EXIST IN ALL KINDS OF SOCIETIES 155

primitive, have undergone certain modifica-
tions.
(b) That all societies, even the least conservative,
exhibit instances of reduced institutions,
and of vestiges of institutions which have
disappeared.
We will take the second point first, as it can be
more briefly dealt with.

1. We know that imitation of the past and
respect for tradition and custom are reduced to a
minimum in modern societies, especially in the
countries of the New World. Even in these
recently formed States, however, instance of sur-
vival may be found.

In the first place, there are legal and religious
survivals of European origin. Jews settling in the
United States kept up the practice of circumcision,
while Christians introduced the Eucharist. Spencer *
has shown that forms of greeting are vestiges of a
primitive ceremonial demonstrating submission to
the omnipotence of others. Then take the Calendar
system which is universal; we know that the names
of months and clays of the weeks are survivals from
Polytheistic times, and it seems certain that both
circumcision and the celebration of the Mass are
true survivals which originated in religious sacrifices.
Independently of these imitative survivals vestiges
remain in the United States of reduced institutions
which were fully functional in the last century.
1 Essays on Progress.

156 UNIVERSALITY OF DEGENERATIVE EVOLUTION

Take for instance the town meetings of Boston and
Newhaven.1

When the inhabitants of a town attained to a
certain number, the town became a city, and the
Assembly of inhabitants was transformed into a
Common Council. In some instances, this old
system persists though in a reduced condition.

In Boston, which continued to be a town,
governed by an Assembly of all the inhabitants up
to 1821, the present Charter of the city authorizes
the convocation of a town meeting wherever the
Mayor and Aldermen consider it advisable ; the
latter, however, never make use of this privilege.

In Newhaven (Connecticut), the old town meeting
continues to exist side by side with the Common
Council which was established in 1784, but,
Levermore says, " This ancient institution nowadays
is a meeting together of a small number of citizens
to conduct the business of several thousands. The
few people connected with the affairs of the town
(which is very poor), meet together to discuss
matters in a friendly way, decide what money is
required for current expenses, and then adjourn.
Not one in seventy of the inhabitants attends these
meetings. Few know when they take place, and
the papers make brief, if any, mention of them."

2. We have now to show that the simplest
societies have undergone modifications, and exhibit

1 Bryce, The American Commonwealth, i., pp. 598 and following.
London, Macmillan, 1893.

SURVIVALS EXIST IN ALL KINDS OF SOCIETIES 157

instances of survival. We will take as examples
those rudimentary types most nearly approaching to
the primitive type,1 i.e. the Veddahs of Ceylon, the
Fuegoes of Cape Horn, and the Australian tribes.

(a) The Veddahs, who have lived in the jungles
of Ceylon for centuries, either as separate families,
or in groups of two or three families, appear to
have formerly possessed a much more complicated
social organization. According to Max Mliller,
they were not formerly so low in the scale of
humanity ; he says that their language, if not
their blood, betrays their " distant connection with
Plato, Newton, and Goethe."

In their language, folk-lore, and clothing, these
retain characteristic vestiges of a former condition.
Take for instance the carefully observed practice of
piercing the ears of children at the age of three or
four years, although eventually only a small number
of them could wear ornaments in them, others
having to be content with small pieces of twig,
coiled leaves, or bits of straw.

" This custom," says Deschamps, " is extremely

old, and we may suppose — as there is no other

signification in it than the prospect of ultimately

wearing jewels — that it dates back from a time

when the people were not in so low and destitute

1 " Aggregates formed by a simple repetition of hordes or clans
without any such interrelations between them as to form inter-
mediate groups between the whole collection and the individual
clans." Durckheim, les Regies de la mitlwde sociologique, Paris,
F. Alcan, 1895.

158 UNIVERSALITY OF DEGENERATIVE EVOLUTION

a condition as they are now. Having in more
civilized times worn jewels in their ears, the
custom of piercing the ears in youth persists,
though the jewels may be lacking."1

(6) Bridges says that according to a tradition
which is probably true, the Fuegoes, until quite
recently, submitted their young men to a sort of
initiatory trial when they attained to adolescence.
They were taken into a hut (the kind) set apart
for the purpose, and there underwent certain tests,
including a rigorous fast. Bridges adds that the
kina was also the theatre of mysterious and bizarre
scenes of very ancient origin, the roles of which,
now relegated to men only, were entirely performed
by women. Contrary to Giraud-Teulon who cites
these facts as evidence of the former existence of
a matriarchy, the fetes of the kina seem to have quite
disappeared from among the natives of Orange Bay.

Dr Hyades, however, mentions a survival of the
old custom. " The custom is still observed of
submitting young girls to a fast at the time of
puberty, but this fast is less severe than that
already mentioned as undergone by the boys ; the
same good advice is then given them by their
parents, as was formerly bestowed upon the boys
in the Kina." 2

1 Emile Deschamps, V Anthropologic, 1891, t. ii., pp. 297 and
following.

2 Mission scientifique du cap Horn, 1882-1883. t. vii. Anthro-
pologic, Ethnographic, by P. Hyades and J. Deniker ; Paris,
Gauthier-Villars, 1891, p. 377.

SURVIVALS EXIST IN ALL KINDS OF SOCIETIES 159

(c) Eecent researches into the family system
among the Australian tribes has brought a number
of survivals to light. This is especially the case
with regard to the careful researches of Fison and
Howitt1 who have shown that, independently of
their tribal divisions — which are really territorial
groups — the Australians are divided up into clans
or sexual groups comprising all the individuals with
the same Kobong.2

The members of these groups are regarded as
members of the same family, and may never,
under any circumstance, intermarry, under pain
of being driven out of the clan and hunted like
wild beasts. Sometimes individuals of antagonistic
tribes living at several hundred miles' distance from
one another and speaking different languages have
the same Kobowj. The law of classes remains
active ; a captor may not violate a prisoner belong-
ing to his group, but a stranger may enter into
relations with the women of another tribe, so long
as the tribe belongs to a class related to his own.
This system of relationship can only be explained
as being a survival from a former period in which
all persons with the same Kobowj belonged to the
same group. This is a disputed point,3 however,

1 Fison aud Howitt, Kurnal and Kamilaroi {Journal of the
Anthropological Institute, 1884).

2 "The Kobong of a man is the animal or plant, the name of
which he bears and reveres as a protecting spirit" (Starcke).

3 Starcke, la Famillc primitive (Bibl. sciens. intern., Paris,
F. Alcan, 1891, p. 22).

160 UNIVERSALITY OF DEGENERATIVE EVOLUTION

for besides this very likely hypothesis, undoubted
survivals remain of intermarriage by groups or
sexual groups. In the writings of Fison and Howitt,
we find the two following instances of this in two
tribes which, according to them, severally represent
the highest and lowest in the scale of civilization,
among those with which they came in contact.1

(r/) The tribe called Kunandaburi was divided
into two exogamous classes : Mattara and Yungo.
Theoretically all the Yungos whether male or
female were regarded as the males of the Mattaras,
and vice versa. In point of fact, however, only
one vestige of the primitive communal marriage
remained — the jus primce noctis which was the
prerogative of all the contemporaries of the
husband belonging to the same group.

(e) The tribe called Narrinycri which repre-
sented a more advanced stage of civilization, was
equally divided into two sexual groups, but in
reality, marriage was strictly individual. One
survival remained, however, of the former system.
When a man captured an alien bride, all the men
of his own generation and belonging to the same
group possessed the right of jus primce noctis.

3. We have seen that instances of survival are
rare in some countries because modifications are
only effected slowly, and in others because changes
are effected very quickly and useless institutions

1 Fison and Howitt, Journal of the Anthropological Institute,
1882, p. 35.

SURVIVALS EXIST IN ALL KINDS OF SOCIETIES 1 6 1

are eliminated root and branch. It is in countries
like England, where modifications are brought about
with a due respect for old customs and traditions
that ceremonies, institutions and customs exhibit
the greatest number of survivals.

§ 2. Survivals of ancient forms of marriage and of
the family in Modern Europe.

We think we may regard it as proved that
all societies exhibit instances of survival, but in
order to further demonstrate the universal character
of retrogressive evolution, we shall show, by means
of a careful study of one particular institution, that
vestiges of former institutions are neither rare nor
exceptional, taking as examples the various forms
of marriage and of the family throughout Modern
Europe.

1. Forms of Marriage. — From archaic times* up
to our own, we find that among modern nations,
marriage by capture, marriage by purchase, and
marriage with the consent of the woman have been
successively followed by marriage by simple consent,
religious marriage {in facie Ecclesioz) and civil
marriage,1 and that survivals remain of all the
forms of marriage anterior to civil marriage.2

1 Paul Viollet, Histoirc du droit civil Francais, p. 424 and
following. Paris, Larose & Forcel, 1893.

2 Westermarck, The History of Human Marriage, 1892, p. 418
(when the mode of contracting a marriage altered, the earlier mode,
from having been a reality, survived as " ceremony ").

L

162 UNIVERSALITY OF DEGENERATIVE EVOLUTION

Marriage by Capture. — Traces of this early form
of marriage, which is still to be met with in certain
parts of Bulgaria,1 are exhibited in the nuptial rites
and customs of Ukraine, where the ceremonial is
quite a museum of reduced institutions.

Vestiges of what originally was marriage by
capture occur too in France and Belgium, and other
countries. The nuptial games of Lower Brittany
and Chimay have obviously been derived from it.2

Marriage by purchase. — The retrogressive evolu-
tion of this form of marriage exhibits the following
stages :

1 Th. Volvok, Bites et usuges nuptiaux de V Ukraine (V Anthro-
pologic, 1891, p. 169). " In certain parts of Bulgaria (Kustcndil),
the capture of young girls takes place even in the present day and
constitutes a form of marriage (Vlatcheny monny)."

2 Monseur, Bulletin du Folklore, January and March 1895,
Coutumes, p. 1 ; les notes, p. 18.

"In some villages of Chimay (Hainault), when a young man
chooses a bride from a neighbouring village, the young people
proceed in cavalcade to the home of the bride. The leader of
them presents her with a whip and a large cake crowned with
a bouquet. The bride then takes up her position on the door-
steps while the bridegroom and his friends pass and repass her
at full gallop seeking to dispossess her of the whip with which
she lashes at them. All this may be regarded as the last remain-
ing phase of marriage by capture, the bridegroom arriving at the
dwelling of the bride with a cortege, being more or less figurative
of a marauding expedition."

Baudrillard (in Stances et travaux de V Academic des sciences
morales, Jan. 1884, p. 36) connects the customary nuptial games
of Lower Brittany with the primitive marriage by capture. In
these games the bride hides before going to church, and the
bridegroom has to search till he finds her.

SURVIVALS EXIST IN ALL KINDS OF SOCIETIES 163

1. The payment made was originally a pecu-

niary compensation to the family of
the captured bride. By degrees, actual
capture gave place to mere symbolical
capture and then the system of compensa-
tion became transformed into the purchase
system.

2. The price paid for the bride which was origin-

ally the property of the whole clan under
the name of wergelt, became the perquisite
of the bride's father.

3. It next became modified into a marriage

dowry given to the wife by her husband
and the ceremony of purchase became
purely symbolical. In the Merovingian
period, for instance, the future husband
presented the father of the bride with a
sou and a denier (marriage per solidum et
denarium)}

Other survivals still exist of these three stages
in the evolution of marriage.

(a) In Ukraine, where the signification of the
purchase system was purely one of the vira
or compensation (wergelt), the bridegroom
has to give a. present to each member of
the bride's family ; this is the custom too
in the valleys of the Caucasus where the

1 Vanderkindere, Condition de lafemmeA Vipoque mirovingietwie,
\>. 12. A. Heussler, Institution' n des dcutschen Pricalrcchts,\i., p.
280.

164 UNIVERSALITY OF DEGENERATIVE EVOLUTION

relations make a further demand of pay-
ment in money.1

(b) The custom of purchasing the bride from the

father still exists — although with no legal
recognition among the Ossetes and in certain
Russian villages.

(c) Lastly, the marriage per solidum ct denarium.

Traces of this are left even in modern
France. This particular purchase signified
the payment of thirteen silver deniers
according to a certain value of the sou.
The introduction of these thirteen deniers into
the ceremony of marriage can be traced through
the middle ages up to the marriage of Louis XVI.
when they still appeared. In some parts of France
such as Dijon, Bordeaux and Barrois, they may
still be met with even nowadays. With the excep-
tion of this figurative number of thirteen, character-
istic of the primitive origin of the ceremony, this
form of marriage has undergone such changes as
almost to entirely obliterate its primitive character.
This is so, too, with other survivals of the system
of marriage by purchase — they have come to be
only intelligible by means of the comparative
method, or by a knowledge of their historical
antecedents.2

1 Volvok, Journal V Anthropologic, 1892, p. 579. Kowalevsky,
Droit coutumier ossiticn, p. 176.

2 See Paul Viollet, Histoire du droit cicil francais ; Paris, 1893,
p. 403.

SURVIVALS EXIST IN ALL KINDS OF SOCIETIES 165

Marriage fairs — now transformed into kermesses,
and still held at Lierre in Belgium and other
places — appear to have been originally regular
markets for the purchase of young women.1

The custom of offering the wrong young woman
to the bridegroom as his future wife — a custom
still in vogue in the department of Landes in
France — formed part of the nuptial ceremonies
among the ancient Hindoos, and is probably a
vestige of the tricks which were played upon the
bridegroom after the purchase of the bride. Several
vestiges of this kind may be noticed among the
customs of modern peasants. It is by no means
infrequent for a wife to be sold by her husband,
on the principle that what has been bought may
fairly be sold, and the transaction rendered legally
binding by being drawn up on stamped paper.
This is merely a survival of the old system of
marriage by purchase.

1 "The second Sunday of this fair, which commences on the
Sunday after All Saint's Day, is called the veersensmarkt (Heifer-
fair or market), and the third Sunday is called the brullenmcrll ,
a name derived from the word brid, and signifying a heifer lowing
noisily. The reerscnsmerkt is the day especially set apart for the
young girls who attend the fair to find husbands. The brullen-
merkt is the day dedicated to older women as a sort of forlorn hope
for those who have hitherto failed to get married. No one seems
to know when these " markets " first came to be held. I am more
than sixty years old, and they were old when I was young, with
this difference only, that in my youth, a young girl who respected
herself would not have been seen at the brullrnmerkt, which is not
the case nowadays." (Taken from a letter from the Secretary of
the Commune at Lierre.)

16(5 UNIVERSALITY OF DEGENERATIVE EVOLUTION

Marriage by consent of both parties. — In French
civil marriages of the present day, traces remain of
the two other forms of marriage by common con-
sent which preceded it.

(a) The marriage by simple agreement, which held
good throughout Christian Europe until the Council
of Trent, is still valid in Scotland. Gretna Green
marriages (Gretna Green being a village situated on
the Border, near Carlisle) were notorious. In ac-
cordance with an old custom, the blacksmith of
Gretna Green kept the register of these marriages,
and the union was contracted in his presence.1

After the Council of Trent, marriages in facie
Ecclesice came to be alone recognized by the
Church : all marriage to be valid must be sanc-
tioned (if not actually celebrated) by the parish
priest of one contracting party, in the presence of
one or more witnesses. The legislation of the
Council of Trent became French law after the
Ordinance of Blois in 1574, until the law was
revised in September 1792, when civil marriage
was definitely established in France.

1 Viollct, Histoirc du droit civil fin meals, p. 428.

''These marriages, about which everyone has heard, were not
an invention of Scotch law. Like most legal curiosities, they
find an explanation in survival from a former condition of
tilings, persisting in a singularly original and peculiar form.

"The blacksmith and his register were not necessary in them-
selves to contract a marriage ; they merely constituted evidence
that it had taken place. In 1804 a Scotch man and woman
merely declared themselves man and wife in writing, and in
1811 they were recognized as legally married by Scotch law."

SURVIVALS EXIST IN ALL KINDS OF SOCIETIES 167

From that time forth, religious marriage may be
regarded in the light of a survival, having lost all
legal importance, while civil marriages are greatly
on the increase. Priests, then, the " groomsmen "
of Lower Brittany, and the blacksmith of Gretna
Green may alike be regarded as plain evidences of
an institution in process of decline.

II. The family system. — Archaic forms of the
family still exist, sometimes as mere vestiges,
sometimes as exceptional cases, in countries where
only separate families are legally recognized.

1. The Matriarchy. — Traces of the matriarchy,
i.e. of the exogamous family of blood relations
through the mother, abound among the customs
of the inhabitants of the valleys of the Caucasus.
Kowelevsky, in his book on the customs of the
Ossetes, has dealt with this subject. The vestiges
remaining even in France of marriage by capture
and the prohibition of certain marriages in Monte-
negro, derived doubtless from an exogamous period,
are alike survivals of this primitive family system.
Further, although both facts and their interpreta-
tion are rather doubtful, some authorities regard
the couvade, a custom which is still in practice
among the Basques and also in the Isle of Mark
(in Holland), as a vestige of the transition period
between the matriarchy and paternal affiliation.1

1 See Viollet, Precis de Vhistoire du droit francais, ii. 326, and
Giraud-Teulon in Origines du marriage et de la famille primitive,
Paris, 1881, p.. 138 ; and Starckc, Famille primitive, p. 49.

168 UNIVERSALITY OF DEGENERATIVE EVOLUTION

2. The Patriarchy. — The Patriarchy, which was
a system of family community, is still exhibited in
the participanze of Italy, the companias de Galicia
of Spain, the parsonneries of France, the Haus-
genossenschaften of Germany, and the zadrugas of
the Balkan peninsular. Besides these, the family
system of to-day, according to Sumner-Maine,
affords constant evidences of a patria protestas
in process of decay and of declining male property
rights.

Survivals from the old patriarchal system abound
in modern legislation. Take the following examples :

(a) The limitations imposed on a testator with

regard to leaving his property away from
his family (Civ. Code, 213 and following).

(b) The legal opposition to a woman's equality

in succession.
The more barbarous laws, if not actually
excluding women from succession, at least
excluded their succession to landed estate,
in order to keep the family property intact.
In France the privilege of sex was main-
tained in some respects, even among the
peasants, up to the close of the old
regime, and it still exists in the present
day in different degrees in Scandinavia,
Russia, Servia, and some of the Swiss
Cantons,
c) The inequality of the sexes as regards con-
jugal fidelity.

SURVIVALS EXIST IN ALL KINDS OF SOCIETIES 169

The privileges of the husband over the wife
in this matter are survivals from the time
when these obligations were wholly on one
side.
(d) Affiliation following on a double marriage. —
In some countries, under the old system, a
child could only succeed to the property of
his parents if he resided with them (ex-
cepting in quite exceptional cases). Hence
there arose marriages by exchange. In
order to compensate the children, the two
families, if each had a son and daughter,
exchanged them, and bestowed the rights
of one upon the other. Under the new law
these rights could not be legally claimed,
and yet even in the present century they
are not wholly unknown. The last mar-
riage by exchange was probably that
mentioned by Dupin1 which took place at
Gacogne (Nievre) in 1839.
In conclusion then, we have found that while
the legislative systems of modern races tend to
become more and more alike in main principle, we
can yet find vestiges, more or less faint and dis-
torted, but quite recognizable, of the different
institutions which dominated the earlier conditions
of the different races.

1 Viollet, ffistoire du droit civil frangais, p. 491.

PART III

SUMMARY AND CONCLUSIONS

The examples we have been able to give in the
first part of this volume make it plain that de-
generative evolution exists everywhere. It must be
noticed, however, that biological investigation shows
that in the evolution of organs certain parts may
disappear completely, but also that in the evolution
of organisms certain organs may also disappear.
This last phenomenon is most common in embryo-
logical development, when it is known as ontological
abbreviation.1 Sometimes it is the adult stage that
is suppressed, this being possible by what is called
pedogenesis a precocious appearance and ripening
of the sexual organs.2

1 In Scalpellum Stroini, a deep sea Cirripede, the nauplius stage
of the larval life is suppressed at least so far as that is a free
swimming larva. Here is at least a physiologically complete sup-
pression of a whole larval stage.

2 Axolotl. Most salamanders pass through a larval stage in
which they are aquatic and perform their respiration by means of
external gills. In this condition they are incapable of reproduc-
tion, and must undergo metamorphosis to secure propagation of
the species. In the case of Amblystoma, however, a Mexican
salamander, the larval form of which is called the Axolotl, repro-
duction is possible in the larval stage. Thus most individuals of

SUMMARY AND CONCLUSIONS 171

Sometimes a degenerative transformation becomes
still more complete and wonderful ; not only may
a larval stage or an adult stage be completely
suppressed, but a multicellular organism may even
lose its power of dying. It is known that the
simplest forms of life are practically immortal :
when a microbe like micrococcus divides nothing
dies, and throughout the whole series of successive
divisions the primitive life is preserved. On the
other hand, in the case of higher animals such as
man there are both mortal somatic cells and
reproductive cells which by means of conjugation
become practically immortal. The mortality of the
somatic cells is evidently an acquisition, an advant-
age fixed by natural selection ; but there exist
multicellular organisms evidently derived from
creatures which had acquired the division into
mortal somatic and immortal reproductive cells and
which have lost it since. All the cells of their
body are able to avoid death by conjugation.
This occurs in many conjugate algae like spirogyra

this species do not actually reach the adult stage. According to
Boas writing on Neotenie in Gegenbaur's Festschrift, 1896, this
probably happens in the case of all the perennibranchiate urodeles.
Eanunculaeece. On page 85 we showed that in Ranunculus
aquatilis there are produced first submerged leaves, and afterwards
floating lobed leaves, and that the flowers are produced in the
axils of the floating leaves. Some forms of the plant living in
deep water produce only lacinated leaves, in the axils of which by
a kind of pedogenesis the flowers are produced. In other species
[Ranunculus fluitans and B. ttivaricatus) the predogenesis has
become definitely established and no floating loaves are formed.

172 UNIVERSALITY OF DEGENERATIVE EVOLUTION

and in some of the Volvocinere (Stephanosphsera,
Eudorina).

Plainly then, the further one examines the facts,
the more enlarged becomes the conception of de-
generative evolution. It is not confined to unusual,
abnormal or pathological cases. Degeneration is
not an accident in evolution : it is the obverse of
progressive evolution and the necessary complement
of every transformation whether anatomical or
social.

Whatever transformation may be studied, it will
be found that change is always accompanied by an
elimination of some parts and that in the interests
of the organism as a whole these useless parts
gradually degenerate. When a whole organization
begins to undergo retrogressive evolution and to
decay, it is frequently in the interests of some still
larger organization. Individuals or species out of
harmony with their surroundings disappear to make
room for others. August Comte has shown how
death is a progressive agency in the social organi-
zation removing the worn-out tissues and leaving
room for new and more plastic intelligences. All
progress implies necessary eliminations. In the
domain of society, those who are victims and who
from birth, education, or interests, attach themselves
to the decaying institutions naturally see only the
degenerative side of the change ; but those who
regard the process as a whole and do not concentrate
their attention upon the injured interests and

SUMMABY AND CONCLUSIONS 173

individual sufferings will see the other side of the
movement.

When a social organism is degenerating there is
considerable opposition to its complete disappearance,
and so as Houzeau has said (see the summary of
Book III.) it is to be expected that living and
superior civilizations drag behind them a trail of
debris from dead civilizations.

BOOK II

THE PATH OF DEGENERATIVE
EVOLUTION

PART I

THE SUPPOSED LAW THAT DEGENERATION RETRACES
THE STEPS OF PROGRESS

It is a common opinion, supported partly by the
etymology of the word, that retrogression is a tracing
backwards of progression.

" In the degeneration of organizations fallen out
of use,': M. A. Lameere has said, " it is to be ob-
served that the structures formed most recently
and most specialized are the first to disappear, and
that the most fundamental characters are those
which persist longest : that in fact degenerative
evolution retraces the steps made by progressive
evolution. Peculiarities recently acquired, if they
become disused, rapidly disappear, while dispositions
of a more ancient kind have a persistence almost
exactly proportioned to their age.1

This supposed biological law of retracement has

1 A. Lameere, Esquisse clc la Zoologie, Bruxelles, Rosez, p. 184.

176 THE PATH OF DEGENERATIVE EVOLUTION

penetrated to psychology and sociology. In 1868
Hughlings Jackson, in the study of certain maladies
of the nervous system, had arrived at the conclusion
that, "In the degeneration of this system the higher
functions, those more complex, specialized and volun-
tary, disappear more quickly than the lower, simpler,
less specialized and more automatic functions." x

Starting from this point, and expressing it in
terms of physiology, Eibot formulates as follows
the law of degeneration of will and memory :
" The dissolution of the will occurs in a retro-
grade fashion, from the more voluntary and com-
plex to the less voluntary and simpler — that is to
say, towards automatism." 2

So also in progressive loss of memory, the
degeneration proceeds from the less stable to the
more stable. " It begins with recent acquisitions
not firmly rooted in the brain, rarely repeated, and
so not firmly associated with others, in fact with
the least organized parts of memory. It ends
with sensory memory which is instinctive, and is
deeply rooted in the organism, or is indeed a
part of the organism itself." 3

These retrograde transformations of the nervous
centres have their echoes in the modes in which
ideas and feelings are expressed. Paul Heger, in

1 Ribot, Maladies de la Mimoire, p. 29. Dallemagiie (Degen&ts
et Ddse'quilibre's), p. 430.

2 Maladies de la Volonte, p. J.50. Paris, F. Alcan.

3 Maladies de la Memoire, p. 94.

DEGENERATION RETRACES STEPS OF PROGRESS 177

particular, has shown this in his investigations into
the degeneration of writing and speech.1

In a lecture on the evolution of language, de-
livered a't the University of Brussels, he said as
follows : " For several years I have studied the
degenerative evolution of writing, and I have
shown how the writing of the insane resembles
that of children. All that I have said with
regard to writing applies to speech, and just as
drawing lasts longer than writing, so rhythm and
music survive after the power of expressing ideas
by words has been lost." " The gradual degenera-
tion of speech may be traced in the case of old
men who gradually pass down the incline into
senility. Study of the speech of such persons
shows that the degeneration of their faculty re-
traces the steps by which it had been progressively
acquired."

The labours of Heger were in a field where the
social element was important. It is a small step
from them to social affairs themselves. A number
of authors, including Ferrero, Colajanni and Degreef,
base their ideas upon this law of retrogression, which
they regard as established and applying to sociologi-
cal phenomena.2

1 Sur revolution Regressive de Vficriture chez certains alUnis
(Bull, de la SociiU d' Anthropologic de Bruxelles, v., 1885-1886).
Sur revolution du Langagc (Revue Universitaire, Bruxelles, 1892-
1893, p. 143).

2 Degreef, Le Transformisme Social. F. Alcan, Paris, 1895,
p. 365.

M

178 THE PATH OF DEGENERATIVE EVOLUTION

Before describing and discussing the special facts
to which they apply their theory, it is necessary to
examine its biological foundation.

CHAPTEE I

THE PATH OF DEGENEEATION IN BIOLOGY

We have now to consider if the degeneration of
organs retraces the steps taken in their progres-
sive evolution. According to Hughlings Jackson
and Bibot, in the cases mentioned in the preceding
chapter, degeneration proceeds by successive atro-
phies occurring in the order opposite to that of
ontological formation. Is the same order to be
found when we compare the degeneration of
organs or of individuals with their phylogenetic
development ? To answer this question, we must
employ both morphology and embryology. Using
the morphological method, we shall study the re-
duction of a homologous organ in several species
descended from the same type, and compare the
different stages of reduction with the different
stages of phylogenetic development of the organ.1

1 In this investigation it will be necessary to compare absolutely
identical organs — for instance, not to compare the pineal eye with
the paired eyes. It will be necessary also to choose animals of
common parentage — to avoid, for instance, comparing a Vertebrate
eye with a Crustacean eye.

THE PATH OF DEGENERATION IN ANIMALS 179

Using the method of embryology and the prin-
ciple, so far as it can be followed, that ontogeny
recapitulates phylogeny, we shall investigate the
mode of origin of some reduced structures. We
shall thus learn if the organs in degenerating
resume any of their ancestral stages.

Reduced organs may appear in two different
forms. They may be atrophied after having
reached a more or less complete development,
and in this case we shall have to compare the
course of the atrophy with the course of the
development. They may be rudimentary, that
is to say. their development may have been
arrested at a given point, the adult state never
being attained. In this case, so far as ontogeny
repeats phylogeny, the arrest of these organs at
different stages in different species should furnish
a series with greater or smaller lacunae, but a
series which will be the reversed series of the
original stages in phylogeny.

Section I.
The path of degeneration in animals.

1. Morphology and Embryology. The law of re-
capitulation. — It has been so often repeated that the
individual development of an organ is a resume more
or less compressed of its historical evolution, that
people are apt to attempt too exact an application

180 THE PATH OF DEGENERATIVE EVOLUTION

of this principle to every individual case. Such an
exact application is, however, impossible. Every
living organism is plastic, and in its development
presents individual variations which serve as
material for the operation of natural selection.
In consequence, the recapitulation cannot be more
than a repetition more or less vague of the essential
phases of phylogeny.1

Moreover, there is nothing inevitable in the law
of recapitulation, for most plants develop directly.

With these limitations, however, we may state
that among animals, the ontogeny usually repeats
in a modified fashion the main ancestral stages.
This is certainly the case when we compare the
development of the brain of man with the probable
ancestral stages as displayed in the series of verte-
brates.

1 Lang, Anatomic compared. As throughout the whole course of
time, adaptation, that is to say, the preservation of what is most
useful in the struggle for existence, is a force modifying heredity,
it is plain that a species instead of resting stable must change.
According to its circumstances, moreover, the successive stages in
the ontogeny of a creature are under the influence of conditions
different from those that affected the corresponding ancestral stages.
We shall call the process of embryology palingenetic so far as it is
based upon inherited legacies, and co?nogenetic so far as it is modi-
fied by adaptation.

Baldwin in his Treatise on Mental development in the child and
in the race (London, 1895), also shows that the development of the
individual is not an exact repetition of ancestral stages. The
development of the child exhibits "short cuts" and phases of
direct development due to adaptation and destroying the exactness
of the parallel with phylogeny.

THE PATH OF DEGENERATION IN ANIMALS 1 8 1

In Fish
and Batrachia.

The cerebral hemispheres
do not cover the region
of the third ventricle
from which the eyes arise
(thalamencephalon).

In Reptiles. The hemispheres cover
the thalamenaphalon but
leave unco vered the region
of the optic lobes (mesen-
cephalon).

In the human
embryo (fig.
61, A) of the
seventh week.

Same
aspect.

In the human
embryo (fig.
61, b) of the
middle of the
third month.

Same
aspect.

In mammals.

The hemispheres cover
the thalamencephalon,
the mesencephalon, some-
times the metencephalon
(cerebellum and medulla),
and the olfactory lobes.

In the human
embryo (fig.
61, c) of the
fifth month.

Same
aspect.

In some mam- The hemispheres are

mals even of smooth.

higher orders

{e. (j. some

Hapalidre).

In the human Same
embryo (fig. aspect.
61, D) of the
midddle of the
fifth month.

Within such limits, the law of recapitulation
may be applied, and the embryonic history of an
individual may be considered roundly as a repeti-
tion of the essential phases of its ancestral history.
We have now to consider how far a reduction by
atrophy or by arrest represents a retracing of steps
in evolution (fig. 61).

From this point of view, we may study the

182 THE PATH OF DEGENERATIVE EVOLUTION

degeneration of the pineal eye in the slow-worm,
and in a series of lizards.

The pineal or median eye in the slow worm and

Fig. 61. — A, Brain of a human embryo of seven weeks ; h, cerebral hemispheres;
ci, intermediate brain or thalarr.encphalon ; cm, midbrain ; cp, hind-brain.
B, Brain of a human embryo about the beginning of the third month ; h, cere-
bral hemispheres; Ig, region of the corpora quadrigemina; cm, mid-brain.
0, Brain of a human embryo at the middle of the third month; h, cerebral
hemispheres ; (q, corpora quadrigemina; ce, cerebellum. D, Human brain
of the fifth embryonic month ; A. cerebral hemispheres; olf, olfactory lobes ;
S, fissure of Sylvius; ce, cerebellum. (After Mihalkovics, Jb'niwkkelungs-
gcschichte des Gehirns. Leipzig, 1877.)

the lizard passes through the following stages in
its individual development.1

(1) Formation of a hollow outgrowth from the

roof of the third ventricle of the brain

(fig. 62, d).

1 P. Francotte, Rccherchcs sur le ddveloppement dc Vfyipliysc.
( These presentee d la Faculle de medecine de Bruxel/es. ) Arch, de
Biologie, 1888.

THE PATH OF DEGENERATION IN ANIMALS 183

(2) This little sac elongates, changes its direction,

B

^ ,r

c,r

E

,cr

jSL

/CP

Fig. 62.— Diagram indicating the progressive evolution and the degeneration of
the pineal eye.

A. Perfect pineal eye, as found in the slow-worm before birth, or in the adult
Sphenodon (Hatteria) ; c, lens; r, retina; n, optic nerve; d, divertialum of the
thalamencephalon. B. Pineal eye in first stage of degeneration as it exists in
Chamceleo and as it was in the slow-worm before stage A. The lens (c), and
the retina (/•), are not differentiated. C. Pineal eye in the degenerate form
found in Calotes and Leiodera; c, lens; ;•, retina; ti, optic nerve in fatty de-
generation. P. Very degenerate pineal eye as in Cyclodus and like the earliest
stage in the slow-worm ; there is no differentiation of the divertialum from the
thalamencephalon. E, F, G. Other modes of degeneration of the pineal eye.
The eye lies within the skull and there is no parietal foramen ; cr, cranial
membranes; E. Ceratophora. F. Birds; g, mammals. (After Baldwin
Spencer.)

and becomes divided into a proximal and
distal portion. The cells lining the distal

184 THE PATH OF DEGENERATIVE EVOLUTION

part, that farthest from the brain, become
differentiated into the cells which will
form the lens, and the cells which will
form the retina.

(3) The distal part becomes specialized, the lens,

the retina, and the stalk of the optic nerve
are mapped out.

(4) The lens, the retina, and the optic nerve

become fully formed (fig. 62, a).

At this stage the third eye has reached its limit
of development.

There is a well-formed retina connected with the
brain by a special optic nerve. The organ projects
strongly from the surface of the head, but from
this point, owing to the development of the
cerebral hemispheres, degeneration begins. The
nerve (fig. 52, c), becomes broken and fatty, and
pigmentary degeneration occurs in it. At the
same time, the pineal eye having become useless
or even harmful to the animal possessed of it,
before the power of receiving perceptions of light
has been lost, and before the organ has been far
reduced by phylogenetic destruction, a veil of
black pigment is formed over it, completely shut-
ting it off from the outer light. The nerve
disappears completely before birth, its degenerate
cells becoming lost in the mesoblastic skeletal
tissue of that region. At the time of birth
the whole eye is enclosed in a thick membrane
which isolates it. The deposition of pigment has

THE PATH OF DEGENERATION IN ANIMALS 185

destroyed any functional activity in the lens and the
retina, but these parts none the less retain traces
of a complicated structure recalling their condition
when functional.

In the Rhynchocephala and Lizards examined
by W. B. Spencer,1 there is to be found a series of
types representing the various stages of the
degeneration of the eye in the slow-worm.

In the type Sphenodon {Hatteria, fig. 62, a), the
organ in the adult is in the complete form. The
eye possesses a lens (c), a retina, (?*), with com-
plicated histological structure. A nerve (n) places
the retina in communication with the brain.

In Chamceleo (fig. 62, b), the degradation of the
organ has reached the following stage : the epi-
physis has a distal portion corresponding to the
eye, but the histological differentiation of this is
incomplete, neither the retina nor the lens being
distinct. Nervous fibres connect this with the
proximal portion which is hollow and in com-
munication with the brain. It thus represents
the second stage in the formation of the eye in
the slow-worm.

In the types Leiodera and Ccdotes (fig. 62, c), the
chief degeneration is in the optic nerve, which has
partially disappeared, and no longer connects the
eye with the brain. The eye itself is not quite

1 W. B. Spencer, On the presence and structure of the pineal
eye in Lacertilia (Quarterly Journal of Microscopical
1886).

H

^r\r:

186 THE PATH OF DEGENERATIVE EVOLUTION

so degenerate : the retina has not a complex
structure, but both retina and lens are present.

In the type Cyclodus (fig. 62, d), the degeneration
is still greater : the epiphysis is a vesicle attached
to the thalamencephalon. The walls of this
vesicle show only the smallest symptom of primi-
tive differentiation into lens and retina. The
proximal part remains hollow, and shows no trace
of differentiation at all.

This, however, is not the only fashion in which
degeneration of the pineal eye proceeds. In
another series of creatures it retains its connec-
tion with the thalamencephalon, but remains inside
the skull. The parietal foramen closes, thus
completely shutting off the eye from the light ;
the eye becomes useless, degenerates, and the
optic nerve loses its function as a conducting
channel, Ceratophora (fig. 62, e). The pineal
organ then becomes a degenerate structure in
which it is exceedingly difficult to see traces of
its original condition, and which is usually marked
by an abundance of blood-vessels : Birds (fig. 62, f),
Mammals (fig. 62, g).

Thus, the degeneration of the pineal eye shows
that the optic nerve, the last organ to be completed,
is the first to disappear.1

In studying the degeneration of the pineal eye,

1 The same happens in the case of the ordinary paired eyes. In
the degenerate eyes of the mole, the optic nerve is more reduced
than are the other structures.

THE PATH OF DEGENERATION IN ANIMALS 187

we have seen the path of progressive evolution
being retraced, at least in the case of these reptiles.
The present state of knowledge does not permit an
extension of the enquiry to the conditions found in
higher animals. However, according to Flesch,
traces of a sensory epithelium have been found in
the pineal body of man, the horse, the sheep, and
the bat : that is to say, that in these creatures too
the oldest parts of the structure have resisted
degeneration longest. We cannot refrain from the
conclusion that in this series degeneration retraces
to a large extent the steps of original advance.

2. Degeneration of the organs of sight in deep-
sea Crustacea. — We cannot however establish the
conclusion of the last paragraph as a general
principle.

The fauna of the deep sea includes a large
number of Crustacea, and in these the eyes, which
are relatively useless, are often degenerate. The
course of the degeneration is generally definite, and
of all the structural parts the most long-lived are
the eye-stalks, although we know that these are
a recent formation. A number of examples chosen
from Decapod Crustaceans, which are specially
abundant, will illustrate this point.1

Nephropsis, which lives in the Atlantic and
Indian Oceans at moderate depths, is a relative of
the Lobster. The optic stalk is short and carries

1 See Pelseneer, V Exploration zoologique den mers profondcs
(Conferences Universitaire de Bruxelles). 2 Annfo, 1890.

188 THE PATH OF DEGENERATIVE EVOLUTION

a rudimentary eye which has neither pigment nor
cornea and is coloured like the general surface of
the body.

Eryonicus (fig. 63) belongs to the same group,
and comes from the region of Saint-Thomas in the
Antilles, where it lives at a depth of about 825
yards. This animal has a reduced optic stalk,
but at the extremity of this, where in littoral

forms the eye
is borne, there
is only a de-
pression as if
the eye had
been carefully
scooped out.

Willemcesia
(fig. 64), a re-
lative of the
marine cray-

Fig. 63. Eryonicus cozens. Bate? (After W. Faxon, fish and ail ill-
The Stalk-eyed Crustacea, Mem. of Mus. of Comp.,
Zool. Harvard College, vol. xviii., 1895.) habitant of the

Atlantic at a depth of about 3500 yards, is com-
pletely devoid of eyes in the adult condition,
although it possesses them in the larval stage.

ScolopJithalmus (fig. 60), which lives down to
4000 yards, is quite devoid of eyes, but possesses
eye-stalks which terminate in spines.

It seems, then, that different species of deep-sea
Crustacea may present different degrees of degenera-
tion of the eye. One species in itself exhibits all

Kii,. 64 —Willematia, (After Pelseneer.)

100 THE PATH OF DEGENERATIVE EVOLUTION

grades of degradation according to the depth at
which it lives. This creature — Cymonomvs —
which, when near the surface, has fully formed
eyes upon movable stalks, at a depth of a few
hundred yards exhibits movable stalks without
eyes; and at 1500 yards the stalks are fixed and
end in spines.

Isopod Crustacea, which live in the deep sea,
present similarly degenerate eyes. Many are blind

Fig. 65. — Seolophthatmus ludfugui, Fax.

a, optic peduncle transformed to a spine. (After W. Faxon, The Stalk-eyed
Crustacea, Mem. of Mus. of Comp. Zool. Harvard College, vol. xviii., 1895.)

and display all kinds of optic degeneration. Noesa,
for instance, simply has eyes devoid of pigment.
Thus, in abysmal Crustacea, the degeneration of the
eyes is in no sense a retracing of developmental
stages.

Another instance chosen from examples of the
atrophy of organs in individuals, shows that the
supposed law of retracing cannot be made uni-
versal.

4. Atrophy of the branchial vessels in man. —
Examination of a human embryo of about three

THE PATH OF DEGENERATION IN ANIMALS 191

weeks old allows the presence of a series of slits on
the sides of the neck, the slits not being parallel,
but converging towards the ventral surface. Be-
tween these slits are swellings, or pads, which pass
up towards the dorsal surface and appear like the
beginnings of hoops or ribs enclosing the visceral
cavity ; the elevations are the branchial arches, the
slits are the gill-slits.

In the human embryo (fig. 66, A), as in fish,
these slits appear

from above
downwards, and
as they are
formed, the cor-
responding blood-
vessels arise.

These vessels,
or aortic arches,
arise from a ven-
tral aorta (a.)
which gives off
six lateral
branches (c.) at each side. These lateral branches
run up between the gill-slits and form two main
trunks on the dorsal side which converge to form
the descending aorta (ad.).

In man the branchial arches are transformed,
parts of them entering into the structure of the
face, and during the transformation parts of the
aortic arches atrophy (fig. 66, b). But the order

Fig. 66. — Diagram of branchial arches in mammals.

A. Embryonic stage, a, aorta; c, aortic arches;
AD. dorsal aorta. B. Adult stage. The parts
represented by dotted lines have degenerated.
A, aorta; v, carotid; ap, pulmonary artery.

192 THE PATH OF DEGENERATIVE EVOLUTION

of this atrophy does not correspond in any way to
the order of the formation of the vessels.

The median parts of the anterior two lateral
branches (m1 and m2) disappear, and the vertical
parts remain as the internal and external carotid
vessels. The vertical piece which joined the
posterior parts of the third and fourth arches dis-
appears : the internal and external carotids thus
acquire a stem of their own. The parts of the
fourth arch remain ; the fifth arch disappears at
each side, and the sixth arch forms the pulmonary
artery (ap.).

Thus the degeneration of these vessels represents
in no way whatever a retracing of their develop-
mental history. All that occurs is that the useless
parts disappear and the useful parts persist. A
comparative study of this example would only
enforce our conclusion.

In ontogeny the neurapophyses are more ancient
than the vertebral centres. None the less, as we
have already seen, the examination of any vertebral
column from head toward tail shows a gradual
disappearance of all parts except the centra,
although the centra are the last to be formed.

Section II.

The path of degeneration in plants.

] . Rarity of cases of recapitulation in the
organogeny of leaves. — We have already said that

THE PATH OF DEGENERATION IN PLANTS 193

recapitulation seldom occurs in plants, the develop-
ment of the whole and of its organs being usually
direct. When it does occur, it is generally limited
to characters coming from comparatively recent
ancestors and not even in the most transitory
form entering into the formation of the fundamental
parts of the plant.1

1 The rarity of recapitulation among vegetables is the result
partly of their fixed condition in the soil, and partly of the more
rigid nature of their cells.

The immobility of a plant forces the adult to live in the same
place as the embryo. Among animals, on the other hand, it
frequently happens that the young pursue a manner of life
different from that of the adult and resembling that of the
ancestor. Young Cirrepedes are vagrant and have the same
needs and use the same organs as other vagrant Crustacea ; larval
frogs inhabit the water like their fish-like ancestors. In plants
there is nothing similar ; all the aquatic flowering plants are
derived from terrestrial ancestors, but if at the beginning of their
existence these aquatic plants were to bear leaves adapted to
aerial life they would ensure their own destruction. The ex-
ceedingly rare ancestral traits to be found in 'a few species are
naturally of a kind not to incommode their possessors. It is
improbable that these are a legacy from distant ancestors ; they
would not have been spared by natural selection had they not
come from ancestors of very much the same habit. The absence
of locomotion in plants has also produced a greater adaptability
than among animals. Animals, when conditions are unfavourable
can remove in search of more suitable localities, plants being fixed
in the soil must become modified or perish. Plants, therefore,
offer numerous cases of individual adaptation. We do not know
if these adaptations are transmitted by heredity,' but natural
selection has at least secured the widest range of plasticity.
Thus plants rapidly rid themselves of ancestral legacies which
have become useless.

The transitory organs of animals are employed for the service
N

194 THE PATH OF DEGENERATIVE EVOLUTION

In consequence, vegetable embryology is of little
use for investigation of the supposed backward path
of degeneration, for the rudimentary or reduced
organs of plants do not generally represent ancestral
stages.

The seedling of Lathyrus tenuifolius (fig. 67), a
vetch, possesses rudimentary organs which cannot
be ancestral stages as their development is direct.
In this plant a whole series of leaves are formed
between those arising at germination and the adult
leaves. This intermediate series displays many
arrests of development.

The adult leaf has a pair of stipules, foliage
leaflets, and tendrils (fig. 67, J). The leaves just
before these, have a pair of stipules (fig. 67, i),
which are absent in the leaves next before (iig. 67,
h). Still earlier leaves are produced with fewer
leaflets and tendrils (fig. 67, d-g), leaves without
leaflets and with a single tendril (fig. 67, c), and
leaves entirely without tendrils (fig. 67, b). Lastly,
at germination very rudimentary leaves are pro-
of the whole body, the branchial arches of mammals are employed
in the formation of important parts of the head and neck. The
tail of the tadpole is reabsorbed by phagocytes and its substance
used for the nutrition of the body. In the case of plants, such
occurrences are rare and limited ; the cells are enclosed in a rigid
wall which resists displacement or alteration ; the protoplasmic
contents may be absorbed and used as nutritive material by
another part, but the cellulose cell-wall remains. A useless organ
can be eliminated only at the expense of loss of material.
J. Massart, La Recapitulation et V Innovation en embryologie
vigitale {Bull. Soc. Roy. Bot. Belg., t. xxxiii., p. 150, 1894).

Fio. 67. — Latfiyrus tenuifolius.
A, B, seedlings In two stages of growth. C to J, different forms of successive leaves

196

THE PATH OF DEGENERATIVE EVOLUTION

duced, the sole function of which is to protect
the axillary bud (fig. 67, b, 1, 2). We have now
to show that this series is by no means a retracing

of ancestral stages.
All, or nearly all,
the Papilionace-
ous plants have
stipulate leaves ;
this shows that
the exstipulate
leaves of L. tenui-
folius do not re-
present an ances-
tral stage. More-
over, before the
acquisition of
tendrils, vetches
had a terminal
leaflet (see Vicia
Pyrenaica, fig.
44) ; none of the
reduced leaves in
L. tenuifolius re-
produce this

Fig. C8. — Seedling Vicia of monanthos. sta^e ' moreover

the simple leaves without leaflets do not represent
an ancestral condition ; the winged petiole is not an
ancestral character.

The primary leaves of another vetch Vicia
monanthos confirm our conclusion. In this case

THE PATH OF DEGENERATION IN PLANTS 197

the first few leaves formed in the seedling remain
rudimentary, aud serve only to protect the axillary
buds (fig. 68, leaves 1 and 2). Contrasted with
the condition in L. tcnwifolius, although all the
vetches had probably a common ancestor, these
primary leaves have three little projections, the
two lateral of which are reduced stipules.

We do not know of any vegetable example of
recapitulation in the case of an organ reduced by
arrest of development. Plants exist, however,
which after having produced leaves of typical
structure begin to produce leaves the development
of which remains arrested. Such a plant is the
Acacia which bears phyllodes. At first the leaves
are like those of other Acacias) next it bears leaves,
the blades of which are rudimentary ; and finally
leaves with normal stipules, but with no trace of
lateral leaflets on the petiole.

In this case the ancestral conditions are known
and are quite different.1

2. Organogeny of flowers. — What we have said
about leaves applies to the organogeny of flowers.
Here also in the cases of atrophy produced by
arrest of development there is no indication of

1 Many other reduced plant organs might be instanced, such as
the stipules of Sambucus or the teeth of the calyx, in many Com-
posite and Umbelliferre. After their formation such organs grow
very slowly and exhibit no trace of recapitulation. This happens
with the leaves of Sempervwum (fig. 49). Without doubt these
leaves are derived from leaves normally divided into hypopodium
and epipodium, but they show no trace of this division.

198 THE PATH OF DEGENERATIVE EVOLUTION

ancestral stages, and thus there is no evidence that
degeneration retraces the path of progressive evolu-
tion.

In the cauliflower (Brassica oleracea, var. Botrij-

tis), a cultivated
variety, the
inflorescence
branches ex-
uberantly; most
of the flowers
produced on
these branches
are arrested in
their develop-
ment. Of the
immense num-
ber of flowers
produced on
each plant only
a few attain
sexual maturity
and produce
seeds; the others
abort at differ-
ent stages. Most of these remain in a very primitive
stage and do not develop sepals ; but, scattered
among them, may be found more fully developed
flowers, so that the same plant presents almost
every possible stage of flower development.

The organogeny of these flowers is quite like

Fig. 69. — Development of flowers of (he Cauliflower.

sa, anterior sepal ; sp, posterior sepal ; si, lateral
sepal ; p, petal; «, long stamen ; ec, short stamen ;
o, ovary. A, very young flower with only rudi-
ments of two sepals; B, flower with rudiments of
four sepals ; C, older flower with rudiments of
petals; D, still older flower with rudiments of
stamens and ovary. (The flowers A — D are seen
from the hide. In the figures C and D the lateral
sepal has been removed.) E, flower seen from
behind; the posterior sepal has been removed.

THE PATH OF DEGENERATION IN PLANTS 199

that of other Cruciferae. If in the cauliflower the
arrested flowers corresponded to different ancestral
stages the case would be striking ; but this does
not occur. Primitively the flower contained only
the stamens and pistils, the essential organs of
reproduction. But we see that the cauliflower
produces first the anterior and posterior sepals
(fig. 69, a), then the lateral sepals (fig. 69, b), then
successively the petals (fig. 69, c), the four larger
stamens (fig. 69, d), and the two shorter stamens
(fig. 69, e). Moreover, the flowers display a special
readiness to the suppression of certain parts such
as the petals, as in the flower d (fig. 69).

3. Progressive degeneration of the prothalhis in
phanerogams. — rAlthough embryology gives us few
examples, morphology proves clearly enough that
in plant degeneration there is no return to ancestral
types.

This appears clearly from a comparison of the
progressive evolution of the prothallus in crypto-
gams with its degeneration in phanerogams.

Terrestrial vegetation has been derived entirely
from aquatic life. The Bryophyta (mosses and
liver-worts), the Vascular Cryptogams (ferns), and
the flowering plants all have sprung from aquatic
algae probably not very different from Coleochcete.
Such aquatic forms are reproduced by means of
true ova and spermatozoa. The terrestrial plants
which were derived from them had also spermatozoa
with vibratile locomotor hairs and impregnation

200 THE PATH OF DEGENERATIVE EVOLUTION

took place in a fluid. A special organ, the arche-
gonium, was developed ; this contained the ovum
and made the approach of the spermatozoa more
easy.

For the present purpose we may omit considera-
tion of the mosses and liver-worts as it is improb-
able that they are in the line of ancestry of the
flowering plants. It is necessary only to say that
in them, while a single egg is produced in the
archegonium a large number of spermatozoa are
produced in the antheridium.

The same condition is found among the fern-like
plants, but in their case, owing to the development
of special channels for the passage of nutritive
materials, it is possible for a much greater size to
be reached. In these circumstances it would be
unlikely that the spermatozoa should find at the
summit of a comparatively lofty plant the drop
of water necessary for the task of fertilization.1
Accordingly the sexual cells are produced on
prothalli, which hardly reach above the surface of
the ground, and are in a favourable position for the
necessary moisture.

In the less specialized ferns (fig. 70) a single
prothallus bears both male and female organs
(antheridia and archegonia). In the Equisetums

1 The smaller forms like the Selaginellidae and the club-mosses
were represented in the past by plants of much greater size, as is
seen from fossil remains, and it is probable that the modern forms
have descended from these giants.

THE PATH OF DEGENERATION IN PLANTS

201

the prothalli are unisexual, but the spores from
which they are produced are alike. Finally, in the
allies of Selaginella which are the Cryptogams most
nearly approaching flowering plants, the prothalli
which bear male organs
are quite distinct from
the prothalli which bear
female organs ; more-
over, the spores which
give rise to the two
kinds of prothalli are
quite different. A
small number of large
spores arise in special
sporangia, termed
macrosporangia, and it
is from these that the
female prothalli arise.
A larger number of
smaller spores are pro-
duced in microspor-
angia and these micro-
spores produce the male
prothalli.

The spores of ferns are very small and contain
only the nutritive matter required for germination ;
the spores of Selaginella, even the microspores, are
very much larger and contain a large supply of
reserve material at the expense of which the
development of the prothallus takes place.

Fig. 70. — Seedling of Fern (Aspleniutn
Filix-foemina.

The prothallus (j>) bears antheridia at its
edges and towards the centre an arehe-
gonium which has been fertilized. The
young seedling developed from the
fertilized ovum has already given rise
to roots and to two leaves.

202 THE PATH OF DEGENERATIVE EVOLUTION

We may give a brief account of the formation of
the sexual cells in Selaginella. The first stages in
the appearance of microsporangia and macro-
sporangia are identical. Later on in the one case
each of the spore mother-cells breaks up into four
microspores, in the other case only one mother-cell
breaks up into four macrospores, the others dis-
appear. On germination the macrospore becomes a
fairly large prothallus with several archegonia : the
microspore forms only one cell as the representative
of the prothallus, the rest of the structure becoming
the antheridium.

In Phanerogams (flowering plants) a drop of
water is no longer necessary for fertilization. The
spermatozoon reaches the ovum, not by swimming
movements, but by a kind of growth. This
difference has resulted in profound modifications.
Fertilization may occur in the air, and it is no
longer necessary for the macrospore to become
detached : it remains fixed on the individual from
which it arose and there proceeds to develop.
In the Gymnosperms the archegonia are fully
formed in a prothallus of considerable size, but the
Angiosperms which are more specialized, retain
only traces of these structures. The microspores
(pollen grains) exactly as in Selaginella are pro-
duced in great numbers, but when they leave the
parent plant they develop only if they reach suit-
able conditions. In germination there is shut off,
as in Selaginella, a single cell to represent the

THE PATH OF DEGENERATION IN PLANTS 203

prothallus ; later on in a few Gymnosperms it
undergoes a few divisions representing the forma-
tion of the antheridium. In Angiosperms the
reduction has proceeded further, and each pollen
grain besides the prothallus cell produces only one
or at most two reproductive nuclei. In all
Phanerogams the reproductive nucleus reaches the
ovum by being carried in a long tube which grows
out from the pollen grain at the expense of
nutriment derived from the female tissues.

Thus, the prothallus may be traced through
marked stages of reduction from the condition in
ferns, through Selaginella to Gymnosperms and
higher flowering plants. But these phases of
reduction are by no means to be regarded as
repetitions of ancestral conditions.

In conclusion, it is plain that we cannot assert
as a general law, that degeneration retraces the
steps of evolution. In the vegetable kingdom no
facts support such a conclusion. In certain special
cases in the animal kingdom, the most recently
developed structures are the first to disappear when
atrophy sets in, but this proves no more than that
these particular structures happened to be less
stable, and to offer less resistance. It is quite
impossible to make such facts support the generally
held view, that degeneration is a kind of inverse of
evolution.

It is very seldom that a living apparatus with
complex functions loses all of them, but usually

204 THE PATH OF DEGENERATIVE EVOLUTION

preserves one or other ; for instance, the leaves of
parasitic flowering plants continue to serve as
protectors of the buds, and degeneration does not
affect the part that has remained functional. It
is highly probable that the original function of
leaves was assimilative rather than protective, and
yet here it is the later function that is retained.

Even when all function is lost, and the whole
structure degenerates, there is no reason why the
degeneration should retrace the evolution. In the
case of the atrophy of an organ in an individual
especially in such pathological instances as those
mentioned by Ribot, it may be that the latest
formed parts are the most fragile, and the most
ready to disappear ; but the path of atrophy is
quite different in the case of the gradual reduction
of an organ in a species. When an organ becomes
useless to a species, as in the case of the eyes of
deep-sea Crustacea, the only thing that matters to
the species is that it may be got rid of. Any
individual variation tending towards reduction will
be of advantage, and may be retained by natural
selection. There is no reason to suppose that such
individual variations appear in any inverse order ;
in fact we do not know that the appearance of
variations follows any law at all. Perhaps the
apparent inverse order of the degeneration of the
pineal eye in lizards may be explained from the
fact that the most recently acquired characters are
frequently the most variable.

THE PATH OF DEGENERATION IN SOCIOLOGY 205

However, even when the most recent organs
disappear first, we cannot speak safely of a law
of degeneration inverse to evolution. In the
actual evolution of many organs, parts have
appeared and then completely disappeared. If
degeneration were a true inverse of evolution,
it would be necessary that such lost parts should
reappear only to disappear again. Such observa-
tions apply both to ontogeny and phylogeny.

CHAPTEE II

THE PATH OF DEGENERATION IN SOCIOLOGY

§ 1. Investigation of facts.

We have now to see if degenerative evolution in
social matters retraces the steps of progressive
evolution.

In the first place the question cannot be even
entertained with regard to some cases, and this for
a general reason which will be dealt with later on.

When a complex institution — such as a com-
mercial society or an administrative organization —
becomes useless and ceases to be functional, it usually
disappears either by voluntary dissolution or else it
is legally suppressed. In either case there is no
slow retrogressive degeneration retracing inversely
the steps of progress, for all the parts cease to

206 THE PATH OF DEGENERATIVE EVOLUTION

exist simultaneously. If certain parts of the sup-
pressed institution are allowed to persist, these are
by no means necessarily the oldest parts, but quite
the contrary.

When, for instance, the Provincial States of
Dauphiny and Normandy were suppressed by the
French monarchy, only the titles with their cor-
responding emoluments were allowed to remain, and
they were obviously of more recent origin than the
States themselves.

It must be borne in mind that all the parts of
an institution rarely become simultaneously useless
and non-functional. Those which retain their utility
longest are by no means always the most ancient in
origin.

English sheriffs have gradually become of less
and less functional importance; and now fulfil no
other role than that of presiding over elections and
accompanying the judges when on circuit. Both of
these functions have been acquired recently com-
pared with all those which the sheriff discharged in
the days when the care and protection of the whole
county practically devolved upon him.

The question then of the pathway of degeneration
only arises in those cases where the same cause
of dissolution simultaneously affects all parts of
the institution, and where, without sudden inter-
ruption, degeneration is effected slowly but surely
through many successive stages. This, of course,
happens in the degenerative evolution of individual

THE PATH OF DEGENERATION IN SOCIOLOGY 207

societies or institutions, and not in the disappearance
of complete classes of institutions.

These reservations being understood, we will
mention a few more or less obvious cases in which
degeneration does retrace the footsteps of progressive
evolution.

1. Tithings, hundreds and counties in England. —
In the chapter dealing with the pathway of degenera-
tion in Transformisme social, G. Degreef mentions
the following interesting facts : —

" Mr Herbert Spencer, after describing the forma-
tion of tithings, hundreds and counties in England
under the Anglo-Saxon regime, observes that the
tithings along with their courts of justice were the
first to disappear, then the hundreds followed, though
some vestiges of their old courts of justice remained,
and only the counties and the county courts were
left intact. Now we have historical proofs that
English counties along with their courts of justice
were created before the hundreds, and the hundreds
before the tithings." x

2. Order of elimination of various racial elements
in a country. — In his interesting work Civiliza-
tion et de'popidation,2 Dumont mentions certain
facts which go to show that the inhabitants of poor
districts, who are nevertheless of pure racial descent,
have a birth-rate higher than that of the members
of the population who are not aboriginal, and who

1 Degreef, Le transformisme social, p. 450.

2 P. 156.

208 THE PATH OF DEGENERATIVE EVOLUTION

for the most part dwell in the towns and fertile
plains. From this he concludes that the various
racial elements of a nation are eliminated in inverse
order to that in which they were introduced. In
France, for instance, the Frank has been com-
pletely absorbed in the Gaul.

3. TJie degenerative evolution of political organiza-
tions.— The progressive and degenerative evolution
of political organizations has been described by
Herbert Spencer as follows 1 : —

" Political integration, as it advances, obliterates
the original divisions among the united parts. In
the first there is the slow disappearance of those
non-topographical divisions arising from relation-
ship, as seen in separate gentes and tribes —
gradual intermingling destroys them. In the
second place, the smaller local societies united
into a larger one, which at first retains their
separate organizations, lose them by long co-
operation ; a common organization begins to
ramify through them. And, in the third place,
there simultaneously results a fading of their
topographical bounds, and a replacing of them by
the new administrative bounds of the common
organization.

" Hence, naturally, results the converse truth that
in the course of social dissolution the great groups
separate first, and afterwards, if dissolution con-

1 Herbert Spencer, " Political Institutions," Part iv. of Principles
of Sociology, p. 286.

THE PATH OF DEGENERATION IN SOCIOLOGY 209

tinues, these separate into their component smaller
groups. Instance the ancient empires successively
formed in the East, the united kingdoms of which
severally resumed their autonomies when the coer-
cion of keeping them together ceased. Instance
again the Carlo vingian Empire which, first parting
into its large divisions, became, in course of time,
further disintegrated by subdivisions of these. And
when, as in this last case, the process of dissolution
goes very far, there is a return to something like
the primitive condition, under which small preda-
tory societies are engaged in continuous warfare
with like small societies around them."

We may conclude then that political integration
is attended by degeneration ; primitive institutions
disappear and make way for fresh institutions, and
their disappearance is permanent. In the course
of the dissolution of the Carlovingian Empire
there was no reappearance either of the gentes
or of the primitive tribal system ; but when this
vast organization broke down, it was natural that
the more recently formed social bonds, having had
the least opportunity of becoming consolidated,
should be the first to be sundered.

4. Degeneration in monetary systems. — The prin-
ciple that degeneration retraces the steps of pro-
gress applies equally to a very different range of
ideas, — the evolution of monetary systems. Stanley
Jevons says that there is little doubt that every
system of coinage was originally identical with a

o

210 THE PATH OF DEGENERATIVE EVOLUTION

system of weights. A survival of this primitive
condition existed in Eoman law, and even when no
use was made of them, the custom of bringing a
pair of scales survived as a legal formality in the
sale of slaves at Eome.

After the time of the Punic wars, the ass,
which originally equalled a Eoman pound in
weight, diminished rapidly, until it became re-
duced to the weight of an ounce. The Eomans
had naturally reverted to weighing the metal, and
the ces grave was money reckoned by weight, and
not by tale. Generally speaking, whatever be the
inconveniences of the method, currency by weight
is yet the natural and necessary system to which
people revert whenever the abrasion of coins, the
intermixture of currencies, the downfall of a State,
or other causes, destroy the public confidence in a
more highly organized system.1

It is plain then that the more recent developments
in the coinage system are the first to disappear.2

The disappearance of money altogether and the
return to a system of exchange would represent a
much farther advanced stage in degeneration.

1 See Stanley Jevons in Money, International Scientific Series.

2 There is no silver money and only a little copper in China.
Nowadays, Mexican piastres, on reaching the country in payment
of commercial transactions, are melted down into bars as soon as
they fall into the hands of the merchants, and these bars are then
imprinted with the Chinese stamp. This was the usual system
employed amongst civilized peoples before the invention of money.
See Thorold Rogers in ' ' The Economic Interpretation of History. "

THE PATH OF DEGENERATION IN SOCIOLOGY 211

5. Degenerative adaptation in colonial legislation.
— In his treatise (Annalisi delta proprieta capita-
lista), Loria furnishes another striking example of
the law of degeneration : " When English colonies
were first formed in America, the colonists hesitated
to establish any legislation other than that of the
mother-country. They were habituated to it ; it
was written in their own language, and therefore
seemed best to correspond with their national
characteristics. But, from the outset, the greatest
difficulties were met with in the application of this
legislation to the colonies.

" In the first place the Statute law of England, the
most recent addition to the legislation, was found
to be quite unsuited to the economic condition of
a colony, and so common law alone came to be
established, which, being the more ancient, was
better suited to the social organization of a newly-
formed society. But even this form of legislation
did not remain permanent under social conditions
profoundly different to those in which it had been
originally established, and the construction of a
special legislation was found to be necessary. In
this way the common law of England came to be
regarded as unsuited to her colonies, excepting in
such cases as were unprovided for in the new
colonial law." *

Loria then proceeds to give numerous examples
of how these colonial statutes — owing to the simi-
1 Loria, Annalisi delta proprieta capitalista, ii. 48.

212 THE PATH OF DEGENERATIVE EVOLUTION

larity of circumstances between those for whom
they were severally fashioned — came to resemble
the primitive law of England.

6. Degenerative evolution of the corporations of
Western Flanders. — In a treatise published by one
of our number in May 1892, entitled I' Evolution
regressive des corporations de la West-Flandre} it
was shown that among these associations, the insti-
tutions the first to degenerate were those most
recently established.

The twelve corporations which still exist, though
in more or less degenerate conditions in Bruges,
Fumes, Eeghem, and Iseghem, were formerly con-
structed on similar principles and fulfilled the
following functions : —

1. The furtherance of sociability (i.e. the holding

of banquets and fetes).

2. The encouragement of religious feeling (i.e.

frequent celebrations of the mass and the
building of new churches).

3. Mutual assistance (i.e. insurance against loss

of work through illness, or against funeral
expenses).

4. Mutual protection of professional interests.

5. The furtherance of certain political and mili-

tary interests.
These various functions were established not
simultaneously but in succession. The Flemish
corporation of the fourteenth or fifteenth century
1 La SocUU nouvelle, Mai 1892, Bruxelles, Moimom.

THE PATH OF DEGENERATION IN SOCIOLOGY 213

represents the last stage in a long series of corpora-
tions of different kinds, and of increasing complexity.
It will be as well to glance briefly through these
various kinds of corporations in order to compare
the different stages of dissolution through which the
corporative system eventually passed with the pro-
gressive evolution exhibited in the course of its
establishment.

(a) Associations formed for the holding of
banquets and fetes (cojivivia) which were origin-
ally distributed throughout Northern and Western
Europe.

(b) The convivia assumed a religious character
when the Church, unable to suppress them, deter-
mined to transform them.

(c) Guilds (etymologically significant, according
to Brentano, of repasts where all expenses were
shared) where the original conviviam was accom-
panied by religious ceremonies, but a feature of
which was the addition of institutions of mutual
insurance.

(d) Corporations (ambachtcu). — These embodied
all the primitive institutions that had gone before :
(1) the banquets; (2) religious ceremonies; (3)
mutual assistance ; (4) fresh means for the protec-
tion and development of professional interests.

(e) Finally, at the commencement of the four-
teenth century, these corporations assumed both
political and military functions. The concession of
keure to the people in 1304 by Philip of Thiette

214 THE PATH OF DEGENERATIVE EVOLUTION

was the first act by which was recognized the right
of the corporations of Bruges to take part in com-
munal administration and to provide a military
contingent of their own.

When these corporations began to decline mili-
tary and political functions were the first to be
eliminated, in other respects the corporation con-
tinued to exist till the close of the eighteenth
century in the form of economic groups, and groups
for mutual aid and religious development.

An investigation of such corporations as still
survive shows that it is in the economic functions
formerly discharged that degeneration has made
furthest strides, while the religious character is
maintained and banquets are still held. Of the
twelve corporations of Western Flanders six preserve
the old corporative institutions almost intact ; seven
maintain a scale of charges and regulations con-
nected with them ; ten provide organizations for
assurance against illness ; eleven hold annual re-
ligious ceremonies ; twelve, i.e. all of them, continue
to hold banquets.

These figures alone show that the original and
earlier functions have remained longest in force, but
to further demonstrate this point it will be necessary
to enter more fully into details and to study each
group separately.

The first group — consisting of associations con-
tinuing to protect professional interests — consists of
three branches : the four offices of Bruges (including

THE PATH OF DEGENERATION IN SOCIOLOGY 215

vendors of lime, coal, seeds and beer), the two offices
of Fumes (street porters and vendors of beer), and
the community of the bakers of Bruges.

All the ancient statutes of the four offices have
been preserved, including trade monopolies, the
freedom of the city, and scales of charges, help
in times of illness, accident, or when out of work ;
religious ceremonies and banquets. With the com-
munity of bakers, however, this is not the case ;
here the economic functions discharged are reduced
to a minimum. The association continues to exist
in a triple capacity : as a syndicate to keep up a
fair price in wheat and bread, as a mutual assur-
ance association in times of distress among the
members, and as a confraternity imposing religious
obligations and holding an annual banquet and fete.

In the second group consisting of mutual aid
associations such as the corporations of tailors, shoe-
makers and weavers of Bruges, all trade interests
have disappeared, and the corporations only exist
in the capacities of confraternities and mutual aid
societies. At this particular stage of degeneration
these corporations resemble in a striking degree the
old guilds which preceded the ambachten en neringen.
It is interesting in connection with this, that the
corporation of wool weavers (wollewevers) in Bruges
has lost its orginal professional character, and quite
heterogeneous elements have been introduced ; there
are only twenty-five weavers in Bruges, and their
society numbers nearly two hundred members. The

216 THE PATH OF DEGENERATIVE EVOLUTION

guild of weavers in Eeghem represents the third
stage only — that of a religious confraternity. The
office of mutual aid has disappeared, and only the
Saints' Day Fete and the convivium remain : " The
first Sunday after the fete of the Trois-Bois, which
is the annual fete day of the weavers, the guild
proceeds to the parish church, headed by a banner,
accompanied by a jester, and with drums beating,
where mass is celebrated in honour of Saint Severin.
The rest of the day is passed in diversions."

The guilds of Saint Crispin at Iseghem, all that
remains of the old corporation of shoemakers, are
representative of the final stage of degeneration :
the association has resumed its most primitive
character, and is reduced to a mere dining society—
the primitive convivium.

There are from 1500 to 2000 shoemakers in
Iseg-hem. At the time of the Be volution their
ambacht, having resumed its archaic form, was
divided up into six or seven guilds. Some years
ago a vain attempt was made to reconstruct these
guilds and adapt them to modern requirements.
At a time when success in this project was still
hoped for, one of the promoters of the reconstruction
wrote as follows : —

" The members of the guild still recognize Saint
Crispin as their patron saint, but they no longer
assemble in church to do him honour by the cele-
bration of mass. They keep the anniversary instead
by going from one public-house to another, with

THE PATH OF DEGENERATION IN SOCIOLOGY 217

flying banner and drums beating, and the day has
become merely a day of copious libations, and serves
as a pretext for poor workmen, the fathers of families,
to spend all the week drinking in public-houses."

In conclusion then the cycle is complete ; a
corporation, unless suddenly dispersed, ends as it
began ; in the last stage of decline it resembles
the associations from which it originally developed,
the most recently established functions having been
the first to decay and disappear.

§ 2. A criticism of the supposed inverse path of
degeneration.

These few examples suffice to show that in
certain cases the more recently formed institu-
tions are the first to decline and disappear, while
the older persist to the end.

It must be remembered, however, that the con-
trary is at least as frequently the case. All changes
of legislation, either juridical or religious, follow,
but never precede, the economic transformations to
which they relate, whether these be social or ethical,
unless the transformations are ephemeral. " Imita-
tion," says Tarde, " proceeds from the more obvious
to the less obvious ; that is to say, ends and feel-
ings are imitated sooner than their means and
expressions."

Title-deeds and armorial bearings survive no-
bility ; houses continued to be held as personal
or moveable property long after the disappearance

218 THE PATH OF DEGENERATIVE EVOLUTION

of nomadic tribes, which, living as they did in
tents, originated the conception.1

Among peoples where the system of marriage by
groups has existed, family nomenclatures continued
to persist long after the disappearance of the family
system to which they owed their origin. " The
family," says Morgan, " is an active element, never
stationary ; it keeps pace with the development of
society in the march of progress. On the other
hand, the reckoning of kinship changes very
slowly ; only after long lapses of time does it
register the progress actually made by the family
in the course of ages, and does not undergo any
radical transformation until long after the family
itself has been completely changed." " And," adds
Karl Marx, whose critical annotations on Morgan's
book were carefully preserved by Engels, " this also
applies to systems of politics, law, religion, or
philosophy."

These systems, formed after the completion of
the social organization which they express, survive
after the organization itself has disappeared. Their
elimination is not of such importance to society
as is that of the economic or family institutions
themselves, as these, when they become useless and
disadvantageous, are a drag on future development.

1 Viollet, Hisfoire du droit civil //'., p. 617.

"Although houses were for centuries treated as moveable pro-
perty, they continued to be legally treated as such for a still
longer period of time ; it is characteristic of judicial ideas that
they lag far behind economic progress."

THE PATH OF DEGENERATION IN SOCIOLOGY 219

It cannot be established, however, as a general
principle, that the pathway of degeneration as
regards societies or institutions, is inverse to that
pursued by their progressive evolution. In the
first place, the mere explanation of this supposed
law shows that it is quite untenable.

What reason is given for supposing the decline
of memory or will power, the degeneration of writ-
ing and speech, the decadence of societies and
institutions, to be a retracing of the steps of
progressive development ? The reason given is
that, other things being equal, the more fragile,
unsteady and complicated structures are the first
to fall.

Now, although the most fragile structures are
frequently those most recently formed, and which
have not had time to settle down and firmly estab-
lish themselves, it is also true that in many cases
the more recent acquisitions and structures attain
a more solid basis than those which have preceded
them.

There is nothing invariable about the pathway of
degeneration. It can no more be said to retrace the
pathway of progress in an inverse direction than it
could be said that in a country abandoned by its
inhabitants the more recently formed paths of com-
munication would be the first to become effaced. It
is quite true that the broader roads, which would
naturally last longer, are frequently the oldest
paths of communication ; whereas the footpaths,

220 THE PATH OF DEGENERATIVE EVOLUTION

which are the first to disappear, are usually of
more recent origin. Very frequently, however,
the new roads follow a rather different direction,
and, although more recently constructed, are not
the first to disappear.

It is the same with great commercial crises. It
is quite inexact to say with Ribot, who is respon-
sible for the analogy : " Old houses offer the best
resistance to the storm ; it is the new houses
which, being less solid, crumble and fall." *

After the time of the cotton famine, during the
American war of Independence, the greater part of
the old firms of Gand became bankrupt, whereas
most of the large, newly-established joint-stock
companies survived the crisis.

Moreover, in those cases where the most recently
formed structures are the first to decay, it cannot
be deduced that evolution is reversed, and that
the institution returns to its primitive condition,
for there is no reappearance of the intermediate
structures.

1 Ribot, Les Maladies dc la Memoire, p. 99.

PART II

The irreversibility of degenerative evolution

Most authorities on the subject are agreed that
evolution is not reversible,1 and that institutions
or organs which have disappeared or been reduced
to rudiments do not reappear and develop afresh.
It would be a useless extension of this volume
to cite many facts in favour of a view which is
almost without supporters, but it will be useful to
examine the exceptions, real or apparent, and to
discuss —

1. If an institution or organ which has dis-

appeared may reappear.

2. If an institution or organ which has been

reduced may resume its primitive function.

3. If an institution or organ which has been

reduced may redevelop and assume a func-
tion other than its original function.

1 L. Dollo, Les his cle revolution (Soc. Belg. Giol. Pal&mt.
Hydr., t. vii., 1893, proces-verbaux, pp. 164-166.

222 THE PATH OF DEGENERATIVE EVOLUTION

CHAPTER I

DO INSTITUTIONS OR ORGANS WHICH HAVE DIS-
APPEARED REAPPEAR ?

Section I
Disappeared organs

In biology we are almost unaware of indisputable
examples of the normal reappearance of disappeared
organs.

1. Plants. — As the embryonic development of
plants is usually direct, it is impossible to decide
whether an organ which forms a component part
of the embryological history represents an ancestral
organ. However, in a few rare cases, artificial
selection causes an actual reversion of evolution.

Typical geraniums possess two whorls of five
stamens, as, for instance, in Geranium. In Erodium
there is only one cycle of five. In Pelargonium one
cycle of five is complete ; the other is represented
by two stamens and three filaments which have
lost their anthers. But in certain varieties with
very large flowers the two complete cycles re-
appear, five stamens having long, and five short,
filaments. In this case there is no doubt as to
the reappearance of the three stamens lost in
typical Pelargoniums.

DO INSTITUTIONS OK ORGANS REAPPEAR ? 223

This reappearance is the result of artificial
selection. The typical Pelargoniums have a bi-
lateral symmetry, but horticulturists set a higher
value on flowers with radial symmetry, and in
consequence have produced flowers with such a
symmetry. As a matter of fact, they have paid
attention only to the symmetry of the petals, but
in modifying that, they have also modified the
symmetry of the stamens.1

The Privets (Ligustram), like most of the Oleacete,
possess only two stamens. Nevertheless, it is not
uncommon to find among normal flowers of the
common Privet specimens with three or four
stamens. In this case, however, it is uncertain
whether there is a real reappearance of lost organs,
or if the loss has not actually become complete.

Animals. — In the case of animals, teratology and
embryology furnish a few exceptional cases of an
apparent reversion of degenerative evolution.

As an abnormality in the horse, the first, second,
fourth, and fifth digits may reappear.

Adult man has lost the complete covering of
downy hair. According to Ecker, however, hyper-
trichosis is an abnormality really due to the re-
appearance of this ancestral condition, as may be
seen from the mode of its distribution in whorls.
Hypertrichosis is a trait frequently inherited, and
in this connection the Mauchamp variety of Merino

1 The flowers upon which these observations were made were
kindly provided by H. Canuell of Swanley.

224 THE PATH OF DEGENERATIVE EVOLUTION

sheep are interesting. This variety was obtained
by breeding from a sport which appeared in a
normal flock, and which transmitted its peculiarity
to its descendants. From time to time, in normal
flocks, variations occur which are similarly capable
of giving rise to Mauchamp breeds.

As recorded by Willett and Walsham, there
have been found in human children cases of a
bone stretching from the scapula to the sixth and
seventh cervical vertebrae. According to these
authors, the bone represents the suprascapula of
the tailless amphibia, which the normal homologue
in man is the merest edge of the scapula ossified
from a separate centre.

Such cases, as well as cases of polydactylism and
of supernumerary mamma?, are usually set down as
atavistic. However, the attempt to explain by
atavism such pathological and teratological peculi-
arities must be made with caution. Such inherited
anomalies occur very frequently in degenerate
families — the neuropathic families of F^re * — and
are associated with other abnormalities equally
heritable and certainly not due to atavism. Such
are pigmented retinitis, congenital cataract, chro-
matic asymmetry of the iris, asymmetry of the
pupil, ichthyosis, pigmented erectile spots on the
skin, and congenital disposition to bleeding.

As in a degenerate line of heredity these
abnormalities may replace one another indiffer-
1 F^re, La Famille n&vropathique, 1894. Paris, F. Alcan.

DO INSTITUTIONS OR ORGANS REAPPEAR ? 225

ently it is very doubtful if any of them ought
to be regarded as the inheritance of ancestral
traits. Teratology contains no undoubtful case
of the reversion of evolution.

On the other hand, it seems certain that in the
individual development of some species there is a
real reappearance of lost organs. The larval his-
tory of certain Crustacea Malacostraca 1 (cray-fish,
shrimps, etc.), seems to provide instances.

In Stomatopoda, the youngest erychtheus larvae
(fig. 71, a) are formed of three parts : the head,
five anterior thoracic segments, each bearing a pair
of biramous swimming limbs (i.-v.), the three last
decreasing in size from before backwards (these five
pairs of appendages represent the five pairs of buccal
appendages of the adult) ; three terminal posterior
segments (vi.-viii.), and a caudal fin, all without
appendages. Tn older larvae the first and second
pairs are profoundly modified, losing a joint and
acquiring gills (fig. 71, B, i.-ii.) ; the third, fourth,
and fifth pairs disappear completely, or at most
occur as minute saccules (hi., iv., v.). New thoracic
segments are formed, and later on in the third
larval stage the thoracic appendages reappear in
their final form.

Similar facts occur in the development of the
Decapoda Macroura, such as Palinucrus and
Scyllarus. While within the egg the creature
passes successively through nauplius and phyllo-

1 Lang, Anatomie comjwirdr, vol. i., p. 458.
P

226 THE PATH OF DEGENERATIVE EVOLUTION

some stages, and possesses all its thoracic appen-
dages (three pairs of jaws and five pairs of
ambulatory appendages). In this larval life the
exopodites of the second and third pairs of jaws

—J/

Fig. 71. — Erychtheus larvae of Stomatopoda.
A, The youngest known Erychtheus larva : I — V, buccal appendages well developed. —
B, Young Erychtheus larva : I and II modified buccal appendages; III— V, degenerating
buccal appendages; VI — VIII, segments devoid of appendages. — C, Older Erychtheus
larva: 1 — II, modified buccal appendages; III — V, reappearing buccal appendages;
VI — VIII, ambulatory appendages in course of formation. (After Claus, in Lang's Traiti
d'anatomie comparee, vol. i.)

atrophy. Shortly before hatching, the first pair
of jaws atrophies completely ; the two pairs of
antennae and the two posterior pairs of ambula-
tory legs become very degenerate. In the young
phyllosome stage these organs are rudimentary,
but in the older phyllosome larvse the first pair

DO INSTITUTIONS OR OEGANS REAPPEAR ? 227

of jaws and the two posterior pairs of ambulatory
limbs are reformed ; the posterior two pairs of jaws
reacquire exopodites, and gills are formed on the
ambulatory limbs.

As a matter of fact, in most Crustacea, the
ambulatory appendages appear when they become
necessary, that is to say at the end of larval life, but
in the Stomatopoda and in the Decapoda Macroura,
owing to inheritance, they appear much sooner.
But when these appendages are useless during the
larval life they disappear again to reappear at the
end of larval life as in most Crustacea. This
adaptation of the larva to special conditions is of
great importance, as the larval life is most im-
portant from the point of view of the species.

Section II.
Disappeared Institutions.

The apparent revival of hygone institutions. — It
seems, at first sight, as if there were many instances
of the subsequent revival of bygone institutions.

Those of ancient Borne and Greece, for instance,
appear from time to time to have been recon-
structed. In feudal Home of the fourteenth century,
Cola di Rienzi, by turns tribune and senator of the
people, re-established the old republican constitu-
tion. During the Renaissance period the ancient

228 THE PATH OF DEGENERATIVE EVOLUTION

schools of science were resuscitated, and during the
French Revolution similar attempts at revival were
made, especially in the department of politics. When
Herault de S^chelle, being ordered to draw up a
scheme of legislation, revived the laws of Minos,
in the constitution of the year VIII., the Tribunal,
Senate and Consuls reappeared. During the First
Empire, Napoleon, in imitation of Augustus, affected
a respect for republican institutions, and had the
coinage stamped with his own effigy and that of
the Republic. In Germany, the Holy Empire
which nominally ceased to exist in 1806, reap-
peared in 1871. In Greece, the Olympic games,
suppressed in 1525, were re-established in 1896.

It is hardly necessary to insist upon the essen-
tially superficial nature of these revivals. It is
always possible to bestow upon new social systems
the ceremonial code of an institution long since
abandoned, but it is quite impossible that the
institution itself should be resuscitated in the
midst of surroundings which have been completely
transformed. The consuls of the year VIII. and
the emperors of modern times do not resemble the
consuls and im%)eratorcs of ancient Rome more than
the Christian societies of the present day resemble
those of the middle ages. With regard to outer
form in the drawing up of statutes, in all which
constitutes, so to speak, the decorative side of the
institution, the organizers imitated minutely the
keures of the old ambachtcn • beneath this appar-

DO INSTITUTIONS OR ORGANS REAPPEAR ? 229

ent similitude, however, were structures of an
essentially different nature. Just as the new rail-
way station at Bruges, in spite of its towers and its
pointed arches, is far more like any other railway
station than a Gothic cathedral, so the Christian
societies of to-day, in spite of the archaic caprices
of their founders, resemble more closely modern
associations than ancient associations. We see then
that it cannot be said in any of these cases that
the actual revival of a decayed institution took
place. The empty form reappeared, but the founda-
tions and the essential parts had become completely
transformed.

2. The apparent disappearance of institutions, —
There are other instances, however, showing the
opposite of this phenomenon. The essential parts
remain unchanged, but the form itself is modified ;
the institution persists, but its existence is dis-
sembled. Sometimes even when the dissolution
of an institution has been enforced, there has been
a . reconstruction on the first opportunity.

Aii instance of this is the reappearance of poly-
gamy among the Mormons, the last traces of
polygamy having disappeared during the middle
ages from the people of the West.1

1 In Bigorre there was maintained up to the fifteenth century,
a kind of system of concubinage called massipia, which, though
not actually polygamy, was the contraction of an inferior union in
conjunction with real marriage.

In Marseilles, too, polygamy seems to have reappeared in the
middle ages, owing to the frequent intercommunication between

230 THE PATH OF DEGENERATIVE EVOLUTION

In ceasing to be legal, however, polygamy has
by no means ceased to exist, even in the present
day, and the Mormons in instituting polygamy
have only given official recognition to what had
never really disappeared.

In the process of social transformation, periods of
transition are frequently characterized by reactions
in favour of bygone institutions, which reappear
although apparently permanently abolished. This
has been the case with certain corporations, and
with numerous other institutions of the old regime
which reappeared after the revolutionary crisis.
Some years ago a large landed proprietor in the
domain of Peterhoff, in Eussia, attempted, in the
interests of his serfs, to introduce the rural system
of European countries. He divided up the land
into independent allotments, and built at his own
expense a separate house for each family ; but no
sooner was the abolition of serfdom declared than
the peasants proceeded to re-establish the primitive
community, and to rebuild their houses on the old
sites in spite of the considerable labour which this
entailed.1

3. Instances of convergence. — It sometimes happens

that town and the East. It was never, however, established there
officially as was the massipia in Bigorre. The municipality
promptly suppressed it by issuing the following mandate : ' ' Quod
(vir) non habeat duas uxores, vel mulier duos viros." (Viollet,
Histoire du droit civil, fr. p. 388.)

1 De Laveleye, De la ProprUte" et ses Formes primitives. Paris,
F. Alcan, 1882, p. 23.

DO INSTITUTIONS OR ORGANS REAPPEAR ? 231

that after the lapse of several centuries, an institu-
tion seems to reappear.

An exact analogy to the primitive contubernium
(the community of the cabin) is exhibited in modern
slavery. Only a few years ago, in the Spanish
Antilles, marriage between slaves was recognized by
neither church nor state. When a negro wished to
become united to a particular negress he asked per-
mission of his owner to share his cabin with her,
and these unions could only be dissolved with the
consent of the master. It is hardly likely that
these slave marriages of the Spanish Antilles are
survivals or rather resuscitations of the Roman
contubernhim. They rather represent a case of
convergence : identity in circumstance has been
productive of identity in institution. In this case,
as in all others of the same kind, it cannot be said
that a bygone institution has reappeared, for the
new institution has quite a different origin. Further,
in the other instances which have been mentioned,
an institution which has reappeared has never really
ceased to exist ; a real dissolution has never been
followed by a resuscitation. For this to happen, the
whole social surroundings would have to be trans-
formed into their former condition, which is obviously
impossible.

232 THE PATH OF DEGENERATIVE EVOLUTION

CHAPTER II

CAN RUDIMENTARY INSTITUTIONS OR ORGANS RESUME
THEIR PRIMITIVE FUNCTIONS ?

There is no break of continuity between a rudi-
mentary organ and the complete ancestral organ :
the rudimentary organ is the ancestral organ trans-
mitted by inheritance. It is the same organ not
only because it has the same form, but because it is
actually a part of it. For the reduced organ to
return to its ancestral condition and resume its
ancestral function, it must retrace the series of
steps in degeneration along which it has passed.

So also in sociology there are many instances of
institutions which are rudimentary, but which retain
their original form because of uninterrupted imitative
transmission. In their case also resumption of the
primitive functional activity would imply a retracing
of the degenerate steps.

This necessity of retracing shows at once that
after a certain amount of degeneration resumption
becomes impossible. We shall find in the few cases
we are able to adduce, that when organs retrace
their steps and resume their ancestral functions,
degeneration had not gone very far in them.

Section I.
Rudimentary organs.

I. Animals. — Among animals it is very unusual
for a rudimentary organ to become active again.

CAN ORGANS RESUME THEIR PRIMITIVE FUNCTIONS? 233

The only cases show that in them no great amount
of degeneration had taken place.

1. Muscles of the ear in man. — It is known that
the human ear possesses a number of intrinsic and
extrinsic muscles reduced to delicate fibres, and
incapable of producing movement of the whole ear
or of one part of the ear on another part. In some
abnormal persons, however, certain of these muscles
may be well developed, making movements of the
ear possible.

2. The abdomen and appendages in deep-sea
hermit-crabs. — The hermit-crabs of the deep sea are
another instance of reversion to an ancestral form.
Littoral hermit-crabs inhabit the spiral shells of
Gastropods, and to suit this mode of life the body is
unsymmetrical, the appendages of one side being
rudimentary. In the depths of the ocean such
spiral shells are rare, and the crabs either abandon
this mode of life or live in straighter shells. In
consequence the limbs and the abdomen become
nearly symmetrical again. It is plain, however, that
in the littoral crabs these structures are not truly
rudimentary.

II. Plants. — In plants it is very difficult to
distinguish between the reappearance of lost organs
and the formation of new organs.

1. Hermaphrodite flowers in Melandryum. — The
Hermaphrodite flowers of melandryum (fig. 59) may
be flowers which after being unisexual have again
become hermaphrodite, or they may have retained

234 THE PATH OF DEGENERATIVE EVOLUTION

the primitive type. We cannot decide between the
alternatives.

2. Branches of Colletia cruciata, Crataegus, Vicia
Faba, etc. — Some cases, however, point clearly to
a renewed development of rudimentary organs.
Here are some examples. (See also, further on,
page 244 on hybrid individuals of Pentstcmon.)

Colletia cruciata (fig. 72) in the normal adult
condition bears large flattened branches, which
serve for assimilation and possess only very rudi-
mentary leaves. Sometimes, however, the plant may
give rise to more slender branches with normal
assimilating leaves. These branches and leaves are
probably the reappearance of the ancestral condition.

Wild pear and apple trees produce small lateral
branches which are transformed into spines. These
thorns have evidently arisen from normal lateral
branches which originally bore leaves. In the
cultivated varieties these lateral branches have
resumed the leaf -bearing habit.

In the hawthorn {Crataegus) the lateral branches
are similarly modified into spines. None the less,
while these spines are still young they may be
artificially stimulated to produce leaves by cutting
the principal stem.

The branches of Vicia faba bear low down a set
of rudimentary leaves. If the main stem be lopped
while still quite young, the usually rudimentary
leaves grow to the normal size.1

1 Goebel, Beitra/je zur MorpJwlogie unci Physiologic des Blattes.
Bot. Zeit., 1880.

Fio. 72. — Colletia cruciata (after Goebcl, PflanzenbioJogiache Schihlerungen, vol. I., p. 17).
A, normal branch. B, branch recurring to ancestral form ; (natural size).

236

THE PATH OF DEGENERATIVE EVOLUTION

Many plants such as Sempervivvum1 (fig. 73)
normally possess a very much shortened stem.
However, if the plant be grown in a saturated
atmosphere the internodes of the stem lengthen out.

Fig. 73. — Sempervivum (ectorum.
A (left figure), normal branch; B (right figure), branch grown for several months in a
saturated atmosphere.

Certain Veronicas (fig. 74) bear only small scaly
leaves. Cultivation of these plants in an atmosphere
saturated with water, results in the appearance of
normal leaves.

Lastly, we may quote again the instances given
by Goebel of the production by Equisctum arvense
of normal leaves under special conditions.2

1 The branch figured here was grown by G. Clautriau, in the
Brussels Botanical Institute.

2 Goebel, Uebcr die Fruchtsprosse dcr Equiseten. (Ber. d. d. Bot.
Gen.), vol. iv. p. 184, 188G.

CAN ORGANS RESUME THEIR PRIMITIVE FUNCTIONS? 237

Section II.
Rudimentary institutions.

It seems, at first sight, as if some societies of
the present day furnished instances of a return to
primitive conditions.

Modern developments in finance seemed to tend
towards a return to the exchange system. Politi-
cal institutions,
after a period of
absolutism, point
anew towards de-
mocratic equality.
Corporations re-
appear in the form
of syndicates or
religious societies.
Landed property,
formerly collective
and now individual, seems to be again tending
towards collectivism.

The same phenomenon occurs in the evolution
of matters relating to maritime rights. The sea,
according to Eoman law, was equally open to all
maritime nations. Later on it has been from time
to time practically in the hands of a few nations,
and we have now returned to a condition in which
it is equally open to all.1

1 Tarde, Transformed in h <1h droit, jijt. 161-162, Paris, F. Alcan,
1890.

Fig. 74. — Veronica cupressoides (after Goebel,
Pflanzenliologische Schilderungen, vol. i. p. 19).
End of a branch grown under a bell-jar in a

saturated atmosphere. In the older parts (B),

the leaves are small and applied to the stem ;

in the younger parts (A), the leaves are larger

and protrude from the stem.

238 THE PATH OF DEGENEKATIVE EVOLUTION

These, however, are not cases of true revival.
The resemblance goes little farther than the name.

A comparison of modern institutions with such
survivals of primitive institutions as continue to
exist will demonstrate this point. The difference
between them is so wide that it would be hardly-
possible to utilize the old as a basis upon which to
form the new.1

1 Cf. Durcklieim, Les Regies de la me'thocle sociologique. Paris,
F. Alcan, 1895.

In sociology, dealing as it does with things familiar to us all,
such as the family, property, crime, etc., it is useless to attempt
to adhere to exact definitions. The exact meaning of some words
in common use in conversation cannot be defined with any pre-
cision ; the common acceptation of these words cannot be avoided.
Now this common acceptation is frequently very ambiguous, so
that two totally different things are often referred to under the
same name, causing hopeless confusion.

There are, for instance, two different kinds of monogamous
unions — those so only in point of fact, and those which are
legally so. In the first case, a man has only one wife, though
legally entitled to several ; in the second he is only legally
entitled to one. These two kinds of conjugal conditions are
quite different, and yet the same word serves to express both ;
it is commonly said of some animals that they are monogamous,
although there can be nothing approaching to a legal contract
between them. Spencer, when dealing with the subject of
marriage, makes use of the term monogamy without defining
it in its common and equivocal sense. The result of this is
that the evolution of marriage seems to him to represent an
incomprehensible anomaly. It seems, according to him, that
the superior or monogamous form of union was prevalent during
the primitive phases of historic development ; that it then dis-
appeared during an intermediate period, to subsequently reappear.
From this he concludes that there is no regular connection between

CAN ORGANS RESUME THEIR PRIMITIVE FUNCTIONS? 239

It may be definitely asserted then that a re-
duced, but still persistent, institution never again
becomes actively functional. The following are a
few examples which will serve to illustrate this
point : —

1. The truck system and clearing-house. — Some
forms of the primitive system of exchange sur-
vive, not only in countries where money is
unknown, but in certain industries where the
workers continue to be paid in kind (the truck
system).

On the other hand, there seems to be a modern
tendency towards the elimination of money as an
instrument of exchange. The clearing-house system
is singularly analogous to the old exchange system.
"The truck system," says Stanley Jevons, "represents
the first and the last stage ; but it appears for the
second time in a very different form. Gold and
silver money continue theoretically to be the
instrument for buying and selling, but practically
metal no longer constitutes the real medium of
exchange, and has ceased to pass from the hands
of the purchaser into those of the vendor."

In this transformation there is obviously no
return to primitive systems, the last vestiges of
which, far from being revived, are rapidly dis-
appearing.

social development in general and a progressive advance towards
an improved system of family life. A more exact definition would
have prevented this erroneous conclusion.

240 THE PATH OF DEGENERATIVE EVOLUTION

The suppression of the truck system coincides
with the development of the clearing-house system.

2. Corporations and syndicates. — The radical
differences existing between the corporations of
former days and the greater part of modern pro-
fessional associations has already been pointed out.
The ecclesiastical associations, however, of the
present day are modelled as closely as possible
upon mediaeval institutions. It does not follow
that the last remaining vestiges of the latter have
been revived. There seems to be evidence that
quite the contrary has taken place.

At Bruges no attempt was made by the founders
of the guild of ambachten to resuscitate such
mediaeval corporations as continue to exist in a
state of decline.

At Iseghem, a small town in the west of
Flanders, we have already seen that the corpora-
tion of shoemakers was divided up into six or
seven guilds at the time of the Eevolution. An
attempt was made a few years ago to reconstruct
and modernize these guilds, but the scheme fell
through. A new corporation — wholly disconnected
with the guilds of Saint Crispin, and with no
structural resemblance to them — was established
instead.1

3. Archaic collectivism and modem collectivism. —
Societies of the present day exhibit numerous

1 Emile Vandervelde, EnqtiUe sur les Associations professionelles
d'ouvriers el d 'artisans en Bclyiquc, i., p. 17, Bruxelles, 1891,

CAN ORGANS RESUME THEIR PRIMITIVE FUNCTIONS? 241

vestiges of archaic collectivism. The question
arises as to whether there is a tendency in the
modern school of collectivism to resuscitate such
vestiges as remain of the old archaic form of col-
lectivism. Far from this being the case, collective
property, as conceived by the modern socialist,
implies the suppression of the few existing remnants
of archaic collectivism.

Inheritance, ab intestat, for instance, is a survival
from the days of the family community, which
itself arose, as we have already seen, from the
primitive community. If the modern collectivist
school had any desire for a return to the old
primitive community, it would make for the re-
construction of the family community by re-estab-
lishing the law of collateral succession. Now it
is just the opposite with the collectivists. In
order to establish a universal system of collective
property they demand among other things, the
suppression of inherited succession, ab intestat, at
least as regards the collateral line of descent.

4. The survival of elective sovereignty in Eng-
land.— The above examples apply to institutions
which have degenerated without having completely
ceased to be functional.

It very rarely happens, however, that having
arrived at that condition, they renew their vitality
and all their former functions, and this still more
rarely occurs in cases of genuine survival.

In the English coronation ceremony vestiges

Q

242 THE PATH OF DEGENERATIVE EVOLUTION

remain of the old democratic system in which the
king was elected by the people.1

The English sovereignty of the present day is
merely a decorative institution, the real head of the
Government being the Prime Minister, who is
nominated in fact if not in theory by the public.
This system may almost be regarded as a return
to bygone democracy. Nobody would wish, how-
ever, to revive the old system of elective sovereignty,
and to retrace in an inverse direction the various
stages of its degeneration.

CHAPTER III

CAN RUDIMENTARY ORGANS OR INSTITUTIONS RE-
DEVELOP AND ASSUME NEW FUNCTIONS ?

The few facts which we are able to cite on this
subject must be received with considerable caution.

1 The formality of an election disappeared during the Tudor
period. The coronation of Henry VIII. was the last occasion on
which the formula was read which set forth the national agree-
ment with and recognition of, the succession. The king was, in
fact, declared chosen and elected. This formula of election, which
disappeared after the coronation of Henry VIII., is recalled to mind
by the conclusion of the coronation ceremony of the present day.
The archbishop, walking in succession to all four corners of the
platform upon which the throne is placed, addresses the people in
the following terms : ' ' Gentlemen, I herewith present to you the
undisputed sovereign of the realm. Come all who are present and
offer homage to him. Are you prepared to offer it ? " and the
people signifying their assent by acclamation, cry, ' ' God save the
Queen " or "God save the King." (De Franqueville, Le gouvcrne-
meiit et le parlcment Britanniqucs, i., p. 291.)

CAN ORGANS REDEVELOP NEW FUNCTIONS ? 243

Section I.
Rudimentary organs.

1. Animals.— In Birgus latro (a land-crab of the
Philippines), the gills are atrophied and the bronchial
chamber is very richly supplied with blood vessels,
while a kind of incipient lung is formed from the
lining membrane of the reduced bronchial chamber.1

However, it is by no means certain that the
atrophy of the bronchial apparatus has preceded
this development of a pulmonary apparatus. In
the following case it rather seems to be one in
whicli a rudimentary structure has redeveloped in
order to assume a new function. In the develop-
ment of the urinary organs, it appears that the
ducts of the mesonephros are quite independent of
those of the pronephros, although these mesonephric
ducts become functional later in the embryonic life
than the pronephric ducts. They are, neverthe-
less, formed at an earlier stage, and their rudiments
have appeared before there is any trace of the others.
From this fact it would appear that in some
ancestors of existing vertebrates there existed simul-
taneously mesonephric canaliculi and canals homo-
logous with them, but exercising a different function.
Such a condition actually exists in Amphioxus : in
the branchial region of that animal there are pro-
nephric urinary canals and genital chambers which

1 Semper, "The Natural Conditions of Existence as they afreet
Animal Life." (International Scientific Series.)

244 THE PATH OF DEGENERATIVE EVOLUTION

are homologous with mesonephric spaces ; but these
latter do not exist as genital chambers in higher
vertebrates.1

It must be noted that the homology between
mesonephric spaces and the genital spaces of
Amphioxus, as made by these writers, is not uni-
versally accepted.

2. Plants. — The Scophulariacese, which have
usually four stamens, are derived from ancestors
which possessed five. Usually the fifth stamen is
only represented by a tiny process which rapidly
atrophies. However, in Pcntstcmon the fifth or
posterior stamen is developed, not as a functional
stamen, but as a staminode, the function of which is
to stretch open the flower to make it accessible to
hymenopterous insects with short probosces.

Can it be said that in such cases a rudimentary
organ has really become redeveloped to assume a
new function ? To establish this it would be
necessary to show in the case we have just
mentioned, that the stamen did not become trans-
formed directly into a staminode, but that it first
became rudimentary and then developed afresh into
a staminode.

An interesting fact is, that in some hybrid
varieties of Pcntstcmon the staminode becomes fertile

1 Boveri, Die Nicrencanalchen der Amphioxus. {Zool. Jahrbuch.
Abth. Anat. unci Ontogcnie der Thiere, vol. v., 1892.)

Wiedersheim, Grundziigc der Vergleichcnden Anatomie der Wir-
belthicre. Jena, 1893.

CAN ORGANS REDEVELOP NEW FUNCTIONS ? 245

again. In some flowers sent to us by Mr Cannell
of Swanley, the number of petals was increased to
six, seven, eight, or nine. In some of these the
posterior stamen was sterile and like a staminode ;
in others there were five fertile stamens. It is
obvious that in this case the staminode had resumed
its original function after having lost it.

Section II.
Rudimentary institutions.

The Levirat. — In his work entitled Tableau des
origines et de Involution de la famille et de la
proprie'te', Kovalewsky mentions an instance of a
reduced institution which, without having first
ceased to be functional, became transformed into
another institution.

" The custom mentioned in the Bible of alloting
a woman to the brother of her deceased husband, is
explained by the primitive condition of things with
regard to the relations between the sexes ; all the
women were the common property of the men
belonging to one group of relations. Under the
name of levirat, this custom survived for several
centuries, owing to the idea which arose later on
that a wife was property. Consequently, on the
death of the husband, the widow, along with his
other belongings, was treated as the inheritance of
the person whom the death promoted to the rank
of chief or head of the family community."

246 THE PATH OF DEGENERATIVE EVOLUTION

The levirat, a family institution, thus derived
from the old system of marriage by groups, was
transformed by degrees into an economic institution.
It is important to notice, however, that this trans-
formation was effected without the institution having
even been reduced to the condition of a mere
survival.

PART III

SUMMARY AND CONCLUSIONS

From all the facts that we have brought together,
the general conclusion becomes plain that retrogres-
sion, notwithstanding the etymology of the word,
does not imply a return to the ancestral condition.

Eudimentary organs and institutions resemble the
primitive states of these, in so far as they no longer
possess certain parts which the primitive stages did
not yet possess. None the less, profound differences
exist between the primitive and the reduced forms.
In the primitive condition the institution or organ
is capable of varying in the direction of new uses ;
in the reduced form, after a certain degree of
atrophy, there is no longer the possibility of
redevelopment to resume old or to acquire new
functions. These observations apply equally to
biology and to sociology.

Magnan and Legrain, in their work on de-
generate persons, came to similar conclusions.
They came to regard degenerate persons as
abnormal, chiefly because they were devoid of
the power to reacquire the normal condition and
quite unlike their primitive ancestors, who, although

248 THE PATH OF DEGENERATIVE EVOLUTION

possibly brutal and unintelligent, were normal
beings with the activity and stamina necessary
for future progress.

The following two diagrams, borrowed from these
authors, represent clearly the differences between the
initial and reduced condition of an organ or institu-
tion : —

In the diagrams the ascending lines represent
the progressive evolution of an organ or institu-
tion ; the descending lines represent the degenera-
tive evolution. From the point a, representing
the primitive condition progressive evolution passes
towards o, an imaginary perfect condition of the
organ. Along the upward line, however, the points
a, b, c, d, etc., represent obstacles to further pro-
gress— that is to say, factors tending towards
degeneration. From these points lines of de-
generation pass towards z, and the condition at
z, although representing that at a, is not identical

SUMMARY AND CONCLUSIONS 249

with a, and is not reached by a sliding backwards
down the line o, a.

Thus, although the most recently acquired
features may disappear first, degeneration is not
an actual retracing of steps until the point of
departure is reached. The degenerate condition
is a new point, and really the term retrogressive
evolution is misleading.

BOOK III

CAUSES OF DEGENERATIVE
EVOLUTION

PART I

ATEOPHY OF ORGANS AND INSTITUTIONS

The factors of atrophy

The causes which are active in producing degenera-
tion are various, but they may all be referred to the
limited nature of the means of subsistence, that is
to say, of nourishment in the case of organisms,
and of capital and labour in the case of institu-
tions. This limitation produces a struggle between
the individuals (societies or organisms) and between
their component parts.

In the course of the perpetual struggle for
existence among the different parts of an indivi-
dual, the institutions or organs which have ceased
to be functional tend to disappear, their nourish-
ment being absorbed by the active parts.

1. Biology. — In biology the struggle for existence
among component parts appears clearly as a factor
of degeneration in the case of accidental atrophy.
This is to be seen, for instance, in the atrophy of

2SI

252 CAUSES OF DEGENERATIVE EVOLUTION

the leaves of an etiolated plant or of the muscles
of a limb which has been immobile for long, or in
the case of muscles which have become inactive
from disease of the central nervous system. The
results are similar in cases of normal atrophy.
In frogs, toads, and other Batrachia Anura,2 the
disappearance of the tail before the adult state is
reached is the result of a struggle amongst the
cells. The active protoplasm of the muscular
fibres develops specially, and gives rise to many
cells, which enter the contractile material and
separate its elements. Gradually all the contrac-
tile material is absorbed by these isolated cells.

Many plants, especially Sempervivum (see fig.
73, A, p. 236) possess a reduced stem with the
leaves closely massed upon it. This reduction
of the stem, which is nearly constant in the

1 When a cutting from a potato or a seed (fig. 75) is allowed to
sprout in the dark, the young stems assume characters different
from those of plants grown in light. The absence of chlorophyll
produces important modifications of growth. In light the stem
is short, and the leaves are large and expanded ; in darkness the
stem is very long, and the leaves are much reduced. This atrophy
of the leaves is the result of the struggle for existence amongst the
organs of the plant. Light increases the rate of transpiration,
which is chiefly due to the presence of chlorophyll. As chloro-
phyll is most abundant in the leaves, the transpiratory current
sets strongly towards them, carrying in it the nutritive materials
for the formation of new cells. On the other hand, in etiolated
plants, transpiration is slower, and the nutritive materials delayed
in the stem give that the opportunity for specially active growth,
which takes place at the expense of the leaves.

2 Metchnikoff, Annales Inst, Pasteur, January 1892.

THE FACTORS OF ATROPHY

253

species, is the result of a struggle amongst the
organs produced by a scanty water supply. The
leaves attract to themselves the greater part of
the water absorbed by the roots,
and thus retard the growth of 7

the stem. But if the plant be
cultivated in an atmosphere satur-
ated with water, the struggle be-
tween the leaves and stem is
stopped, and the stem grows to
a much greater extent (fig. 73,
b, p. 236).

Vicia Faba, like most vetches,
produces at germination rudi-
mentary leaves, and similar leaves
are borne at the base of each
branch. Goebel has shown that
these rudimentary leaves may be
made to grow by cutting away
the terminal buds at a young
stage. In this way the struggle
between the first formed and later
leaves is suppressed.

In the case of the individual, ^t^t&T^Z
atrophy results from the struggle
for existence amongst the organs.
In atrophy throughout a species it is the struggle
for existence amongst individuals that plays the
chief part. Clearly in this struggle, useless organs
become impediments and burdens. If any organs

Fig. 7-r>. — Two seedlings
of Cicer arietinum.

seedling grown in light.
B, seedling grown in
darkness.

254 CAUSES OF DEGENERATIVE EVOLUTION

are useless they are harmful as they use nutrition
without conferring any advantage upon the whole
organism. Darwin pointed out that when through
changed environment a structure became useless,
its degeneration became certain, as it was a dis-
advantage to the individual to squander nourish-
ment upon a useless part. Weismann has shown
that such a reduction or disappearance of a useless
organ is the result of variation and natural selec-
tion.1

Variation results in the appearance of individuals
with the useless organ in various stages of imperfect
development ; natural selection perpetuates these
advantageous stages by giving advantage to indi-
viduals which tend to produce the organ in the
most degenerate condition.2

1 See, however, Herbert Spencer's Social and Moral Problems, as
he differs from Weismann on this point.

2 The following are good examples of the operation of variation
and selection in producing atrophy in a species : —

(1) Loss of constant colour among domesticated animals. — Wild
animals, especially birds and mammals, have a colour which is
constant for a whole species. Frequently the colour is protective
in rendering the animal little distinguishable from the environ-
ment in which it lives. As soon as such a species has been
domesticated man becomes its protector, and protective colouration
is no longer necessary, and soon disappears. In the wild state,
the colour is quite as variable as in the domesticated state, but
the abnormal individuals become the prey of enemies and are
removed from the species. This applies not only to animals
which are preyed on by others, but to predatory animals them-
selves. The wild-cat, for instance, will have less difficulty in
stalking its prey if its colour makes concealment easy. In the

THE FACTORS OF ATROPHY 255

2. Sociology. — With societies this elimination of
useless structures is effected much more easily than
with organisms for several reasons : —

In biology a special factor, heredity, gives to
specific characters a force which does not exist in
the same degree with social institutions. Now
functional organs common to a whole line of
descent are not easily effected by the influence
of individual surroundings. Further, the trans-
domesticated condition man provides food and the colour being
unimportant all variations may survive.

(2) Loss of spines in plants on oceanic islands. — It is well known
that the presence of spines protects plants from the ravages of
herbivorous animals, particularly mammals. But in oceanic islands
bats are generally the only mammalian inhabitants, and so, accord-
ing to Wallace (Darivinism), there are no spiny plants in the
indigenous flora of St Helena. The much richer flora of the
Hawaian islands includes only a very few prickly plants. All
the endemic genera are unarmed, as also are most of the endemic
species of other genera ; even genera like Xanthoxylum, Acacia,
Xylosoma, Lycium, and Solanum, which are so frequently armed
in other countries, are there represented by unarmed species.
The two species of Hubus bear prickles reduced to the merest
points and the two palms are devoid of spines. How is the
absence of spines to be explained in these plants ? The plants
have been derived from the mainland, the seeds being brought by
the wind, by currents, or by birds, and having found soil have
germinated. In their new country they are not attacked by
herbivorous animals, and it is immaterial to them whether or no
they bear spines. The individuals badly armed are at no dis-
advantage compared with those possessing the normal armature ;
on the other hand, they have the advantage of being without
useless organs to support. Spines, in consequence, gradually
disappear.

This struggle for existence may cause the disappearance of some
organisms themselves, and not only the atrophy of parts of them.

256 CAUSES OF DEGENERATIVE EVOLUTION

mission of acquired characters is, to say the least,
doubtful. There is no proof of individual atrophy
being hereditary, while with societies modification
may be transmitted by imitation. Institutions
which have fallen into disuse rarely recur in
freshly-formed societies. Natural selection plays
an all-important part in biology, but it is artificial
selection which almost exclusively governs the social
domain.

Many vegetables, as for instance the carrot (Daiocus Carota) are
natives of France. The seeds of the cultivated carrot must fre-
quently be carried to waste lands or uncultivated soil. The
domesticated variety, however, is never found Avild although the
wild variety is abundant. This vegetable has lost the power of
struggling against weeds ; it nourishes only when it is protected
by man and when by repeated weedings its wild competitors are
removed. When it is returned to its original wild haunts the
plant dies out at once.

Most cereals, although we may not know their wild ancestors,
are in a similar condition. For instance, if man were to cease
cultivating Wheat (Triticum sativum), or Rye (Secale cereale),
there is no doubt but that these would completely disappear.
Their fate would be shared in Belgium at least by many species
which are reaped with them at harvest, such as Centaurea cyanus
the Corn-flower, Agrostcmma Githago the Rose-Campion, Specu-
alaria speculum, and others. If a corn-field were left to the free
operation of nature, weeds would soon intrude and cause the dis-
appearance of the plants usually present in it. What would
happen in Belgium would happen with other plants in other
countries. Thus, near Bergen in Norway, some plants, such as
Melandryum album, Silene inflata, Vicia cracca, etc., occur only
in cultivated fields. In Java, many aquatic plants such as Limno-
charis Plumieri, Ludwigia perennis, Jussiaea suffruticosa, etc.,
live only in the rice-fields which are artificially watered and
manured. The cessation of tillage would cause the disappearance
of all these plants from those localities.

THE FACTORS OF ATROPHY 257

An institution which has become useless and
burdensome is generally suppressed before its
complete degenerative evolution is accomplished.
This suppression may be either voluntary — as in
the liquidation of a commercial company, for
instance — or it may be enforced. By the terms
of Article 73 of the Belgian Company's Act,
" the dissolution must be declared upon the
demand of all those interested at the termina-
tion of six months from the time when the
number of shareholders has been reduced to less
than seven."

The downfall is generally effected in this sudden
way, either voluntarily by the interested parties, or
by the intervention of legislative means.1

Sometimes, however, artificial selection does occur,

1 The occurrence of autotomy or self-mutilation in animals, as
in crabs, has analogies with what we have been discussing.
Similarly some plants brought into a new locality suddenly
shed their leaves. Ranunculus aquatilis, cultivated in water,
produces long divided leaves without stomata and with chloro-
phyll in the epidermic cells. If, from some chance, the water
falls below the level of the plants, the adult leaves become dry
and perish. The very young leaves growing unsubmerged are
still divided, but to a lesser extent ; they have stomata, and the
epidermis is devoid of chlorophyll. If the plants be again sub-
merged, this form of leaf dies, and there is a new development
from the youngest leaves of the normal aquatic type.

Other plants show similar occurrences. Thus, when a Fuchsia
that has been cultivated in a conservatory is exposed to the air,
all its leaves are shed and are replaced by new leaves. These new
leaves again fall at once if the plant be brought back into the
conservatory. This is a real case of autotomy in plants.

R

258 CAUSES OF DEGENERATIVE EVOLUTION

and then the degenerative evolution of useless insti-
tutions is brought about in a similar way as that
of non-functional organs. Atrophy of this kind may
be, as in biology, either accidental or normal.1

In a besieged town cut off from all outside com-
munication, all train service is necessarily stopped
and the railway staff rendered useless. On the
other hand, the defence of the city requires both
men and money. Under these circumstances the
resources of the railway naturally come to be
absorbed in the service of the defence.

As a good example of normal atrophy may be
mentioned the disappearance, at a certain point

1 See Durckheim on the difference between normal and acci-
dental sociological phenomena in Les Regies de la me'thode
sociologique.

"All sociological phenomena, like biological phenomena, are
liable, while remaining the same individuals, to revert to
different forms. Now, of these forms, there are two kinds of
reversion : —

"The one is common to the whole species, and is to be found,
if not in each individual, at least in the greater part of them.
The cases are not always identical, varying slightly with the
individual, but individual variation is restricted to very narrow
limits.

" The other kind of reversion is exceptional, being of a nature
rarely met with, and, when occurring, is seldom permanent
throughout the life of the individual. Cases of this kind are excep-
tional in point of duration as in other respects.

"Here then are two distinct, varieties of phenomena which
should be distinguished from one another by different terms.
Individuals exhibiting only common characters are called ' nor-
mal, ' while those exhibiting exceptional characters are designated
as 'morbid' or 'pathological.'"

THE FACTORS OF ATROPHY 259

of social development, of the popular assemblies
in which lay the origin of future societies, i.e.
the comitia, assemblies at the market-place, the
witenagemot, May Day games, rustic assemblies,
the Landsgemeinde of the Swiss cantons, the
parochial assemblies of the Andora Valley, and the
town meetings of Xew England, etc. Some of
these were suppressed, but some of them merely
fell into disuse as other institutions arose which
were better adapted to more modern conditions of
society. This happened with regard to the comitia
curiata of the Eomans during the period of the
Empire, which were gradually supplanted by the
comitia ccntariata and tribunal comitia. At the
time when they are first mentioned in history, they
fulfilled only one function : that of ridding the
laws of all traces of extrinsic customs.1

To sum up then, it is plain that although social
degeneration is brought about by the same general
causes as organic degeneration, the comparative
importance of the factors in degenerative evolution
is far from being identical in the two cases. The
autotomy of organs, a protective self-mutilation,
exhibits only a far-fetched analogy with the
conscious and voluntary suppression of social
structures which have become either useless or
prejudicial.

Direct individual adaptation, which plays a part

i Mommsen, Droit public romain, Le peuple et Ic Senat, vol. i. ,
p. 364.

260 CAUSES OF DEGENERATIVE EVOLUTION

of only secondary importance in the development of
animals and plants, is a dominating element in
sociology. Institutions are able to modify their
structure by assimilating new inventions and im-
provements, and by getting rid of the useless parts.
On the other hand, indirect adaptation spread over
a species plays no part whatever in sociology, for
societies seldom reproduce the structures of the
societies from which they sprang, when the latter
have ceased to exist ; whereas, in animals, when a
useless organ is reproduced by hereditary repetition,
variability and natural selection become agents in
its suppression.

CHAPTEE I

ATROPHY OF ORGANS

Part II

Causes producing atrophy

The ultimate cause of the atrophy of organs is the
limitation in the quantity of nourishment. We
have shown that if there were an indefinite supply
of food there would be no struggle, and, as a result,
no degeneration. We have now to examine more
minutely the course of atrophy and the nature of
its immediate causes.

The atrophy of an organ is a reduction in

ATROPHY OF ORGANS FROM LACK OF SPACE 201

structure, in nutrition, and in functional activity,
but the succession of these three events varies with
the nature of the exciting cause.

Eeduction begins with structure when the exciting
cause is lack of space, due, for instance, to increase
in another organ {atrophy from lack of space).

Atrophy begins with function when an organ has
become useless (atrophy from lack of utility). This
uselessness may arise from two causes ; the function
may be no longer useful to the individual or to the
species, or it may be assumed by another organ.

Lastly, atrophy may begin with a diminution in
the supply of nutritive materials (atrophy from lack
of nutrition). This defective nutrition may be the
result of a general cause such as feebleness of the
whole organism, or it may be due to the hypertrophy
of another organ.

§ 1. Atrophy from lack of space.

Cases of this kind are rare.

1. Development of the teeth. — Among animals, the
development of the teeth furnishes an excellent
example. The number of the teeth in human
beings is reduced compared with the number
present in Lemurs and in Platyrhine Monkeys.
These have six grinding teeth while in man five
is the maximum number. Our posterior molar,
however, appears late in life ; it is smaller than
the others, so that it may be useless for chewing ;

262

CAUSES OF DEGENERATIVE EVOLUTION

frequently it is absent, in the lower races in

nineteen per cent, of
cases examined ; in the
higher races in forty-two
per cent.

The reduction in num-
ber and size of these
teeth is due to a re-
duction in size of the
jaw-bones, a cause which
also frequently produces
a distortion in the
arrangement of the other
teeth. The rudiments
of the wisdom teeth
appear on the maxillary
tuberosity and on the
coronoid process ; it is
only after eruption that
they come into normal
connection with the jaw-

Fig. 7G. — Tip of an ear of Lolitim perenne, •,
with two earlets, the superior bearing DOlieS.

two glumes, the lateral earlet with 0 , 7 „ , ,

only the superior glume. In the I. At V Op II y Of tllC

lateral earlet the lower flower is open . , .

and has two glumules ; all the other SUpCriOV glume. AmOUg

flowers aie closed and exhibit only the , ,, »

inferior giumuie. plants there are tew cases

GI, inferior glume: GS, superior glume; „ , , „

gi, inferior glumule; gs, superior Ot atrophy as a result Ot
glumule. i l t? t

lack or space. In grasses
of the genus Lolium, the earlets are arranged in a
spike, but in such a fashion that only the terminal
earlet has space for both glumes (fig. 76). The

ATROPHY OF ORGANS FROM LACK OF USE 263

superior glumes remain ; the inferior, pressed against
the axis, disappear after the embryonic develop-
ment of the flower.

3. Degeneration of pcUece and of stamens. — Lack
of space is probably the immediate cause of the
disappearance of palere in the receptacles of some
composite flowers and of the posterior stamen in
the flowers of some Scrophulariaceae and Labiates.

In normal racemose inflorescences each floret
grows in the axil of a reduced leaf called a
bract. When the axis of the inflorescence is
shortened and the florets crowded, as in the
capitula of composite flowers, it frequently hap-
pens that the bracts of the florets (termed
palese) disappear. This absence is most usual
when the capitulum is small and the florets are
large.

In Labiates and most Scrophulariacea?, although
the ancestral number of stamens was five, there are
not more than four present ; when only one is
absent, it is the original posterior stamen which
was pressed against the axis of the inflorescence.

§ 2. Atrophy from lack of use.

1. Functional Inutility.

(1) Etiolated plants and immobile limbs. — We
have already quoted as instances of accidental
atrophy, cases of degeneration of leaves in etio-

264 CAUSES OF DEGENERATIVE EVOLUTION

lated plants, and of muscles in unused limbs (see
fig. 75, p. 253).

Qf <S?£

Fig .77. — Nymphaea alba sewn on the mud and at different depths in it. 1, 2, 3, successive
stages of the same seedling. Eau, water; limon, mud.

ATROPHY OF ORGANS FROM LACK OF USE 265

(2) Epicotyl and 'primary leaf of Nym/phaea. —

These structures in the water-lily are good
examples of normal atrophy in individuals. Dur-
ing germination (tig. 77) the cotyledons of the
water-lily remain inside the seed, and a new
organ (at right angles to them) grows vertically
upwards. The lower part of this is the first
internode of the stem (epicotyl), and the upper
part is a primary acicular leaf. It grows up-
wards through the mud until the summit of the
leaf reaches light. The growth of the epicotyl is
then much slower, and its terminal bud begins to
shoot out horizontally. The use of this growth of
the epicotyl and primary leaf is to carry the bud
to the light. When that purpose is achieved, these
structures atrophy. A similar occurrence may be
found in Sagittaria sagittifolia (fig. 40, H, I, p. 72).
In that case, however, it is the hypocotyl which elon-
gates, until light is reached, and then degenerates.

(3) Boots of Utrieularia, cotyledons of parasitic
plants, leaves transformed to spines in Phyllocactus
crenatus. — As instances of atrophy throughout a
species produced by inutility of the parts con-
cerned, we have already mentioned the roots of
Utrieularia and the cotyledons of the parasitic
plants Cuscuta, Orohanche, etc.

The spines of Phyllocactus crenatus produced
from modified leaves are another example (fig. 78).
Above the rounded base by which a branch of
Phyllocactus is attached to older branches, there

266

PAUSES OF DEGENERATIVE EVOLUTION

\

is an angular region, the sides of which are
prominent, and bear leaves modified into spines
to serve as protecting organs, as in the similar
case of Cereus. Higher up the branch the pro-
minent sides become flat, and the
spines are replaced by minute
scales. This degeneration is the
result of loss of utility. The
Phyllocacti are epiphytes, and
their situation consequently places
them out of the reach of crop-
ping animals. The spines near
the basis of the branches are a
survival from the terrestrial an-
cestors of Phyllocactus. The
spines higher up have degenerated.
Animals offer many instances
of atrophy as a result of inutility,
both in individuals and in species.
(4) Atrophy of the branchial
arches in mammals. — As they are
no longer functional, most of
the mammalian branchial arches
atrophy. Three pairs alone persist,
and of these it is only those parts which are useful.
(5) Atrophy of ventral fins. — Instances of atrophy
through uselessness in species are to be found in
the ventral fins of fishes like the Pecliculati, which
live in the mud, or in Protopterus, which for a part
of the year is completely buried in mud (fig. 19, p. 44).

'

i/

Fig. 78.— Branch of
Phyllocartus crenalus.

ATROPHY OF ORGANS FROM LACK OF USE 267

(6) Atrophy of muscles. — Cessation of use is also
the cause of the degeneration of the flexor and
extensor muscles of the fore-limb in Cetacea, and
of the imperfection of the finger joint articulations
in Cetacea and Sirenia (fig. 79). In the latter
cases the surfaces of the articu-
lation which make flexor and
extensor movements easy dis-
appear. When a limb becomes
a paddle, it is necessary that it
should be flexible, but that the
articulations should be immobile.

(7) Atrophy of the tail in
man. — The caudal region of the
human vertical column is com-
posed of four or five very de-
generate vertebrae. The whole
of this organ is degenerate.
When the tail is formed at an
early stage of embryonic life, fw. n.—Giobiocephaius

,t , i i -i • , n melas. Right anterior

the vertebral column consists of nn showing absence of

,! . , . , , ,i , , ■• articular facets for the

thirty-eight vertebrae ; the lesser joints of the fingers.
number of vertebrae in the adult (
is due to reduction of the tail, which in man is
quite useless. Later on in life a further instance
of atrophy may occur in individuals. In old men
the caudal vertebrae are frequently fused, and the
whole region is smaller.

(8) Degeneration of the hyoid apparatus in mam
and birds. — This case shows a close correspon-

268 CAUSES OF DEGENERATIVE EVOLUTION

rlence between atrophy and loss of function. The
second arch becomes connected with the third, the
parts of which are, in the adult, the styloid pro-
cess, the stylo-hyoidean ligament, and the lesser
horn of the hyoid bone. According to the weight
of the tongue, the parts of the second arch become
more or less developed. In man the suspensory
apparatus of the hyoid bone is extremely simple,
and it is still more reduced in birds. " The
tongue, reduced to a minute cartilage, no longer
requires the support of a bony base so that the
hyoidean apparatus might almost be removed from
the anatomy of a bird. It is present, but in a
rudimentary condition " (Geoffroy-Saint-Hilaire).

In the horse, which has a heavy tongue, the
second arch is strong and completely bony.

In fish the hyoidean system is still more im-
portant, although in them this is associated not
with any importance of the tongue, but with the
branchial apparatus. The parts of the second arch
are very strong, as they form a fulcrum against
which the branchial system works, but its main
parts are recognizable. " The hyoidean apparatus
is the same in all vertebrates ; its functions are at
a maximum in fish, and at a minimum in birds,
while in mammals the condition is intermediate "
( Geoffroy-Saint-Hilaire).

II. Transference of function. — Among plants
and animals there are many instances of organs well
developed in younger stages of life, but which

ATROPHY OF ORGANS FROM TRANSFERENCE 269

become rudimentary in later stages on account of
their functions being assumed by other organs.

(1) Atrophy of the tail in Batrachia Anura, and
of the larval gills in some insects. — The tadpole of
the frog has a well-developed tail which acts as
the organ of locomotion ; the adult animal moves
by its limbs, and the tail, useless in the adult, has
been removed by phagocytosis.

The aquatic larva; of many terrestrial insects
possess tracheal gills, that is to say, membranous
expansions of the skin, within which tracheae
ramify. In the adult insect respiration is con-
ducted by normal tracheae communicating with the
air, and the larval organs of respiration atrophy.

In addition to such cases of atrophy occurring
normally in the life-history of individuals, there
are known many cases where the organs of a
species have disappeared on account of the trans-
ference of their functions.

(2) Disappearance of limbs. — Vertebrates which
move by general undulations of the body have
lost their limbs for this reason. Such cases are
Slow-worms, Amphisbiena, Snakes, Eels and Cre-
cilians.

Many parasitic creatures have similarly lost their
organs of progression, as they depend upon their
host for movement from place to place.

Sacculina, a parasite on the carapace of crabs,
has completely lost its organs of locomotion. More-
over, as it takes its food by processes passing into

270 CAUSES OF DEGENERATIVE EVOLUTION

the tissues of the crab, its digestive canal has been
lost.

(3) Atrophy of the leaf. — In many plants the
leaves have disappeared, their function having
been assumed by some other part of the plant,
as, for instance, by the phyllodes in Acacia and
in Phyllanthas (fig. 84).

(4) Atrophy of the, protonema in mosses, and of
the leaves in some xerophilous plants. — At germina-
tion, mosses produce a much branched filamentous
structure which serves as the organ of nutrition,
and is termed the protonema. Later on this gives
rise to buds which develop into the normal leafy
shoots of the plants. As soon as the leaves are
large enough to manufacture food for the plant, the
protonema begins to degenerate, and disappears
completely, except in a few rare forms (Ephe-
meracese) where the leafy shoots remain very
small.

In Muehlenbeckia platyclados (fig. 80), which has
become adapted to arid regions, and in consequence
has the surface from which evaporation may take
place much reduced, the branches which do not
bear assimilating leaves are rounded at their bases,
but higher up flatten into broad blades. These
blades contain chlorophyll and fulfil the functions
of leaves, these latter being present only as minute
scales.

Similar phenomena occur in many Papilion-
aceous plants belonging to the genera Genista

ATROPHY OF ORGANS FROM TRANSFERENCE 271

(broom), Spartium, Alhagi, etc. The leaves dis-
appear, and the stems assume their functions.

(5) The reduction or disappearance of the calyx.
— The disappearance of the calyx in many com-
posite flowers and of roots in
epiphytes are instances of species
losing organs because of the
transference of function to other
organs.

In the simplest Composite,
the fruits (achencs) are dissemi-
nated by the wind, the calyx
usually being modified into a
feathery tuft. In other Com-
posite, although wind dispersal
still occurs, the feathery calyx
is lost, its purpose being fulfilled
by wings on the sides of the fruit
(Florestinia pedata), or it is united
to a large membranous scale
developed from the receptacle
(Dahlia), or the sterile florets
may be turned into wings (Lind-
heimeria texana). In other cases
the fruits are dispersed, not by the
animals which eat them (Clibadium
they adhere to the fur of animals by
on the achenes (Calendula), or by
from the involucre of bracts (Lappa).
cases, and the list might have been

. — Branch of Mue/i-
nbeckia platyclados.

wind, but by

asperum), or

hooks formed

hooks formed

In all these

made longer,

272 CAUSES OF DEGENERATIVE EVOLUTION

the function of the calyx has been reduced, or has
disappeared on account of the transference of its
function.

(6) Atrophy of roots. — Roots in most plants per-
form two functions : they fix the plant in the soil,
and, chiefly by means of the delicate hairs on their
youngest parts, they absorb water and dissolved
mineral substances. Sometimes, however, instead
of having root-hairs, the rootlets enter into a
kind of partnership with a fungus, which lives
in their tissues, and absorbs by its processes the
necessary food materials from the soil. The pine
and beech are examples of this.

In some orchids (Corallorhyza, Myrmechis) the
fungi lodge in the subterranean part of the plant,
and the branches, having no function, disappear.

In most of the epiphytic Bromeliaceae the roots
are useful only to anchor the plants ; the absorption
of water takes place entirely through new organs
developed from the leaves. The roots are few in
number and small, and after the plant has obtained
a firm position their growth almost ceases. In
another species (Tillandsia usneoides) of the same
family the roots have completely^ disappeared.1
This plant, called by the Brazilians, " the plant of
the air," fixes itself to branches of trees by its
leaves, and is easily transported by the wind. Its

1 For further details concerning the Bromeliacese see Schimper,
Die epiphytische Vegetation Amerikas, in Schimper 's Botanische
Mittheilungen aus den Tropen. Jena, 1891.

ATKOPHY OF ORGANS FROM TRANSFERENCE 273

absorption takes place entirely through hairs de-
veloped upon the leaves.

In addition to such examples of plants which
have lost their roots on account of the functions of
the roots being assumed by other organs, there are
also plants in which the roots represent the sole

Fio. 81. — Tce/iiophyllum Zollingeri with branches adhering to bark. The plant
bears a flower and a bud.

vegetative organs. In Twiiiophyllium Zollingeri
(fig. 81), an epiphytic orchid, the leaves are reduced
to minute scales and are of no importance in the
nutrition of the plant, that function being trans-
ferred to the roots which in the form of flat green
ribbons, apply themselves closely to the bark of
trees. In this case the roots have assumed the

s

274 CAUSES OF DEGENERATIVE EVOLUTION

functions of leaves, while in Tillandsia itsneoM.es
the leaves have taken on the functions of roots.

The Podostemacese are aquatic plants which live
in warm regions attached to rocks in cascades.
The stems and leaves are completely absent, the
flowers even being produced on the roots. Some
of the roots become closely attached to the stones.
Others which are green and ribbon-like, float in the
stream, and serve for assimilation.

§ 3. Atrophy from lack of nutrition.

In some cases of degeneration, the organs do not
lose their functions, but become reduced, merely
because adjacent parts rob them of their nourish-
ment. This kind of lack of nutrition which results
in local atrophy, must be distinguished from the
general limitation of food-supply which is the
ultimate cause of all degeneration. The possible
amount of food within the reach of any organism
is limited, but besides this, the share of the absorbed
food obtained by any particular organ or part of
an organ may be limited with a resulting atrophy
of that organ or part. Atrophy of this kind may
be accidental or normal.

1. Parasitic castration. — A good instance of acci-
dental atrophy of this kind is found in Melandryum
album where the ravages of a fungus Ustilago
amthcridarum may produce parasitic castration.1

1 See Giard, La Castration parasitaire in the Bull. Scient. de la
Frame et de la Belgique. 1887, 1888, 1889.

ATROPHY OF ORGANS FROM LACK OF NUTRITION 275

Under the influence of this parasitic fungus which
makes its way to the anthers, the stamens of the
female flowers assume the form of those in male
flowers and in consequence the pistils of these
flowers abort from defect in nutrition.

2. Severe or prolonged compression of a limb. — A
long continued or forcible compression of a limb
results in atrophy of its extremity on account of
lack of nutrition from the compression of the blood-
vessels.

3. Atrophy of the genitalia in neuter bees. — Lack
of nutrition is also the cause of the arrested de-
velopment of the genitalia normal in neuter bees.
The neuters of bees and of some of their allies are
females in a state of arrested development. In
wasps, humble-bees, and hive-bees, it sometimes
occurs that the genitalia of these forms develop
sufficiently to be functional, thus resulting in the
appearance of small females. In most honeycombs
two kinds of cells are formed : in the smaller and
more numerous cells are placed the larvae destined
to become neuters, in the larger and less numerous
those destined to become queens or perfect females.
The food of the two sets of larva? is different ;
those in the larger cells are given " royal food " a
more nutritious substance. When some of the
royal food by an accident gets into a worker cell
the sexual organs of that larva are developed so
that a small female is formed. In this way as
many females as may be desired can be produced,

276 CAUSES OF DEGENERATIVE EVOLUTION

and when a hive has lost its queen, the worker
bees produce another1 (Lacordaire).

Plants afford many instances of degeneration due
to defect of nutrition.

4. Atrophy of the superior flowers in Carcx. — In
the tall spikes of Carex, it frequently happens that
the flowers towards the summit are rudimentary,
and authorities are agreed in regarding this condi-
tion as the result of defect in nutrition.

5. Atrophy of pistils and stamens. — Cases of
atrophy of the stamens or pistils normal in species
may be given.

In Fritillaria persica the flowers "are disposed in
bunches. The lower flowers possess six perianth
members, two cycles of three stamens each, and a
pistil. In the median flowers the pistil is smaller,
and rarely capable of being fertilized. In the
superior flowers the degeneration is complete, the
pistil hardly being formed. It might be shown

1 The transference of a function is not invariably accompanied
by degeneration. Thus, in the functional development of an
individual's nervous system voluntary acts which have been re-
peated frequently become reflex actions, and have their seat in a
different region of the nervous system — as, for instance, walking
and acquired professional dexterities. It has been sought to ex-
plain the development of the instincts of species in this way by
supposing that frequently repeated voluntary acts have become
inherited reflexes. As pathological degeneration in man frequently
affects the higher regions of the brain, reflexes and instincts may
persist after loss of voluntary action. In the cases of transference
of nervous functions to lower centres the higher centres do not
degenerate but remain able to acquire new voluntary functions.

ATKOPHY OF ORGANS FROM LACK OF NUTRITION 277

that this degeneration is due to lack of nutrition,
by removing the inferior flowers from a young
bunch, but we are unaware that the experiment has
been made.

In Viburnum tomentosum (fig. 82) .the flowers
are arranged in a compound corymb and occur in
two forms. The central flowers have a small
corolla, five stamens, and a well-developed pistil

Fig. 82. — Flowers from the same inflorescence of Viburnum tomentosum.
A, sterile flower with rudimentary pistil and no stamens. — B, sterile flower with
very rudimentary pistil and stamens. — C, flower with two fertile stamens; the
other three stamens and the pistil are rudimentary. — D, fertile hermaphrodite
flower.

(fig. 82, d). The eight or nine peripheral flowers
of each inflorescence have the corolla much larger,
but the stamens are absent and the pistil is rudi-
mentary (fig. 82, a). The petals turned towards
the outer side are much the largest, and it is not
rare for the petals turned towards the inner side to
be hardly larger than those of the central flowers.
What is important to notice, as it bears out the
relation between degeneration and lack of nourish-

278 CAUSES OF DEGENERATIVE EVOLUTION

ment, is that a smaller petal is always associated
with a larger stamen (fig. 82, b, c).

In Viburnum opulus, the wild guelder-rose, there
are also two kinds of flowers, the central flowers
which are sexual and hermaphrodite, and the
peripheral flowers which are sterile and possess a
very large corolla. The five petals are, however,
equal in size, and there are no forms transitional
between the sterile and sexual flowers. The
cultivated guelder-rose is a variety of the wild
form in which all the flowers are sterile and
possess a large corolla : the plant has completely
lost the power of sexual reproduction.

§ 4. Atrophy without apparent cause.

In many cases it is impossible to determine the
cause of atrophy. Why, for instance, has half the
inflorescence disappeared from a unilateral cyme ?
Why have some composites like Artemisia or
Rosacea? like Poterium lost their habit of • being
fertilized by the agency of insects, and become
anemophilous without conspicuous perianth ? Nor
can we explain why many Myriapods are blind, al-
though they live in association with species possess-
ing eyes and in conditions where vision apparently
would be useful. Semper discovered, in brackish
water in the Philippines, Crustacea (Cymothoe)
which were completely blind, although they lived
in light. The cause of such atrophies is unknown.

ATROPHY OF ORGANS WITHOUT APPARENT CAUSE 279

Moreover, we know that organs may atrophy
through correlation with other degenerating organs,
or because the whole organism is degenerating
although there are no obvious anatomical bonds
present between the related parts. As instances of
such atrophies we may mention degeneration of the
brain resulting on degeneration of the supra-renal
capsules ; or of the condition of myxodcema
resulting from mechanical or functional disturbance
of the thyroid gland. When the essential sexual
organs of the male or female are congenitally or
accidentally absent, the secondary sexual characters
— the beard, voice, hair, and whole male or female
aspect — of the body degenerate. When one eye
becomes injured or diseased, a frequent consequence
is sympathetic degeneration of the undamaged eye.

To these cases of correlative atrophy we may
add the cases of leaves on many plants with
drooping or horizontal branches. On these, the
leaves on the upper aspect are frequently rudi-
mentary. The appearance is most striking where
the leaves are opposite the pairs being at right
angles to one another. In a branch of Acer
campestrc which is upright, all the leaves are of the
same size. In a drooping branch the leaves turned
towards the sky are small. In plants belonging to
very different families (Acanthacea?, Melastomacece
Urticacere, etc.) oblique branches exist, and the
failure in development of the dorsal upper leaves is
invariable. In Procris laevigata (fig. 83) the reduc-

280

CAUSES OF DEGENERATIVE EVOLUTION

tion of these leaves has gone very far, but in
Klvgia notoniana the leaves in the corresponding

Fig. 83. — Branch of Procris laevigata.
F'/'j F"/"> F"V". The successive pairs of leaves in the axil of a large leaf F'
is a female inflorescence ; in the axil of a small leaf /" is a male inflorescence.
(The original drawing was made at Buitenzorg in Java by Mas Kromohardjo, a
Malay draughtsman.)

position are absent. In another representative of
the same family {Streptocarpus monophylleia), the
degeneration has gone still further ; all the leaves

ATROPHY OF INSTITUTIONS 281

are absent and a single greatly enlarged cotyledon
is the sole assimilating organ.1

It is evident that in these oblique and horizontal
branches, the leaves directed vertically towards the
sky are in the most unfavourable position for assi-
milation, and that, in addition, their presence would
shade the leaves lying under them. In these
species the more or less complete reduction is an
inherited fixation of a sacrifice of these particular
leaves for the benefit of the whole organism.

CHAPTEE II

ATROPHY OF INSTITUTIONS

The causes of atrophy in institutions are more or
less analogous to those which bring about atrophy
of organs. First, there is atrophy from want of
use, when function either becomes useless or is
transferred to another institution. Atrophy from
lack of resources corresponds precisely with
atrophy from lack of nutrition. There is no-
thing, however, in the atrophy of institutions quite
analogous to that which, in organisms, results from
lack of space. But if the development of an
institution cannot be actually impeded by the co-

1 See the figures of plants of this family in Fritseh, Gesacriacccc.
{Engler unci Pranll's naliirlichcn Pflanzenfamilien. Leipzig, 1891. )

282 CAUSES OF DEGENERATIVE EVOLUTION

existence of another institution, certain instances
of atrophy do occur which cannot be said to arise
from either want of use or lack of resources. The
institution degenerates merely from hindrance
offered to the exercise of its functions. Thus, for
instance, the laws of exclusion voted in Germany
against socialists brought about the decadence, and,
ultimately, the downfall of a number of professional
and trade associations which cannot be said to have
been actually suppressed by law. It is the same
with the corporations of Western Flanders ; these
have survived the revolutionary laws, but are no
longer legally recognized, a state of things leading
to many difficulties ; the properties of several of
them have been confiscated, as having no owners,
and allotted to benevolent institutions. This want
of legal recognition must inevitably lead in the
near future to the entire disappearance of these
vestiges of the corporative system.

Cases such as these, however, must be regarded
as exceptional, and it may be said that, on the
broad average, institutions atrophy either from
want of use, or from lack of resources.

§ 1. Atrophy from wa?it of use.
I. Functional inutility.

(1) Offices in connection with the Port of Bruges.
— The disorganization of the train service in a
besieged city has already been alluded to as

ATROPHY OF INSTITUTIONS FROM LACK OF USE 283

furnishing an example of degeneration by sus-
pension of function.

Another incidence of accidental degeneration is
furnished by the office of wharf-porter, which was
formerly exercised in the port of Bruges. When
Bruges, consequent upon the blocking up of the
Zwyn, ceased to be a sea-port town, the wharf-
porters who were formerly employed to carry
grain, lime, and coal, etc., were no longer required
and so abandoned their calling.

(2) The for est -courts of England. — As an instance
of normal degeneration due to the transformation
of an agricultural country into a commercial country,
take the old English forest-courts. In the middle
ages there still existed in England great tracts of
forest land which were Crown property, and sub-
jected to special legislation conducted by three
separate courts of justice : (a) The Court of Attach-
ment x which instituted proceedings ; (b) the Court
of Swainmote 2 before which the culprits were tried,
and the Judge's Court presided over by the Lord
Chief Justice, who pronounced sentence, and from
whose decision there was no appeal. These courts
have lost all importance since the seventeenth
century, and the forest laws are now only functional

1 When a forest law was infringed, it was the duty of the forester
to "attach" the culprit — i.e. constrain him to appear either by
seizing his person or his goods. These attachments were then
submitted to the Court of Attachment.

8 The judges were called verderers ; the jury was composed of
foresters of the reeve, and of four men out of each forest hundred.

284 CAUSES OF DEGENERATIVE EVOLUTION

with regard to the Forest of Dean and the New
Forest. The office of Forest Judge has disappeared,
and but few vestiges remain of the courts of attach-
ment and Swainmote. Those persons who are con-
nected with either the Forest of Dean or the New
Forest, meet together still to transact business in
connection with them, but these meetings are of no
great importance.

II. Transference of function.

Instances abound of the transference of a function
from one institution to another ; but as a rule, when
this change is effected, it is attended by the sup-
pression of the old institution. Sometimes, how-
ever, this is not the case, and the institution merely
atrophies without having been suppressed.

(1) Republican institutions under the Roman
Empire, — As instances of this form of degenera-
tion take the institutions of the Eoman Republic,
after the establishment of the Empire — or the
decadence of the functions formerly discharged
by the Privy Council in England, the political
functions of which have been transferred to Parlia-
ment, and the judicial functions to the Court of
Common Pleas and the Court of Exchequer.1

(2) Special jurisdiction in England. — It is the

1 De Franqueville, Le Gouvemement et le Parlement briitunique,
i., p. 431.

ATROPHY OF INSTITUTIONS FROM LACK OF USE 285

same with special jurisdiction, such as the Eccle-
siastical Court, and the University Courts ; they
have completely degenerated, their functions having
been transferred to the jurisdiction of the Common
Law Courts. In England, however, special juris-
diction has not wholly ceased to be functional, for
there still exist, side by side with the modern
Courts of Justice, a few local Courts and other
exceptional forms of jurisdiction which are still
maintained in support of certain ancient acquired
rights and traditions.1 By far the greater part of
these special jurisdictions have, however, fallen
into disuse, owing to the creation of the modern
Courts of Justice. This happened, for instance,
with the following institutions : —

(a) The Local Courts of Feudal origin. — These
Courts have decreased both in number and impor-
tance since the close of the thirteenth century ; the
only vestiges now remaining of them are the Court-
leets of certain manors. Sir James Stephen makes
special mention of the Court-] eet of the Manor of
Savoy, which extended from near the old city gate
of London (Temple Bar) up to Cecil Street. Some
of the old functions of this Court are maintained
in the present day.

(b) Country Town Courts. — Twenty-seven local
courts are mentioned in modern judicial statistics
before six of which no case has been heard for
twelve years. On the application of a litigant,

1 Idem, Le sys/eme jv.diciaire de la Grande- Bretagne, i., p. 216.

286 CAUSES OF DEGENERATIVE EVOLUTION

however, any one of these would resume its
functions.1

(c) The Court of Stannaries, the vice-warden
of which is appointed by the Prince of Wales.
This Court has greatly degenerated in importance,
and only a few minor cases are now heard before it.

(d) The Court of Piepowder. — This was a Court
of summary jurisdiction, dealing promptly with
disputes arising during fairs and markets. It has
almost disappeared from having fallen into disuse.
Practically only one example remains ; that is at
Bristol, and is becoming merged in another local
court called the Tolzey Court.

(e) The ITusting's Court (or folkmote, scirmote,
a kind of County Court of the city of London). —
The old Hunting's Court has completely lost all its
former attributes, in favour of the Court of the
Lord Mayor and Sheriffs. Up to 1860, however,
it reserved to itself the right of jurisdiction with
regard to matters relating to landed property in
the city ; the only cases which come before this
Court nowadays are those of replevy.2

Finally, there is the House of Lords which

1 According to jurisprudence, the fact that a Court still existing
by virtue of Royal Charter, has ceased to be functional for two
hundred years, is no reason against a citizen having his case tried
before it, if he so chooses, even though the town authorities declare
the funds to be insufficient for payment of the judge.

(Case of Rex v. Mayor of Wells, Doivling Practise Cases, j). 562),
mentioned by Franqueville.

2 Franqueville, Systemejud., i., pp. 235, and following.

ATROPHY OF INSTITUTIONS FROM LACK OF USE 287

formerly held full powers of jurisdiction with
regard to matters relating to members of the
peerage ; this jurisdiction is now limited to cases
of high treason or crimes committed by peers to
the exclusion of mere acts of misdemeanour which
are tried before the ordinary Courts.

§ 2. Atrophy from lack of resources.

The instances we have mentioned are those of
institutions decaying because they had become
useless, and their resources were transferred to
other objects. The lack of resources was the result
of the lack of function. Sometimes, however, it is
the cause and not the result, in which case atrophy
may be due either to an abnormal development,
such as the hypertrophy of another institution, or to
poverty, ensuing on the general decline of society
in general. Of course both factors may act
simultaneously.

1. Local administration at the close of the Roman
Empire. — The decline of local government at the
close of the Eoman Empire is an instance of
atrophy ensuing from the ultra-development of
another institution.

As the demands of the central powers grew
more and more excessive, the fiscal rates had to be
augmented in order to meet them, and the curiales,
which consisted of members of the City Council
who were made responsible for the payment of

288 CAUSES OF DEGENERATIVE EVOLUTION

taxes, finally found it was quite impossible to meet
their engagements, and made every effort to leave
the curia.1

2. The degeneration of Societies in all their parts.
— A number of instances might be mentioned of
general social degeneration bringing about the
atrophy of some one or other institution in
particular.

Besides giving classical examples, such as the
Romans, Peruvians and Astecs, V. Lilienfeld men-
tions the decline of the Negro kingdoms which
existed during the sixteenth and seventeenth
centuries in Southern and Western Africa, and
which are merely represented nowadays by
wretched little tribes.2

There are, according to Waitz, at some distance
from Carimango (the equatorial Eepublic) some
people of pure Spanish blood who have fallen back
into absolute barbarism. Their language is de-
formed past recognition, and their manners and
customs exhibit no traces of their former condi-
tion.3

Space precludes us from dwelling further upon
the various causes — often complex and obscure —
which bring about the downfall of societies,
suffice it to say that they are connected with

1 Lavisse and Rambeaud, Histoire g6nera.lt, I., ch. i. , pp. 14 and
following.

2 Von Lilienfeld, Gedarike fiber de Sozialwissenschaft der
Zukunft, ii., p. 241.

3 Waitz, Anthropologic der Ncdurvolker, 1. B., p. 369.

ATROPHY OF INSTITUTIONS FROM LA.CK OF USE 289

territory or with population, the two factors of
social evolution.

Either the physical surroundings of institutions
undergo unfavourable transformations, or else the
population itself degenerates.

(1) The almost complete disappearance of the
great family communities (zadrugas) of Montenegro
is a characteristic instance of atrophy from lack of
resources caused by the impoverishment of physical
surroundings. The persistent cutting down of trees
in the Black Mountain has had a disastrous effect
on the water supply, and consequently upon the
fertility of the ground. Most of the zadrugas,
having found it impossible to continue their
existence in common upon the same territory,
have split • up into small families (inokos?ias).
These latter represent, in a reduced state, the
old family system from which they have sprung.
Bogisic has shown that these in no way resemble
our modern families, but are to be regarded, from
the judicial point of view, as reduced family com-
munities, each comprised of only a few persons.

(2) Other cases occur where the atrophy of an
institution — of an artistic or scientific society, for
instance — is brought about by the degeneration of
a population which ceases to be interested in the
society and no longer contributes to its support.
A large number of cases of this kind might easily
be mentioned, especially as occurring during the
period of the Byzantine Empire, but it is difficult

T

290 CAUSES OF DEGENERATIVE EVOLUTION

to account for the sudden degeneration of a
people where the physical surroundings had re-
cently undergone no special modifications, and
when there had been no sudden and violent
check upon social development. According to
Lapouge and other sociologists of the Darwinian
school, this social degeneration was merely the
outcome of hereditary influences. The destiny of
a nation is dependent upon the quality of the
elements of which it is composed and by which
it is directed. If a nation is rich in energetic
and intelligent qualities, the greatest of disasters
can only have a transitory and limited influence.
When the contrary is the case, the same circum-
stances may produce an arrest in development
or a complete decline and fall. .Up to the
present time, and especially in antiquity and the
middle ages, these favourable qualities were gene-
rally supplied by a dominating minority establish-
ing itself in a conquered country. In the common
course of evolution, these superior elements, which
are indispensable to social progress, are eventually
eliminated. The inferior elements regain greater
power, and each step of their progress is attended
by a backward step towards barbarism. Although,
at first sight, this seems contrary to the Darwinian
theory, it is strictly in accordance with it. The
superior individuals are relatively inferior when
their chances of success or of posterity diminish.
The superior individuals may not only be swamped

ATKOPHY OF INSTITUTIONS FROM LACK OF USE 291

by a diminution in their birth-rate, but in some
cases there may be a direct elimination of them.1

The tendency of decadence is always towards the
degenerative and eliminatory selection of superior
elements.

It may be said in conclusion that there are
constant calls upon the capital and labour of a
society from its various institutions, and the conse-
quence is that, the resources not being unlimited, a
regular struggle for existence goes on amongst the
institutions. In the course of this struggle, the
decline of an institution may be brought about
in two dilferent ways. It either begins to de-
generate from lack of sufficient means of support,
or degeneration sets in consequent upon the insti-
tution having ceased to be functional by inutility,
by transference of function to another institution,
or by obstacles placed in the way of exercising
that function. In either case the institution dis-
appears. It is only in exceptional cases, which
will be alluded to further on, that existence is
still maintained.

1 G. de Lapouge, La Vie et la Mort cles nations (Re"vue int. de
Sociologie, 1894, pp. 421 and following). Several terms used
in this treatise were borrowed from the above article.

See also Hovelacque and Herve, I'ricis d' Anthropologic, p. 189 :
" War, in its double consequence of the elimination of the strong
and the survival of the weak, is for the more civilized races a
powerful factor in the cause of degeneration and downfall."

PART II

THE CAUSES OF THE PERSISTENCE OF OEGANS
OR INSTITUTIONS WITHOUT FUNCTION

CHAPTER I

SURVIVAL OF ORGANS

We have shown how and why organs may become
rudimentary and tend to disappear. In many
cases the disappearance is complete ; and the organ
may not even reappear temporarily in the course of
the individual development. This disappearance
is, however, by no means universal. Even apart
from the phenomena of recapitulation, rudimentary
organs may persist in the adult, and sometimes, even
although organs have ceased to be functional, they
persist without degenerating. We have now to con-
sider why in such cases degenerative evolution does
not result in complete obliteration of such organs.

§ 1. Unfunctional organs that arc not rudimentary.

Absence of Variations. — There are some plants
such as Ficaria l ranunculo'ides and Lysimachia

i Lysimachia Nummulai-ia occasionally produce seeds in some
valleys of the Pyrenees, and Errera has shown us specimens grown
from'seeds coming from the shores of the lake of Quatre-Cantons,
292

SURVIVAL OF ORGANS 293

Nummularia, the flowers of which hardly ever
produce seeds. How is it that in such species
flowers are still produced ? The probable ex-
planation of this anomaly is, that for the disappear-
ance of flowers there would have to be produced
individuals with this advantageous variation. It
is the case, however, that the Ficaria and the
Lysimachia reproduce most actively by asexual
methods, and variations are extremely rare in
cases of these modes of reproduction. The result
is that these species having begun to form sterile
flowers continue to produce them through simple
lack of variation.

An analogous case is presented by Elodea
Canadensis. This unisexual plant is represented
in Europe by only female plants. These plants
have multiplied asexually so luxuriantly that in
Holland they began to choke up the canals, and
it became necessary to make provision in the
budget of that country for the extermination of
the pest. The plants are, of course, able to multiply
only asexually, as the female flowers cannot be
fertilized, and these useless flowers have been
maintained simply from the absence of variations.

Stratiotcs alo'ides, a plant belonging to the
same family as Elodea, is practically only repre-
sented by male individuals. Females are extremely
rare, and none the less the male flowers are pro-
duced, although in the vast majority of cases they
must be useless.

294 CAUSES OF DEGENEEATIVE EVOLUTION

W. Burck has called attention to, without
endeavouring to explain, other instances of the
persistence of functionless organs. A large
number of Anonaceoe bear flowers which do not
open, and which are self- fertilizing (cleistogamous
flowers). None the less, they have retained the
corolla, the original purpose of which was to
attract insects.1

Burck has called our attention to the circum-
stance that several species belonging to the same
genus produce cleistogamous flowers, and that it is
improbable that this condition has been acquired
independently by these species. One would thus
have to admit that the original type must be very
remote, as it has given rise to descendants of
specific distinction, and yet the useless corolla has
persisted through the long series.

Parallel examples may be found among animals.

Machacritcs is an insect which inhabits caves
in North America. The females are quite blind ;
the males, on the other hand, have preserved, or
seem to have preserved, well-developed eyes ; but
are these eyes real ? An abyssal fish, Ijmojjs,2
seems as if it had enormous eyes extending from
the corner of the snout some distance along the
neck, but these organs are not really eyes ; they

1 W. Burck, Ueber Klcistogamie im weitcren Sinne und das
Knight- Darivinschc Gcsetz {Ann. Jard. Bot. Buitcnzorg, viii., p.
122, 1889).

2 Dollo, La Vie au sein des liters, p. 242, Paris, Bailliere, 1891.

SURVIVAL OF ORGANS 295

are light-producing organs, and the fish are in
reality blind. This may be the case also in the
males of Machaerites. It may also be the case
that the male has an opportunity for using eyes
absent in the case of the female, the males some-
times leaving the caves, the females remaining
within them. Something analogous to this occurs
in the case of eels : the males remain always in
the sea while the females rejoin them only for
purposes of reproduction. Moreover, there is still
a third hypothesis, that the male of Machaerites be-
came an inhabitant of caves later than the female,
and has not yet had time for the loss of its eyes.

§ 2. Unfunctional organs ivhich persist as rudiments.

It is outside our purpose to discuss here the
numerous cases of organs reduced through adapta-
tion, such as the leaves reduced to serve as
protectors of young buds (PhyUocactus, fig. 78), or
the wings of the ostrich which, although much
reduced, are supposed to assist the bird in running.
The utility of such organs explains their persist-
ence ; we are concerned here with organs which,
although useless, persist in a reduced form.

1. Absence of Variations. — In discussing organs
which, although without function, have persisted in
a complete state, we attributed the persistence to
absence of variations. It is probable that the
same cause operates in maintaining useless vestiges.

296 CAUSES OF DEGENERATIVE EVOLUTION

It is to be noticed, however, that the variability
of vestiges is frequently considerable. The flowers
of Asparagus officinalis are sometimes, although
rarely, hermaphrodite. Usually they are uni-
sexual and exhibit the organs of the other sex in
every conceivable stage of degeneration. It is
probable that the unisexual condition has been
acquired recently, and that there has not yet been
time for the operation of natural selection to cause
the disappearance of the useless organs.1

1 We have actual knowledge of the mode of disappearance of an
organ in one case, and can see the part played by variability. In
Phyllanthus spcciosus (Xylophylla arbuscula) the adult plant has
three kinds of branches : vertical branches, with rudimentary
leaves ; oblique branches which spring from the axils of these
leaves and themselves bear in two rows very rudimentary leaves ;
and flat branches which are the chief organs of assimilation of the
plant and which also bear rudimentary leaves in two rows. In the
seedling, on the other hand, there are formed after the two coty-
ledons, one or two completely developed assimilating leaves upon
an upright stem (fig. 84, a).

The flat branches grow from the axils of these, and bear,
unlike the flat branches of the adult, assimilating leaves ; higher
up the vertical stem bears only rudimentary leaves with flat
branches in their axils.

It may happen, however, that the seedlings bear, directly after
their cotyledons, rudimentary leaves (fig. 84, b) and in this case
the reduction of the leaves is present not only in the leaves borne
upon the vertical stem but in those borne on the flat branches.
These latter bear only a small number of assimilating leaves.
Thus, we have in this plant a remarkable example of incomplete
recapitulation : the seedling preserves in a functional condition
organs that are rudimentary in the adult, but the species furnishes
instances where these leaves cease to be functional even in the
seedling.

SUKVIVAL OF ORGANS

297

2. Insignificance of the Rudimentary Organ. —
It frequently happens that rudimentary organs are
preserved simply on account of their insignificance :
the absence of organs so small would not be an
advantage to the plant sufficiently great to be laid
hold of by natural selection.

Many species of Tropaeolwm bear leaves without

Fig. 84. — Two seedlings of Phyllanthns speciosus.
1, 2, 3, 4, successive assimilating branches ; c, cotyledons.

stipules. In Tropaeolum majus there are stipules
only in the case of the first two leaves of the
seedling, and the position of these stipules is very
variable. Sometimes both stipules are at the base
of the petiole ; sometimes only one is present ;
sometimes both are several millimetres from the
base — a position in which functional stipules never
occur.

298 CAUSES OF DEGENERATIVE EVOLUTION

In the same way may be explained the persist-
ence of accessory rudiments of enamel organs in
the development of teeth. Besides the rudiments
of the enamel organs for the milk teeth and
permanent teeth, there are additional organs
present in a very variable condition and number,
nearer the external surface. They are, however,
very generally present, and are exceedingly similar
to the youngest stages of the normal enamel organs.
Kollmann and Gegenbaur believe that they are
abortive rudiments surviving from an ancestral
condition in which teeth were more numerous.

CHAPTER II

THE SURVIVAL OF INSTITUTIONS

We have seen that an institution which ceases to
be functional without dissolution — either voluntary
or enforced — ensuing, tends to atrophy and dis-
appear, while its resources are appropriated by
other institutions. It now remains to account for
the fact that this atrophy does not generally end in
total disappearance. The two following hypotheses
may be made : —

1. The structure of a non-functional institution

may remain intact.

2. The institution may survive, but in a rudi-

mentary condition.

SURVIVAL OF INSTITUTIONS 299

§ 1 . Tlie integral persistence of an institution.

A non-functional institution may survive and
retain its structure and resources in the following
cases : —

1. By the intervention of some superior authority

to prevent its suppression.

2. When, while ceasing to be functional, it con-

tinues to he useful, though indirectly so.

3. When its existence is maintained out of respect

for old traditions.

We will take these three hypotheses in succession,
but it must be borne in mind that when a non-
functional institution is maintained out of respect
to tradition, or by virtue of an indirect usefulness,
it is always by the intervention of legal authority.
This legal authority, however, amounts to an ex-
pression of the public will, whereas an institution
may also be maintained by the exercise of personal
influence on the part of some one person.

It sometimes happens, too, that a non-functional
institution continues to survive because its suppres-
sion would entail important changes in other parts
of the social organization.

1. Maintenance by compulsion. — A useless in-
stitution is frequently maintained by compulsion,
when its conservation is advantageous to those con-
nected with it, or even to other persons.1

The following are a few examples of this : —

1 Spencer, Principles of Sociology, vol. iii.

300 CAUSES OF DEGENERATIVE EVOLUTION

(a) A long list might be made of all the sinecures,
now quite useless, that some governments insist
upon maintaining for the advantage of those occu-
pying the posts. Such were certain offices in con-
nection with the Court in former days or the
avoucrics of the end of the middle ages.

" Like the Fief system," says Errera, when writing
about the Massuirs, " The avoverie afforded an effec-
tive protection — military as well as judicial —
against the various dangers arising in a still barbaric
age. But, in the course of the last centuries of the
middle ages, the obligations of feudal chiefs and
the condition of avoverie disappeared ; the reasons
were that relative security was attained ; militia
was established, and the army, under the command
of the sovereign himself, became better disciplined ;
and that there arose the organization of the justices
scabinalcs, of bailiwicks, and of superior courts of
justice. However, although the ancient offices dis-
appeared, the emoluments attached to them con-
tinued to be drawn.1

(b) It often happens that institutions which have
ceased to be functional are yet maintained as being
a source of profit not only to those in direct con-
nection with them, but to a considerable number of
other persons.

Before the Eeform Bill of 1832, when large
towns like Leeds, Birmingham, and Manchester
were unrepresented in Parliament, the House of

1 Errera, Les Massuirs, p. 75. Brussels, Weissenbruch, 1892.

SURVIVAL OF INSTITUTIONS 301

Commons contained seventy members, nominated
by thirty-five rotten boroughs in which there
were no electors, and ninety members nominated
by forty-six boroughs, containing less than fifty
electors.

The borough of Old Sarum was a mere hillock
belonging to Lord Canalford ; Gatton and S. Michael
had only seven electors ; the borough of Dumwich
had been long since submerged by the encroach-
ment of the sea ; Beeralston, belonging to Lord
Beverley, consisted of one house, and Castlerising of
two. In the county of Bute there were twenty-
one electors, only one of whom was a resident and
who nominated himself.

The preservation of this system of representation
which had long ceased to be adequate, was eminently
favourable to the few persons who benefited by it,
and they vigorously resisted the passing of the
Eeform Bill.

(c) After the provincial states of Normandy and
the Dauphiny had been suppressed, the state func-
tionaries retained their titles and emoluments.

In the Dauphiny the representative institutions
ceased to be functional in 1627, but at the close of
the eighteenth century the Bishop of Grenoble con-
tinued to receive 0000 livres per annum as primate
of the dominion. Two barons, delegates of the
nobility, shared a similar salary, and the Syndic
of the province and two secretaries received lesser
emoluments from the province which continued

302 CAUSES OF DEGENERATIVE EVOLUTION

to make these payments for services no longer
rendered.1

In each of the above cases of survival it is plain
that compulsion intervened — more or less directly
— to secure the maintenance of non-functional
institutions.

In the first cases referred to, the privileged
persons concerned, took advantage of their influ-
ential positions to enforce the maintenance of their
sinecures.

In the second case — that of the rotten boroughs
— the institution was not only advantageous to the
member himself but to the whole of his party, so
that naturally its maintenance met with the cordial
support of the latter.

In the third case, those in authority maintained
part of the institution — that of the mere titles
and emoluments — in order to suppress the remainder
more easily. In this case it was not an institution
which atrophied, but an institution which was caused
to atrophy by compulsion.

2. Indirect usefulness. — It sometimes happens
that an institution, although ceasing to be functional,
yet retains a certain usefulness. This is so in
England, with the office of the Privy Seal. All
the functions formerly discharged by the holder
of this office have long since disappeared, but the
post is reserved as a sinecure for persons who have

1 Babeau, Les assemblies des pays d'Etat sous I'ancien regime ;
Reforme sociale, 1893, p. 704,

SURVIVAL OF INSTITUTIONS 303

distinguished themselves in politics, but who from
advanced age are unable to take an active part in
public affairs. A great many sinecures are main-
tained for a similar purpose — that of furnishing
practical though not nominal pensions to distin-
guished persons in art or science.

" It may well be," says Viollet, " that an
institution, which at first sight seems to be a
mere useless wreck, is really of immense service
to society. The deep roots of a dead tree may
continue to furnish support to a new structure."

It is by reason of this negative usefulness that
— according to Bagehot — the English monarchy
has been preserved, forming as it does a symbol
of unity and coherence amidst the electoral changes
of power. The English monarchy offers a char-
acteristic instance of an institution which has lost
nearly all its former functional importance, while
nominally retaining its power. According to
Bagehot, the Queen would now have to sign her
own death warrant if condemned by both Houses
of Parliament. The outer form, however, remains
almost the same as in the days of absolute monarchy,
when the sovereign took an active part in public
affairs. At within the last few years, the
Queen's signature was required to all official
documents. It was only in 18G2 that a law
was passed deciding that for the future, promotions
in the Army or Navy should be signed either by
the Commander-in-Chief, or by the Secretary of

304 CAUSES OF DEGENERATIVE EVOLUTION

State. At that time, by dint of hard work,
the Queen had signed all commissions up to
1858, and there remained 15,931 documents of
this kind still unsigned.1

3. Respect for Tradition. — The persistence of
some institutions can only be accounted for by a
lack of invention. Mommsen calls attention to
a remarkable instance of this kind in the history
of early Rome :

When a government by praetors — as consuls
were first called — was substituted for a govern-
ment by kings, the new system remained the same
in idea though nominally different. The old idea
of royal authority survived for a long period, and
the praetors enjoyed all the old kingly powers, even
those in plain contradiction to the temporary
character of their office : the king could not be
deposed, but neither could the praetor be con-
strained to depose himself ; the king, when dying,
nominated his successor himself, and this power
remained to the praetor — although the system of
election by the comitia had been introduced —
for the praetor had the right of excluding whoever
he chose from the list of candidates, and also of
annulling the votes given to those who displeased
him. It was only at a later period that a logical

1 Bagehot, The English Constitution, pp. 57 and following ;
London, 1891.

De Franqneville, Gouvernement et Parlement Britanniqucs, i.,
p. 251.

SURVIVAL OF INSTITUTIONS 305

and consistent conception of the consular authority
came to be formed.1

In the same way the maintenance of the insti-
tution of sheriffs in modern England can only be
attributed to an exaggerated respect for tradition.

In the commission which sat in 1888 to inquire
into judicial organization, one of the commissioners
expressed himself as follows : " I cannot see that
sheriffs are of any use whatever, unless it be for
show ; there is not one single function attached to
the post which a sheriff can fulfil himself. I am
a sheriff, but I do not know what a sheriff's duties
are." The institution, however, remains intact.
The sheriff is credited with the discharge of several
functions, which are executed in his name and on
his responsibility. All he does is to receive the
judges, accompany them on circuit, and preside over
parliamentary elections.2

The mode of nominating the sheriffs has remained
unchanged since 1340. The Queen, by means of a
traditional gold pin, is supposed to prick by chance
in the list of candidates which is presented to her,
the name of him upon whom she confers " the

1 Ferrero, Simbolii, p. 53, Torino, 1893.

2 " The two sheriffs of London, who are elected from among the
liverymen of the various city companies, have not to go on circuit,
but are supposed to attend at the Central Criminal Court. Their
functions chiefly consist in escorting the Lord Mayor to all city
ceremonies, and in attending numerous banquets, some of which
are given at their expense." De Franqueville, Systime judiciaire de
la Grand- Br dagne, i., p. 611.

U

306 CAUSES OF DEGENERATIVE EVOLUTION

charge and keeping of the county." This ceremony
is known as the " Pricking of Sheriffs."

§ 2. The Survival of Institutions in a reduced

state.

It has been shown that a number of institutions
in a state of decline continue to be maintained,
because they are still useful in spite of their reduced
condition.

This is the case with the symbolic ceremonies
which in former days accompanied the drawing up
of solemn contracts.1

We now come to institutions which persist in a
reduced condition, but which are not directly use-
ful to any one. In this case their persistence
may be attributed to one of two causes: either to
respect for tradition or to the insignificance of the
vestiges which remain.

1 According to Viollet in Histoirc du droit civil fr., p. 607, the
primitive assembly of the people still survives, though in a reduced
condition, in the Roman maneipatio, and in Scandinavia in a solemn
form of sale called the scotatio.

" I believe," he says, "that it was the primitive sale of German
law a sale concluded and ratified in a popular assembly, that gave
rise, in the middle ages, to the Scandinavian scotatio. So also in
the case of the Roman maneipatio there has long been believed to
exist the remnants of a popular assembly. The dumb witnesses in
the scotatio appear to me the petrified representatives of the
German tribe or village ; and, in the opinion of good judges, the
witnesses in maneipatio are no other than symbolic statues of the
five classes of the Roman people. However, this is mere
hypothesis.

SURVIVAL OF INSTITUTIONS 307

It is difficult, however, to distinguish between
these two causes, for insignificant vestiges of
institutions are especially numerous in very con-
servative environments, and, on the other hand,
mere respect for tradition very rarely ensures the
maintenance of either a harmful or an expensive
institution.

1. Insignificance of the Institution. — Some
of these insignificant institutions are reduced to mere
vestiges, no longer functional, or of use to any one,
others — representing traces of a former system —
retain some local vitality, although not in keeping
with the new conditions ; these are of such small
importance that their very insignificance ensures
their survival.

(a) The two French laws passed in 1835 and
1849 relating to entail have never become func-
tional at La Martinique. The old law of entail is
still active there, and this state of things is allowed
to continue without authoritative interference.1

(b) In England, the sovereign, up to the close of
the sixteenth century, reserved to himself the right
of presiding over the Courts of Justice, and of
pronouncing sentence. Since the Ee volution, no
sovereign has essayed to render justice personally,
and any attempt of the kind would be regarded as
unconstitutional nowadays. Vestiges of the old
system remain, however, in the formula of certain
legal proceedings, such as the serving of warrants,

1 Viollet, Hist, du dr. civ.fr., p. 883.

308 CAUSES OF DEGENERATIVE EVOLUTION

in which an order is given to appear before the
Queen herself, and it has never been suggested that
the old formula should be altered.1

The following are a few examples of local sur-
vivals which owe their existence to their insignifi-
cance : —

(a) Although Cambray has belonged to France
since the close of the seventeenth century,
its diocese includes a small part of Belgium;
the ancient religious organization has in
this instance survived political changes.2
(h) It is not generally known that with certain
properties situated in Artois — consisting
chiefly of marsh land — the law of primo-
geniture still holds good.3
(c) Public attention has recently been directed
to a very curious survival of the political
connections which existed during the middle
ages between Beam and Spain. Every
year, on the 13 th of July, the inhabitants
of the French valley of Baretous in the
Pyrenees, solemnly pay a tribute — for the

1 De Franqueville, Systime judiciaire de la Grande- Bretagne,
p. 23.

2Viollet, Hist. du. dr. civ.fr., p. 882.

3 Errera, Lcs Masuirs, p. 290. Decision of the Council of 25th
February 1779, as regards the castle -wards of Lille, Douai, and
Orchies. " Those portions of land which fall or have fallen to each
inhabitant as the result of division, shall be inalienable ; no person
shall possess two portions. The eldest male of each family, or in
default of the eldest male, the eldest female, shall alone inherit
the said portions of land."

SURVIVAL OF INSTITUTIONS 309

maintenance of peace — to the inhabitants

of the Spanish valley of Ronoal.1

It is plain that this custom, which has gone on

so long as to pass unnoticed, is too inconsistent

with the present relations between France and

1 The ceremony commences at nine o'clock in accordance with
instructions laid down in a document which, according to the
mention made of it in the Proces-verbal, dates from 1375.
First, the French mayors don their scarves of office ; next the
Spanish mayors, advancing from a group of compatriots, proceed
towards the frontier boundary line, accompanied by a peasant
carrying a lance with a red flag tied to it — a symbol of justice —
and draw up within six yards of the boundary. The French
mayors follow suit, but the flag hoisted on the lance which
precedes them, carried by a peasant, is white, as signifying
their pacific intentions. The Mayor of Isaba then says to
them, "Is it peace?" The French mayors reply in the affirma-
tive, and, as a proof of their sincerity, couch their lance upon the
mile-stone marking the boundary. The Spaniards then first plant
their lance in French soil leaning against the stone, and after-
wards place it so as to form a cross with the French lance.
Next the Mayor of Arette places one hand upon the crossed
lances, and the Mayor of Isaba does the same, and together
they utter the formal declaration of peace, which all those
present swear to observe. After the vow, the Mayor of Isaba
cries three times, " Paz davans ! " which means, " May peace con-
tinue."

Peace being thus declared, the Roncalais, in order to ratify
their abandonment of hostilities, order the guards to remove
their arms from the French side. The ceremony being over,
it only remains to pay the blood-tax. This formerly consisted
of three white mares, all exactly alike ; but, owing to the great
difficulty of matching them, three unblemished heifers were sub-
stituted, all of same colour and with the same markings. These
three heifers cost about 600 francs, which is a large sum for that
part of the world.

310 CAUSES OF DEGENERATIVE EVOLUTION

Spain to be continued much longer. " It is to
be expected," says the paper from which this
information was obtained, " that the French
Foreign Minister will in the near future come
to an understanding with the Spanish authorities
to put an end to this iniquitous custom, and it is
to be hoped that this year is the last occasion upon
which a blood-tax will be paid by the valley of
Baretous to the valley of Roncal."

2. Respect for tradition. — In his " Essays on Pro-
gress, Manners and Customs " ( Westminster Review,
1854), Spencer points out the connection between
respect for tradition and custom, and the con-
servatism of those in authority. He says that
certain customs, which have elsewhere died out,
survive in some departments of the government.
The Secretary of State in ratifying acts passed in
Parliament uses old Norman French,1 and certain
legal terms in old Norman French are still used.
The wigs now worn by judges and barristers are
identical with those seen in old portraits, while the
" Beef-eaters " of the Tower of London wear the

1 ' ' For financial Acts the formula is : La Reyne remercie ses bons
sujets, accepte leur benevolence et ainsi le veult ; for general Acts :
La Reyne le veult ; for private bills : Soit fait comme il est desire ;
for petitions : Sdit droit fait comme il est desire. The veto is
announced : La Reyne s'avisera. Cromwell had changed these old
forms ; he gave his consent to Bills in English ; the old custom
was resumed at the Restoration, and the House of Commons in
1706 rejected a Bill passed by the Lords to abolish the French
phraseology." De Franqueville, Gouvcrnement et Parlemeut Britan-
niques, vol. i., p. 279.

SURVIVAL OF INSTITUTIONS 311

same costume as that once worn by the body-guard
of Henry VII.1

Two similar examples may be added : —

(a) At the coronation of English sovereigns two
gentlemen of the Privy Council, chosen " on account
of their appearance," and created knights for the
occasion, are appointed by the Lord Chamberlain to
represent the Dukes of Aquitaine and Normandy.

(b) The First Officer of the Crown was formerly
the Lord High Steward, which title having, in the
course of time, become purely honorary, was
hereditary in the family of the Earls of Leicester.
The post is now in abeyance, but, on the coronation
of a sovereign, or, on the occasion of a peer being
placed upon his trial, this dignity is conferred upon
some important person nominated solely for the
occasion. It is not only in ceremonial — which,
according to Viollet, is the museum of history — that
reduced institutions, which are completely useless,
are tenaciously maintained ; it is the same with
judicial and religious institutions. A few examples
will suffice to show that this is so : —

(a) In its primitive form, the Assembly of the
People included the whole army, and was
necessarily held in some large open space.
The custom survived the necessity of the
choice of some such spot, and up to the
sixteenth century, whenever a new Emperor
whs proclaimed in Germany, it was the
1 H. Spencer, Morals, Science and Art.

312 CAUSES OF DEGENERATIVE EVOLUTION

custom for the electors to proceed to some
mountain for the purpose, probably because
it was the custom in former days to hold a
general meeting there, before the election.

In Iceland, the althing — consisting of
two chambers now — was formerly one large
assembly held in the open air upon the
Logberg (the Mountain of the Law) near to
the Lake of Thingvellir. In the Eepublic
of Andora, criminal sentences are still
pronounced with great solemnity from the
Market-Place.1
(b) Among the Ossetes, where the family com-
munity still flourishes, the only inalienable
and unsaleable property is not the real
property, but certain personal possessions,
such as the great cooking-pot with the
chain by which it is hung over the fire.
" At first," says Kowalewsky, " this may
appear a strange custom, but it must be
remembered that possessions of that kind
were of equal importance to the Ossetic
' family ' as were their tombs to the
ancient Greeks and Romans, which ex-
plains how any infringement of the

1 In Montenegro, it was the custom, till recently, for the Prince
himself to render justice, sitting under a tree in front of his palace
at Cettigne. This mode of jurisdiction, which was prohably a
survival of the old system of which we have been speaking, has not
completely disappeared.

SURVIVAL OF INSTITUTIONS 313

custom is regarded as an infamy by the
Ossetes." x

We now come to cases of purely religious
survival, which offer the strongest resistance to
the inroads of change. Spencer instances the
custom of circumcising with a knife made of
flint, and the vestiges remaining in Catholic
worship of former primitive religions. The
Eucharist, as we have already pointed out, is
reminiscent of real sacrifices, and the symbolic
representation by a clove of the Holy Ghost is
only a rudimentary form of zoolatry.2

In Belgium there are still traces of the old
custom of sacrificing an animal upon the com-
pletion of a new building, with the idea that the
animal's spirit will protect the edifice from harm,3
and if the observer of the following facts is correct
in his interpretation of them, there also remain in
Belgium traces of the ancient sacrifices to the
genius of the earth : —

" It is the custom, round about Florenville
(in the Belgian Ardennes), to offer a sacrifice to
the presiding genius of the road upon the con-
struction of a new road or railway. It is usually
a fowl, or a rabbit, or even a calf which is sacri-
ficed. ... In some parts of Luxembourg animals

1 Kowalevsky, Droit coutmnier Osseticn, p. 105.

2 Spencer, Principles of Socioloi/i/.

3 Folklore, Wallon, No. 1526, p. 115 (Bulletin de Folklore, ii.,
177).

314 CAUSES OF DEGENERATIVE EVOLUTION

are also offered in sacrifice to the genius supposed
to preside over a newly-purchased field, with a
view to ensuring abundant crops.1

Eemains, either undoubted 2 or only probable,3
of phallic worship, are scattered throughout Europe.

In Brittany 4 and in Belgium,5 for instance,
strange old customs still exist showing that here,

1 Rivue dcs traditions pqpu I 'aires, 1893, p. 394.

2 Th. Volkov, Bites ct usages nuptiaux en Ukraine (I'Anthro-
pologie, 1891, p. 167).

Only a short time ago it was the custom in Tver, on the day
dedicated to Yarilo (the phallic God of Spring), for the parents
of young daughters to send them to join in games similar to
those of the ancient Slavs, with a view to their getting married.

3 Note sur un vestige du culte de la terre mere (phallism) en
Provence, by Berenger-Feraud (Re'mie (V Anthropologic, 1888, p.
563).

" At Luc, in Provence, upon the 1st of May, which is a country
holiday, the young girls proceeded to a place where two roads met.
Here they assembled around an olive tree, and after each dance
they struck the olive tree three times with their backs.

" This fete, a survival of the floral fetes of the month of May
which are still celebrated in Provence and Italy, continued to be
held until quite recently, and appears to have been a lingering
vestige of the ancient worship of creative Nature, Mother-earth
— in short, of phallic worship.

" The three knocks given by the young girls to the tree trunk
is a survival of the ancient virginal sacrifice to the phallic
emblem. The original meaning was not quite lost, for the Pro-
vencals still realized, though vaguely, that the three knocks were
somehow connected with the idea of marriage."

4 ' ' Les Me'galithcs de Locmariaquer et de Carnac, et les amours, by
Bonnemere {Revue dcs traditions populaires, 1894, p. 123). "In
former days it was the custom for all the young women who wished
to get married to climb (on the night of May 1), to the top of the
great menhir where they lifted up their clothing that their bodies

SURVIVAL OF INSTITUTIONS 315

as elsewhere, a belief in the influence of fetiches
once prevailed, and particularly in the form of
megalithic monuments relating to the fecundity of
women.

The custom still prevalent in the African Congo,
of driving a nail into a fetich, with the view of
reminding it of a request, has not disappeared from
Europe. The young men of Couvin (Namur) still
stick pins into the wooden saints of the little
chapels round about in order to draw a lucky
number in the military lottery, and young girls in
Brittany do the same with a view to getting

might come in direct contact with the stone, and then slid from
the top to the hottom."

At Carnac, young girls wishing to marry undressed completely
upon the same night, and proceeded to rub their abdomen against
a special menhir. In that part of Brittany, where only French is
spoken, similar customs have erpially prevailed.

5 Fete de Notre-Dame de Ride-cul (Jules Lemoine, in the journal
Le petit bleu, of October 18, 1896).

Similar to- the above Breton custom is that of sliding down the
Rocher Ridc-cul, which is situated near to Landelies, in the valley
of the Sambre. Here, as in many other places, old customs have
been Christianized, and a Christian chapel now stands close to the
ancient shrine. Young people of both sexes used to seat them-
selves upon the top of the stone upon little fagots of boxwood
gathered in the neighbourhood, and then slide down to the bottom.

According to the old saying an upset meant waiting ; an em-
brace signified mutual affection ; a collision, indifference ; and an
embrace followed by rolling over indicated matrimonial suitability.
Similar customs prevailed at Trou-deux-Trous, situated near to
the Rocher Ride-cul. These two megalithic temples disappeared
about forty years ago, their materials have been used as fluxes in
smelting works.

316 CAUSES OF DEGENERATIVE EVOLUTION

married,1 and in Belgium,2 as in Brittany,3 if the
image prove too hard, the reminding pin is stuck
instead into a fissure, or into the door of the niche
containing the image. Besides these mere vestiges
of pre-historic customs and belief, by referring to
certain illustrated documents bearing upon the
subject, it would be easy to work out the connec-
tion between the worship of saints which is pre-
valent in Belgium in the present day, and the
pagan worship of the ancient Celts and Germans.4

1 Bulletin de Folklore, i., 250-251.

2 Ibidem.

3 The menhir of the Pierre-Frite in the valley of Lunain. In
nearly every hole or fissure of this monument, a nail or pin has
been stuck by the young people of that part in the belief that it
will ensure them a speedy marriage. (Be'vuc des traditions popu-
lates, 1893, p. 448.)

4 See for Saint Eloi, Mdnsine, viii., 122-132 ; for Saint Martin,
Bulletin de Folklore, i., 309-315 ; for Saint Hubert, Gaidez, La
Rage et Saint Hubert.

PART III

RESUME AND CONCLUSIONS

When an institution or an organ ceases to be
functional or in any way useful, it very soon
disappears altogether. If, as happens in some
exceptional cases, it persists, it is because neither
of the chief factors in causing atrophy, variability
or selection, have intervened.

Sometimes the vestiges are of too insignificant
a nature to call for their removal by either
artificial or natural selection, and sometimes their
existence is ensured by the lack of variability, as
in the case of the persistence of flowers in plants
which multiply asexually. This absence of varia-
tion occurs equally in the social domain, especially
in matters connected with religion, wherein ancient
customs are credited with a divine origin. Eeligions
may pass away, philosophies may be transformed,
and old beliefs cease to prevail, but the remnants
of old creeds, conveyed by popular tradition
through the centuries, defy destruction by modern
innovations.

The ancient winter festival, on which day the

318 CAUSES OF DEGENERATIVE EVOLUTION

dead were supposed to leave their graves and join
the living in a feast around the family hearth, is
still celebrated in the keeping of Christmas and in
the various customary practices on the first two
days of November.

The May-Day festivals — pagan festivals held in
honour of vegetable and human fecundity — are
still held in their early form round about
Locmariaquer and in the village of Campine.
Traces also remain in the picking and wearing
of flowers on the 1st of May, and the same day
is selected by the socialists for the celebration of
their near approach to a life under freer and
happier conditions.

This survival of festivals, customs and traditions,
while the religions and civilizations which produced
them have passed away, is the principal link which
connects us with bygone generations.

" Their value lies," says Houzeau in his Etude de
la Nature, " in the establishment of a chain between
successive generations. The memory of an indivi-
dual may be regarded as constituting his personality.
Take from him the memory of his past, and he is left
at a point in time wherein there is no stability and
complete isolation. To be himself, a man requires
not only his recollections, but a knowledge of his past
habits and traditions. When a savage is removed
from his fellows and transported to new surround-
ings in a distant country, he loses all knowledge of
his former condition. Society itself, made up as it

SUMMARY AND CONCLUSIONS 319

is of customs and prejudices, constitutes history.
The mirror of the past is exhibited in the conscious-
ness of the collective individual which is called
a nation. What link shall we have with former
generations if not a heritage of their ideas — i.e. of
their discoveries and their mistakes ? Nations,
like individuals, are continually modifying this
inherited legacy, but, like the individual, they
cannot get away from it without breaking the
thread which has made them themselves."

GENERAL CONCLUSIONS

All evolution is at once progressive and retro-
gressive.

All modifications of organs and institutions are
attended by retrogression. This occurs equally
in the modifications of organisms and of societies.
All existing forms, whether organic or social, have
undergone certain modifications, and, as a result,
have lost some parts of their structure. This
universality of degenerative evolution may be
proved either by the comparative method, or by
showing that all organisms contain rudimentary
organs, and that all societies contain survivals.

II

Degenerative evolution follows no definite path,
and can in no way be regarded as constituting a
return to the primitive condition.

In some cases — when one cause of dissolution
equally and simultaneously affects all the parts of an
institution or an organism — the most complicated
and delicate structures are the first to disappear ;
but it must not be taken as a general principle
that the most complicated structures are necessarily

GENERAL CONCLUSIONS 321

the most recent, and that consequently degeneration
always retraces the path of progress. Evolution
is irreversible, and accordingly, with a few more
or less obvious exceptions, we draw the following
conclusions : —

1. That an institution or an organ which has

once disappeared never reappears.

2. That an institution or organ once reduced

to the condition of a vestige cannot be
re-established and resume its former func-
tions.

3. Neither can they assume fresh functions.

Ill

Degenerative evolution is brought about by a
limitation in means of subsistence — either in
nutriment, capital or labour. In biology the prin-
cipal if not the sole agents in its accomplishment
are the struggle for existence between the various
organs, and the struggle for existence between the
various organisms.

In sociology it is artificial selection which is the
dominating agent, and natural selection plays only a
secondary part.

The occasional causes of degenerative evolution
are inutility of function, insufficiency of nutri-
ment or resource, and (in biology only) lack of
space.

x

322

GENERAL CONCLUSIONS

An institution or an organ which has ceased to
be functional, and has also ceased to be useful either
directly or indirectly, continues to exist if neither
variability or selection intervene.

TCRNBCLL AND SPEARS, PRINTERS EDINBURGH.

Date Due

1 U

MM-i&JH

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/ERSITY OF B C LIBRARY

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