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Democracy and Dictatorship in Europe
 Democracy and
Dictatorship
Sheri Berman in Europe
From the Ancien Régime
to the Present Day

1
1
Oxford University Press is a department of the University of Oxford. It furthers
the University’s objective of excellence in research, scholarship, and education
by publishing worldwide. Oxford is a registered trade mark of Oxford University
Press in the UK and certain other countries.

Published in the United States of America by Oxford University Press

198 Madison Avenue, New York, NY 10016, United States of America.

© Oxford University Press 2019

All rights reserved. No part of this publication may be reproduced, stored in

a retrieval system, or transmitted, in any form or by any means, without the
prior permission in writing of Oxford University Press, or as expressly permitted
by law, by license, or under terms agreed with the appropriate reproduction
rights organization. Inquiries concerning reproduction outside the scope of the
above should be sent to the Rights Department, Oxford University Press, at the
address above.

You must not circulate this work in any other form

and you must impose this same condition on any acquirer.

Library of Congress Cataloging-in-Publication Data

Names: Berman, Sheri, 1965– author.
Title: Democracy and dictatorship in Europe : from the Ancien régime to the
present day / Sheri Berman.
Description: New York, NY : Oxford University Press, [2018]
Identifiers: LCCN 2018027023 (print) | LCCN 2018044301 (ebook) |
ISBN 9780199373208 (updf) | ISBN 9780199373215 (epub) |
ISBN 9780199373192 (hardcover)
Subjects: LCSH: Europe—Politics and government. |
Democracy—Europe—History. | Dictatorship—Europe—History.
Classification: LCC JN8 (ebook) | LCC JN8.B47 2018 (print) | DDC 320.94—dc23
LC record available at https://lccn.loc.gov/2018027023

9 8 7 6 5 4 3 2 1

Printed by Sheridan Books, Inc., United States of America

To those who have struggled to get rid of the ancien régime.
CON TEN TS

Acknowledgments ix

1 Questions about Political Development 1

2 The Ancien Régime 15
3 English Exceptionalism I 29
4 The French Revolution 49
5 1848 77
6 The French Third Republic 106
7 Italian Unification 125
8 German Unification 146
9 The Struggle for Democracy in Interwar
France 169
10 English Exceptionalism II 185
11 The Collapse of Democracy and the Rise of
Fascism in Italy 214
12 The Collapse of the Weimar Republic and the
Rise of National Socialism in Germany 234
13 Political Development in Spain 257
14 The Consolidation of Democracy in Western
Europe 284
 15 The Transition to Communist Dictatorships in
East-​Central Europe 303
16 The Transition to Democracy in Spain 327
17 The Transition to Democracy in East-​Central
Europe 347
18 Lessons from Europe 376

Notes 409
Index 517

viii   c o n t e n t s
ACK NOW LEDGME NTS

This book has been long—​too long—​in the making. Indeed at many points
I was not sure it would get done. The only advantage of having taken so long
is that I have read and learned more about European political history and
political development than I ever thought possible. During the many years
I worked on this book I tried out parts of the argument on many audiences. To
those who came to my talks, panels, and seminars, I offer immense thanks—​
your questions and criticisms pushed me to think harder about what I wanted
to say and how to say it better. During the time this book was percolating
I have also published versions its arguments in various venues. I would like
to thank the Journal of Democracy and Dissent in particular for publishing
several of these essays and their editors for proving feedback on them. My
greatest intellectual debt, however, goes to the many Barnard and Columbia
students who have taken my “Democracy and Dictatorship in Europe” class
over the last decade. This book grew out of this class; after teaching it for
several years I realized there was no single book that covered what the course
was trying to achieve and that maybe I could be the one to write that book.
And each year I taught the course I re-​worked the lectures to better commu-
nicate important arguments and themes to my students and these lectures
provide the intellectual infrastructure for the book that follows. Without this
process of constant revision and re-​thinking I would never have been able to
put the many pieces of the puzzle that is European political development to-
gether, so again thank you to the students who pushed me to keep revising
and improving. I would also like to thank the many teaching assistants who
helped me teach this class and the research assistants who helped me put
together the figures in the book that follows. And finally, I am grateful to
Oxford University Press and David McBride for publishing this book and
being patient while I struggled to finish it off.
Although they did not directly help with the book, I would like to thank
Isaac and Lucy for pushing me and making me determined to show that
if you work hard and do not give up you can accomplish what you set out
to do. I would also like to thank my wonderful fuzzies for comforting me
when I wanted to throw another book I realized I needed to read against the
wall or chuck another chapter revision in the trash bin. And finally thanks
to the friends who helped me get through my own process of overcoming
the ancien régime. Just as this book shows that liberal democracy cannot be
consolidated without undergoing this long and painful process, so too did
this book’s completion require it.

x   a c k n o w l e d g m e n t s
Democracy and Dictatorship in Europe
 Chapter 1 Questions about Political
Development

You have to know the past to understand the present.

—​Carl Sagan1

I n november 1989 the place to be was Berlin. History was being made.
Dictatorships were collapsing and, as George H.W. Bush put it, “America
[had] won the Cold War.”2 The bloody twentieth century with its horrible
violence and titanic ideological battles was coming to a close. Liberal democ-
racy was triumphant, and the West was about to lead the way towards prog-
ress and prosperity. Many felt in 1989 as Wordsworth did about the French
Revolution exactly two hundred years before: “Bliss was it in that dawn to be
alive, /​But to be young was just heaven.”3
Europe, of course, was particularly euphoric. The downfall of communism
in Eastern Europe combined with the collapse of dictatorships in Southern
Europe during the previous decade left the continent more united than ever
before. For the first time in its modern history European countries shared
the same political (democratic) and economic (capitalist) system; Germany
and Russia—​the great powers that had caused so much instability in the
past—​were no longer threats; and the European Union was on the verge of
incorporating much of Eastern Europe and creating a single currency. At the
end of the twentieth century, the view that a united Europe was on its way
to becoming “the next global superpower,” and the model for a new type of
peaceful, prosperous, post-​national polity, was widespread.4
 How fickle is History. Today the optimism of 1989 is long gone, replaced
by fears that Europe and the West have entered a period of decline. Liberal
democracy has faltered in Eastern Europe and is threatened by populism in
Western Europe and the United States. Scholars and commentators no longer
talk about the triumph of liberal democracy, the “end of history,” and “post-​
national” politics, but worry instead about “illiberal democracy,” “global au-
thoritarianism,” virulent nationalism, and democratic “deconsolidation.”5
As Viktor Orbán, Hungary’s current prime minister, whose political career
began in 1989 as an opponent of dictatorship but who has since morphed
into an opponent of democracy, recently proclaimed: “The era of liberal de-
mocracy is over.”6
How did we get from there to here? How can we understand the fate of
today’s new democracies and the problems facing its older ones? What makes
liberal democracy work well in some places and some times, but not others?
These are questions of the utmost theoretical and practical importance.
The rapid spread of democracy at the end of the twentieth and beginning
of the twenty-​first century combined with its current problems has placed
questions concerning the origins, evolution, and fate of democracy at the
forefront of contemporary debate. This book will add a fresh perspective to
this debate. From historians we have gotten superb studies of particular coun-
tries’ politics at particular points in time. From political scientists we have
gotten theories about what it takes to make democracy work.7 And more
recently, scholars and commentators have provided analyses of democracy’s
current crisis.8 This book draws on and aims to complement the work of
historians, political scientists, and contemporary analysts of democracy. It is
grounded in the view, as Carl Sagan put it, that one “has to know the past
to understand the present.” It will integrate questions being asked about
democracy today with a re-​consideration of European political development
in order to gain a better understanding of how and why democracies and
dictatorships developed in the past. And by reconsidering how democracies
and dictatorships developed in the past it will add a historical perspective
that will enable us to better understand what is going on in the world today.
Europe is the perfect place to ground such an examination: it is the place
where modern democracy was born and is currently the home of a large crop
of well- and not-​so-​well-functioning democracies. In addition, Europe has
also been the home of dictatorships of all kinds—​monarchical, military, pop-
ulist, hybrid, competitive, fascist, and National Socialist. These democracies
and dictatorships emerged, moreover, at various times and in various eco-
nomic and social contexts. Indeed over the course of the modern era, European
countries have differed in almost every way political scientists identify as

2   d e m o c r a c y and dictatorship in europe

causally significant for democratic development: timing, nature, and ra-
pidity of economic development; levels of economic inequality; strength of
national states; degree of ethnic and religious diversity; and more. That so
many types of political regimes and such immense economic and social diver-
sity have existed in Europe provides an opportunity to study how democracies
and dictatorships develop and decay over time and in various contexts.
In addition to placing the European experience in broader context and
bringing a historical perspective to the consideration of contemporary cases,
a re-​examination of political development in Europe can help us understand
the problems its own democracies face today. After a half century of un-
precedented political, social, and economic success, Europe is once again
experiencing profound political malaise and even political disorder. Most
commentary on Europe’s present problems has focused on contemporary
causes like globalization, technological change, the financial crisis, technoc-
racy, immigration, and refugee flows. This book will show, however, that
short-​term analyses misunderstand or at least oversimplify what is going
on. In particular, the historical perspective adopted in this book will help
us see how Europe’s current problems have their roots in the decay of the
foundations upon which liberal democracy was (re)built after the tragedies of
the interwar period and the Second World War.
Before moving on, it is necessary to briefly define some key concepts used
throughout the book: democracy, democratization, and consolidation. The
first seems straightforward, but in fact students of democracy have almost
as many definitions of it as there are regimes in existence.9 The latter two,
meanwhile, although conceptually and practically related, represent different
stages or parts of the political development process; too often contemporary
commentary conflates or fails to clearly differentiate them, leading to con-
fusing analyses, assertions, and policy recommendations. Some conceptual
brush-​clearing, in short, is necessary before the analysis begins.

Key Concepts
The term “democracy” derives originally from the Greek word for popular
rule, demokratia (demos = “common people,” and kratos = “rule” or “power”).10
Although defining democracy as “rule by the people” seems uncomplicated,
it is actually too ambiguous to be used to analyze or compare political sys-
tems. For example, who are “the people”? Does rule by “the people” mean all
“the people” or just some subgroup of them? And how about “rule”? Do “the
people,” however defined, have to rule directly in order for a system to be

questions about political development 3

considered democratic or should indirect, mediated, or technocratic rule also
be considered democratic? Must “the people” agree on all political decisions
or at least share a conception of the “common good” in order for a political
system to be considered democratic or can there be conflicts among “the
people” on what “rule” should entail in a democratic system?
Because of questions like these, social scientists have tried to come up
with definitions of democracy better suited to empirically analyzing and
comparing political regimes. Perhaps the one most oft used by scholars
and policy makers today is a “minimal” or “electoral” definition which
characterizes political regimes based on their procedures, particularly those
used to pick rulers. In this view, the central procedure of democracy—​and
the feature that distinguishes it most clearly from other types of political
regimes—​is the selection of leaders through competitive elections by the
people they govern. The most important modern statement of this defini-
tion of democracy was given by Joseph Schumpeter in his classic Capitalism,
Socialism and Democracy. According to Schumpeter, “the democratic method
is that institutional arrangement for arriving at political decisions in which
individuals acquire the power to decide by means of a competitive struggle
for the people’s vote.”11 For Schumpeter, in other words, the need for leaders
to compete for support or win elections rather than coming to power via
coercion or heredity is the most important distinction between democracy
and dictatorship. It is important to note that while its focus on elections is
fairly simple and straightforward, this definition of democracy also implic-
itly includes a number of civil and political rights, such as some degree of
freedom of speech, assembly, and organization since without them, elections
could not be considered truly competitive, or “free and fair.”
This “minimal” or “electoral” definition of democracy has many
advantages. First, it is intuitive; most people recognize free and fair elections
as the central feature of democracy. Second, it provides concrete, empirical
benchmarks that can be used to differentiate democracy from dictatorship: if
a country denies the right to participate in free and fair elections to its citi-
zenry, then it is not democratic. Third, in addition to enabling us to judge
the nature of any particular political system at any particular point in time,
this definition also helps us analyze whether it is moving in a democratic
or non-​democratic direction: as elections become “freer and fairer” and/​or
increasing numbers of citizens are able to participate in them, countries can
be characterized as becoming more democratic, or democratizing.
The “minimal” or “electoral” definition of democracy thus allows for
relatively clear-​cut and straightforward comparisons of political regimes.
Some scholars, however, think the benefits of this definition’s parsimony are

4   d e m o c r a c y and dictatorship in europe

outweighed by costs in terms of accuracy. In particular, critics assert that
proponents of this definition are guilty of the “fallacy of electoralism”—​of
assuming that relatively free and fair elections indicate a well-​functioning
democracy. Surely anyone who reads a newspaper today can come up with
examples of relatively free and fair elections gone awry. For example, many
currently worry about the growing popularity of populist parties and
politicians in the West whose commitments to liberalism and perhaps even
democracy are uncertain. Similar concerns were voiced during the Arab
Spring when many worried that elections would bring to power Islamists
whose commitments to democracy were weak or instrumental. Indeed,
such concerns were invoked in Egypt to help justify the overthrow of the
Muslim Brotherhood’s Mohamed Morsi and when the Algerian government
voided the results of a democratic election that was about to be won by the
radical Islamic Salvation Front (FIS) in 1992. Commenting on the United
States’ support of the Algerian government in this latter case, then Assistant
Secretary of State Edward Djerejian explained that the United States favored
democratization but did not support groups that believed in “one person,
one vote, one time.”12 Such concerns, of course, are nothing new. Infamously,
both Hitler and Mussolini pursued an “electoral path” to dictatorship; this
book will discuss many other cases from European history where elections
brought to power politicians and parties with at best tenuous commitments
to democracy.
In addition to worrying about elections leading to governments that
subsequently undermine democracy, critics also worry about democrati-
cally elected governments using their power in “non-​democratic” or illiberal
ways. Once again anyone who pays attention to current events could come
up with numerous contemporary examples of such behavior. For example,
Orbán in Hungary, the Law and Justice party in Poland (led by Jarosław
Kaczyński), and Recep Tayyip Erdoğan in Turkey all came to power via rela-
tively free and fair elections and then used their power to reward supporters
and punish detractors, and more generally undermine democratic procedures
and institutions. And also in the still fairly recent European past, democrat-
ically elected leaders in the former Yugoslavia sanctioned or incited violence
against minorities in their own countries. As Richard Holbrooke famously
asked before the 1996 elections in Bosnia, “Suppose elections are free and fair
and those elected are racists, Fascists, separatists? That is the dilemma.”13
Because it is not hard to find cases of democratic elections gone awry,
and because many believe that while necessary, free and fair elections are not
sufficient to make a political system fully democratic, many scholars argue
for going beyond a minimal or electoral definition of democracy and favor

questions about political development 5

a “liberal” one instead. As the term “liberal” indicates, proponents of this
view argue that democracy is not merely a political system that chooses its
leaders in a particular way; it is also a political system where government
and citizens act in particular ways, namely in accordance with the tenets
of liberalism. As the American Declaration of Independence declared: “We
hold these truths to be self-​evident, that all men are created equal, that they
are endowed by their Creator with certain unalienable Rights, that among
these are Life, Liberty and the pursuit of Happiness.” In this view, in order to
be considered truly or fully democratic, a government must be willing and
able to guarantee the rule of law; and must protect minorities and individual
liberties; and leaders and citizens must respect the democratic “rules of the
game,” and treat all members of society as political equals. The “liberal”
definition of democracy is thus deeper than the electoral or minimal one.
It incorporates into our understanding of democracy a set of values or goals
that cannot be secured by political procedures alone. Democracies differ from
dictatorships, in other words, not only in the way in which they choose their
leaders, but also in the way they treat their citizens and their citizens treat
each other.
The liberal definition of democracy has real advantages. First, it avoids
the “fallacy of electoralism”: it does not assume that particular procedures,
most importantly relatively free and fair elections, are all it takes to make de-
mocracy. Second, it explicitly incorporates a wide variety of liberal rights that
many view as inherently part of any truly democratic system. Third, it allows
us to engage in more fine-​grained and multifaceted, if somewhat fuzzier and
more complicated,14 comparisons of political regimes than does the minimal
or electoral definition of democracy. Indeed, the more stringent requirements
of liberal democracy clearly eliminate countries like Hungary, Poland, and
Turkey and many others from the democratic category.15 Precisely to cap-
ture the fact that many democracies are not liberal, another political label—​
illiberal democracy—​is often invoked. Like electoral democracies, illiberal
democracies have relatively free and fair elections but few if any of the other
trappings of liberal democracies: they do not fully or indiscriminately apply
the rule of law, protect minorities, or ensure individual liberties.
European history offers the perfect context within which to examine the
nature, dynamics, and implications of different types of democracy. The fol-
lowing chapters investigate how common these different types of democracy
have been; the factors determining whether a country develops an electoral,
illiberal, or liberal democracy; and whether and how the development of il-
liberal democracy affects the development of liberal democracy down the
road.16 Some scholars, for example, assert that countries that democratize

6   d e m o c r a c y and dictatorship in europe

before the conditions to sustain liberal democracy are in place are likely to
become “stuck” in illiberalism. As one influential treatment of the topic puts
it, “Premature, out-​of-​sequence attempts to democratize may make subse-
quent efforts to democratize more difficult and more violent than they would
otherwise be.”17 Is this true? Has electoral or illiberal democracy precluded
progress towards liberal democracy in the past? Are electoral or illiberal and
liberal democracy “competing” regime types or rather different stages of po-
litical development?
Alongside investigating the nature, dynamics, and implications of dif-
ferent types of democracy, two other key foci of Democracy and Dictatorship in
Europe are democratization and consolidation.
Democratization refers to the transition from dictatorship to democ-
racy. It is a process, not a stable condition. Democratization implies nothing
about the durability or health of democracy. Democratization may occur
and the new democracy may be weak or short-​lived. Once again, European
history provides a great context for investigating how and why democra-
tization occurs and how often it leads to stable democracy. The following
chapters examine many cases of democratization in Europe from the seven-
teenth century on in order to shed light on how and why dictatorial regimes
are replaced by democratic ones and to determine how common it has been
for democratization to lead to consolidation rather than instability or dem-
ocratic failure.
Consolidation is what potentially happens after democratization: it is
when the collapse of a dictatorship leads to the formation of a successful de-
mocracy. But how should democratic success be measured? Some scholars and
observers focus on durability. In this view, democracy should be considered
consolidated if it lasts for some significant period of time. But what should
that period be? Five years? Ten years? A generation? Because any particular
time period might seem arbitrary, some prefer a measure of durability that
defines a democratic regime as consolidated when it has successfully carried
out a certain number of elections.18
If one rejects, however, an electoral or minimal definition of democracy,
then a purely quantitative definition of consolidation focused on durability
or elections will be insufficient. Many scholars and observers accordingly
favor a qualitative definition of consolidation that focuses on the attitudinal
and behavioral shifts that comport with liberal democracy. In this view, a
consolidated democracy requires that all groups are allowed to participate in
political life and voice their demands, as long as they do so within the “rules
of the game”; the basic rights of minorities and individuals are respected
by the government as well as other citizens; and support for democracy is

questions about political development 7

Another random document with
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Fig. 80.—Dorsal view of male Diastylis
stygia, × 12. A, 2nd antenna; Ab.6, 6th
abdominal appendage. (After Sars.)

Order II. Cumacea.[96]

The Cumacea are a group of small marine animals rarely attaining

an inch in length, which agree with the Mysidacea in the characters
noted above as diagnostic of the Division Peracarida; they possess,
however, in addition a number of peculiar properties, and Sars
believes them to be of a primitive nature showing relationship to
Nebalia, and possibly to an ancestral Zoaea-like form. They follow a
habit similar to that of the Mysidacea, being caught either in the
surface-plankton or in great depths, many of the deep-sea forms
being blind. They are, however, not true plankton forms, and they
appear to attain a greater development both in point of variety and
size in the seas of the northern hemisphere. The thoracic limbs may
be biramous, but there is a tendency among many of the genera to
lose the exopodites of some of the thoracic legs, an exopodite never
being present on the last few thoracic limbs of the female and on the
last in the male. In the Cumidae the four posterior pairs in both sexes
have no exopodites. The first three thoracic appendages following the
maxillae are distinguished as maxillipedes; they are uniramous, and
the first pair carries an epipodite and a large gill upon the basal
joints. Pleopods are only developed in the male sex.
The flagellum of the second antennae in the male may be
enormously elongated, as in the Atlantic deep-sea species shown in
Fig. 80, so as to exceed in length the rest of the body.
Fam. 1. Cumidae.—No sharp demarcation between thorax and
abdomen. Four posterior pairs of legs in both sexes without
exopodites. Male with five well-developed pleopods in addition to the
uropods. Telson wanting. Cuma, Cyclaspis, etc.
Fam. 2. Lampropidae.—Body-form resembles that of Cumidae.
All the thoracic limbs except the last have exopodites. The male has
three pairs of pleopods. Telson present. Lamprops, Platyaspis, etc.
Fam. 3. Leuconidae.—Body-form similar to above. Male has
only two pairs of pleopods. Mouth-parts peculiar, much less setose
than in other families. Telson absent. Leucon, Eudorella.
Fam. 4. Diastylidae.—Anterior part of thorax sharply marked
off from posterior part. Male has two pairs of pleopods. Telson
present. Diastylis (Fig. 80). D. goodsiri from the Arctic ocean
measures over an inch in length.
Fam. 5. Pseudocumidae.—Rather similar to Diastylidae, but
differ in reduced size of telson and presence of exopodites on third
and fourth thoracic legs of female. This family is represented by
three very similar marine forms of the genus Pseudocuma; but, as
Sars has shown,[97] the Caspian Sea contains thirteen peculiar
species, only one of which can be referred to the genus Pseudocuma,
while the rest may be partitioned among four genera, Pterocuma,
Stenocuma, Caspiocuma, Schizorhynchus.

Order III. Isopoda.

The Isopoda and the Amphipoda are frequently classed together as

Arthrostraca or Edriophthalmata, owing to a number of features
which they share in common, as, for instance, the sessile eyes which
distinguish them from the podophthalmatous Schizopoda and
Decapoda, the absence of a carapace, and the thoracic limbs which
are uniramous throughout their whole existence. For the rest, in the
presence of brood-plates and the other diagnostic characters, they
are plainly allied to the other Peracarida, and an easy transition is
effected from the Mysidacea to the Isopoda through the Chelifera or
Anisopoda. Only one thoracic segment is usually fused with the head,
the appendage of this segment being the maxillipede; in the Chelifera
among Isopoda, and the Caprellidae among Amphipoda, two
thoracic segments are fused with the head.
The Isopoda are distinguished from the Amphipoda by the dorso-
ventral flattening of the body, as opposed to the lateral flattening in
the Amphipoda, by the posterior position of the heart, and by the
branchial organs being situated on the abdominal instead of on the
thoracic limbs.
The Isopoda, following Sars’[98] classification, fall into six sub-
orders—the Chelifera, Flabellifera, Valvifera, Asellota, Oniscoida,
and Epicarida,—to which must be added the Phreatoicidea.
 Sub-Order 1. Chelifera.

The Chelifera, including the families (1) Apseudidae and (2)

Tanaidae, are interesting in that they afford a transition between
the ordinary Isopods and the Mysidacea. The important features in
which they resemble the Mysidacea are, first, the fusion of the first
two thoracic segments with the head, with the coincident formation
of a kind of carapace in which the respiratory functions are
discharged by a pair of branchial lamellae attached to the
maxillipedes; and, second, the presence of very small exopodites on
the first two thoracic appendages of the Apseudidae.
The second pair of thoracic limbs, i.e. the pair behind the
maxillipedes, are developed both in the Apseudidae and Tanaidae
into a pair of powerful chelae, and these frequently show marked
sexual differences, being much more highly developed in the males
than in the females. The biramous and flattened pleopods are purely
natatory in function, and the uropods or pleopods of the sixth pair
are terminal in position and slender.
Both families, of which the Apseudidae contain the larger forms,
sometimes attaining to an inch in length, are littoral in habit, or
occur in sand and ooze at considerable depths, many of the genera
being blind. Many Tanaids (e.g. Leptochelia, Tanais, Heterotanais,
etc.) live in the algal growths of the littoral zone, and being highly
heliotropic they are easy to collect if a basinful of algae is placed in a
strong light. The females carry the eggs about with them in a brood-
pouch formed, as is usual in the Peracarida, by lamellae produced
from the bases of the thoracic limbs. The males on coming to
maturity do not appear to grow any more, or to take food, their
mouth-parts frequently degenerating and the alimentary canal being
devoid of food. They are thus in the position of insects which do not
moult after coming to maturity; and, as in Insects, the males are apt
to show a kind of high and low dimorphism—certain of the males
being small with secondary sexual characters little different from
those of the females, while others are large with these characters
highly developed. Fritz Müller, in his Facts for Darwin, observes
that in a Brazilian species of Leptochelia, apparently identical with
the European L. dubia, the males occur under two totally distinct
forms—one in which the chelae are greatly developed, and another in
which the chelae resemble those
of the female, but the antennae in
this form are provided with far
longer and more numerous
sensory hairs than in the first
form. Müller suggested that these
two varieties were produced by
natural selection, the characters
of the one form compensating for
the absence of the characters of
the other. A general consideration
of the sexual dimorphism in the
Tanaidae[99] lends some support
to this view, since the smaller
species with feeble chelae do
appear to be compensated by a
greater development of sensory
hairs on the antennae, but the
specific differences are so difficult
to appreciate in the Tanaidae that
it is possible that the two forms of
the male in Müller’s supposed
single species really belonged to
two separate species.
 Fig. 81.—Apseudes spinosus, ♂, × 15. A,
1st antenna; Ab, 6th abdominal
appendage; T, 2nd thoracic appendage.
(After Sars.)

Sub-Order 2. Flabellifera.

The Flabellifera include a number of rather heterogeneous families

which resemble one another, however, in the uropods being lateral
and not terminal, and being expanded together with the telson to
form a caudal fan for swimming. The pleopods are sometimes
natatory and sometimes branchial in function. Some of the families
are parasitic or semi-parasitic in habit.
Fam. 1. Anthuridae.—These are elongated cylindrical creatures
found in mud and among weeds upon the sea-bottom; their mouth-
parts are evidently intended for piercing and sucking, but whether
they are parasitic at certain periods on other animals is not exactly
known. Anthura, Paranthura, Cruregens.
Fam. 2. Gnathiidae.[100]—These forms appear to be related to
the Anthuridae; they are ectoparasitic on various kinds of fish during
larval life, but on assuming the adult state they do not feed any more,
subsisting merely on the nourishment amassed during the larval
periods. The larvae themselves are continually leaving their hosts,
and can be taken in great numbers living freely among weeds on the
sea-bottom. The larvae, together with the adults of Gnathia
maxillaris, are extremely abundant among the roots of the sea-weed
Poseidonia cavolinii in the Bay of Naples. The young larvae hatch
out from the body of the female in the state shown in Fig. 82, A. This
minute larva fixes upon a fish, and after a time it is transformed into
the so-called Praniza larva (B), in which the gut is so distended with
the fluid sucked from the host that the segmentation in the hind part
of the thorax is entirely lost. When this larva moults it may, however,
reacquire temporarily its segmentation. After a certain period of this
parasitic mode of life the Praniza finally abandons its host, and
becomes transformed into the adult male or female. This may take
place at very different stages in the growth of the larva, the range of
variation in size of the adults being 1–8 mm., and it must be
 remembered that when once the
adult condition is assumed
growth entirely ceases. What it is
that determines the stage of
growth in each individual when it
shall be transformed into the
adult is not known. The males
and females differ from one
another so extraordinarily that it
was for long denied that they
were both derived from the
Praniza larvae. This is
nevertheless the case. The change
from the Praniza to the female
(Fig. 82, C) is not very great. The
ovary absorbs all the
nourishment in the gut and
comes to occupy the whole of the
body, all the other organs
degenerating, including the
alimentary canal and mouth-
Fig. 82.—Gnathia maxillaris. A, parts. Indeed, only the limbs with
Segmented larva, × 10; B, Praniza their muscles and the nervous
larva, × 5; C, gravid female, × 5; D, system remain. The change to the
male, × 5.
male (D) is more radical. The
food is here stored in the liver,
which increases in the male just as the ovary does in the female. The
segmentation is reacquired, and the massive square head is formed
from the hinder part of the head in the Praniza, the anterior portion
with its stylet-like appendages being thrown away. The powerful
nippers of the male are not formed inside the cases of the old
styliform mandibles, but are independent and possibly not
homologous organs. The meaning of the marked sexual dimorphism
and the use of the males’ nippers are not in the least known, though
the animals are easy to keep under observation. In captivity the
males never take the slightest notice of either larval or adult females.
Fam. 3. Cymothoidae.[101]—This is a group of parasites more
completely parasitic than the foregoing, but their outer organisation
does not differ greatly from an ordinary Isopodan form. A great
many very similar species are known which infest the gill-chambers,
mouths, and skin of various fishes. The chief interest that attaches to
them is found in the fact that a number of them, and perhaps all, are
hermaphrodite, each individual acting as a male when free-
swimming and young, and then subsequently settling down and
becoming female. This condition is exactly the same as that
occurring universally in the great group of parasitic Isopoda, the
Epicarida, to be considered later. There is no evidence that the
Cymothoidae are phyletically related to the Epicarida, so that the
similar sexual organisation appears to be due to convergence
resulting from similar conditions of life. The general question of
hermaphroditism in the Crustacea has been shortly discussed on pp.
105–106. Cymothoa.
Fam. 4. Cirolanidae.—In this family is placed the largest Isopod
known—the deep-sea Bathynomus giganteus, found in the Gulf of
Mexico and the Indian Ocean, sometimes measuring a foot long by
four inches broad. A common small littoral form is Cirolana.
Fam. 5. Serolidae.[102]—The genus Serolis comprises flattened
forms bearing a curious resemblance to Trilobites, which Milne
Edwards considered more than superficial. The genus is confined to
the littoral and deep waters of the southern hemisphere.
Fam. 6. Sphaeromidae.[103]—These are flattened, broad-bodied
forms, most commonly met with in the Mediterranean and warmer
seas. Without being actually parasitic, they are frequently found as
scavengers in decaying material, and they show some relationship to
the parasitic Cymothoidae. In some of the genera, e.g. Cymodoce, the
ovigerous female shows a degenerate condition of the mouth-parts,
while the maxillipedes undergo an enlargement, and are used for
causing a current through the brood-chamber.

Sub-Order 3. Valvifera.

The Valvifera, illustrated by the Idotheidae and Arcturidae, are

characterised by the uropods being turned back and expanded to
form folding doors covering up the delicate pleopods, which are
mostly respiratory in function, though the anterior pairs may serve
as swimming organs. Arcturus is a typically deep sea genus, many
species, remarkably furnished
with spiny processes, having been
taken by the Challenger in the
southern hemisphere. The
Idotheidae are more littoral
forms, several species of Idothea
being commonly met with off the
British coasts, occasionally
penetrating into brackish or even
fresh water.

Fig. 83.—Munnopsis typica

(Munnopsidae), ♂ , × 2. A, 2nd
antenna; Ab, abdomen; T, 5th thoracic
appendage or 4th leg. (After Sars.)

Sub-Order 4. Asellota.

In this group the abdominal segments are fused dorsally to form a

shield-like caudal region; the pleopods are respiratory in function
and reduced in numbers, the first pair being often expanded and
produced backwards to form an operculum covering the rest. Several
of the Asellota are fresh-water, Asellus aquaticus (Asellidae) being
extremely abundant all over Europe in weed-grown ditches, the mud
of slowly-moving streams, and even on the shores of large lakes.
They are mostly sluggish in habit, but the marine Munnopsidae
(Fig. 83, Munnopsis) are expert swimmers, the swimming organs
being fashioned by the expansion and elongation of the thoracic legs.

Sub-Order 5. Oniscoida.
 The Oniscoida[104] are terrestrial forms in which the abdomen is
fully segmented, the pleopods are respiratory, their endopodites
being delicate branchiae, while their exopodites are plate-like and
form protective opercula for the gills, and the uropods are biramous
and not expanded. The epimera of the segments are greatly
produced. The terrestrial Isopods, although air-breathers,[105] are
dependent on moisture, and are only found in damp situations. It
seems probable that they have been derived from marine Isopods,
since the more generalised of them, e.g., Ligia (Fig. 84), common on
the English coasts, are only found in damp caves and crannies in the
rocks.

Fig. 84.—Ligia oceanica, ventral and dorsal views, × 1. (From

original drawings prepared for Professor Weldon.)

The related Ligidium is found far inland, but always in the

neighbourhood of water. These two genera may be distinguished by
the numerous joints in the flagellum of the second antennae, the
flagellum being in all cases the portion of the antenna succeeding the
long fifth joint. Philoscia muscorum occurs usually near the coast,
but it is also found inland in England under trees in damp moss. This
genus and the common Oniscus, found in woods, are distinguished
by the presence of three joints in the flagellum of the second
antenna. Philoscia can be distinguished from Oniscus by its
narrower body and the pretty marbled appearance of its back. The
genus Trichoniscus has four joints in the flagellum; various species
are found in woods. In Porcellio and Armadillidium there are only
two joints in the flagellum, while Armadillidium, the common
garden wood-louse, can be distinguished from all others by the
flattened shape of the uropods, and the habit of rolling up into a ball
like an Armadillo.
There is also a very peculiar species, Platyarthrus hoffmannseggii,
which occurs in England and Northern Europe, and always lives in
ants’ nests. It is supposed that they serve as scavengers for the ants,
which tend them carefully, and evidently treat them as domestic
animals of some kind. The small creature is quite white and blind,
and has exceedingly short antennae.

Sub-Order 6. Epicarida.

The Epicarida include an immense number of Isopods, parasitic

upon other Crustacea. In the adult state they become greatly
deformed, and offer very few characters of classificatory value, but
they all pass through certain highly characteristic larval stages which
are essentially similar in the different families. All the species are
protandric hermaphrodites, each individual being male while in a
larval state, and then losing its male organisation and becoming
female as the parasitic habit is assumed.
Two series of families are recognised according to the larval stages
passed through, the Cryptoniscina, in which the adult male
organisation is assumed in the Cryptoniscus stage, and the female
condition is imposed directly upon this form, and the Bopyrina, in
which the Cryptoniscus passes into a further larval stage, the
Bopyrus, which performs the function of the male, and upon which
the female organisation is imposed as the parasitic habit is assumed.
The following is a list of the Epicarida with the Crustacea which
serve as their hosts[106]:—
 Microniscidae on Copepoda.
Cryptoniscidae on Ostracoda.
Liriopsidae on Rhizocephala.
Cryptoniscina Hemioniscidae on Cirripedia.
Cabiropsidae on Isopoda.
Podasconidae on Amphipoda.
Asconiscidae on Schizopoda.

Dajidae
Phryxidae
Bopyrina on Decapoda
Bopyridae
Entoniscidae
In all cases the first larval form
which hatches out from the
maternal brood-pouch is called
the Epicaridian larva (Fig. 85).
This little larva has two pairs of
antennae, a pair of curious frontal
processes, and a pair of
mandibles. The other mouth-
parts are missing; there are only
six thoracic limbs, but the full
complement of six biramous
pleopods are present, and at the
end of the body there may be a
long tube of unknown function.
Fig. 85.—Epicaridian larva, probably As a type of the
belonging to one of the Cryptoniscina. Cryptoniscina we may take the
A, 2nd antenna; Ab, abdominal Liriopsidae,[107] parasitic on the
appendages; T, thoracic appendages. Rhizocephala, which are, of
(From Bonnier, after Hansen.)
course, themselves parasitic on
the Decapoda, the whole
association forming a very remarkable study in Carcinology.
Almost every species of the Rhizocephala is subject to the attacks
of Liriopsids, the latter fixing either on the Rhizocephala themselves,
or else on the Decapod host at a point near the fixation of the
Rhizocephalous parasite. An exceedingly common Liriopsid is
Danalia curvata, parasitic on Sacculina neglecta, which is itself
parasitic on the spider-crab, Inachus mauritanicus, at Naples. The
adult Danalia is a mere curved bag full of eggs or developing
embryos, and without any other recognisable organs except two pairs
of spermathecae upon the ventral surface where the spermatozoa
derived from the larval males are stored.
In Fig. 86 is represented a
female of Inachus mauritanicus
which carried upon it two
Sacculinae and a Danalia
curvata, and upon the latter are
seen two minute larval males in
the act of fertilising the adult
Danalia. The eggs develop into
the Epicaridian stage, after which
the larva passes into the
Cryptoniscus stage (Fig. 87). In Fig. 86.—Inachus mauritanicus, ♀, × 1,
this larval form the segments are carrying two Sacculina neglecta (a, b),
clearly delimited; the only mouth- and a Danalia curvata (c), the latter
bearing two dwarf males.
parts present are the mandibles,
but there are seven pairs of
thoracic limbs and the full number of pleopods. This Cryptoniscus
stage is found in all the Epicarida, and only differs in detail in the
various families.
In the Cryptoniscina the Cryptoniscus larva is the male, and at this
stage possesses a pair of large testes in the thorax. The ovaries are
also present at this stage as very small bodies applied to the anterior
ends of the testes. The larval males in this state seek out adult fixed
Danaliae and fertilise them; and, when this is accomplished, they
themselves become fixed to the host and begin to develop into the
adult female condition. The limbs are all lost, and out of the mouth
grows a long proboscis (Fig. 88, P), which penetrates the tissues of
the host. The ovaries begin to grow, and a remarkable process of
absorption in the testes takes place. These organs, when fixation
occurs, are never empty of spermatozoa, and are frequently crammed
with them. After fixation some large cells at the interior borders of
the testes begin to feed upon the remains of these organs and to grow
enormously in size and to multiply by amitosis. These phagocytes, as
they really are, attain an enormous size, but they are doomed to
 degeneration, the chromatin
becoming dispersed through the
cytoplasm, and the nuclei
dividing first by amitosis and
then breaking up and
disappearing. As the parasite
grows, the heart at the posterior
end of the body ceases to beat;
the ovaries increase enormously
at the expense of the alimentary
canal, and on the ventral surface
two pairs of spermathecae are
invaginated ready to receive the
spermatozoa of a larval male. In
the adult condition, after
fertilisation has taken place and
the ovaries occupy almost the
whole of the body, the remains of
the phagocytic cells can be seen
on the dorsal surface in a
degenerate state. They evidently
are not used as food, and their
sole function is to make away
with the male organisation when
it has become useless.[108]
In the series Bopyrina, after
Fig. 87.—Ventral view of Cryptoniscus the free-living Epicaridian and
larva of Danalia curvata, ♂, × 25. Cryptoniscus stages, a further
larval state is assumed, called the
Bopyrus, which is the functional
male, and, after performing this function, passes on to the adult
female condition.
The family Bopyridae is parasitic in the branchial chamber of
Decapoda, especially Macrura and Anomura. When one of these
Decapods is infested with an adult Bopyrid the gill-chamber in which
it is situated is greatly swollen, as shown in Fig. 90. A very common
Bopyrid is Bopyrus fougerouxi, parasitic in the gill-chambers of
Palaemon serratus. The Bopyrus larva or functional male has the
 appearance shown in Fig. 91. It
differs from the Cryptoniscus
stage in possessing a rudimentary
pair of anterior thoracic limbs
and seven pairs normally
developed, while the abdominal
limbs are plate-like and branchial
in function. The male can often be
found attached to the female
beneath the last pair of
incubatory lamellae.
Fig. 88.—Side view of Danalia curvata, The adult female condition,
× 15, shortly after fixation and loss of which is assumed after the
larval appendages. A, Alimentary canal; Bopyrid stage is passed through,
E, eye; H, heart; N, phagocytic cells; O,
ovary; P, proboscis.
is illustrated in Fig. 92. The body
acquires a remarkable
asymmetry, due to the unequal
pressure exerted by the walls of
the gill-chamber. The antennae
and mandibles (Fig. 92, B) are
entirely covered up by the largely
expanded maxillipedes; maxillae
are, as usual, entirely absent.
Very large lamellae grow out from
the bases of the thoracic limbs to
form a brood-pouch, and in this
manner the adult condition is
attained.
The final complication in the
life-histories of these Isopoda is
reached by the family
Entoniscidae, which are
parasitic when adult inside the Fig. 89.—Optical section (dorsal view)
thoracic cavity of Brachyura and of Danalia curvata, in the same stage
Paguridae. The cephalothorax of as Fig. 88. A, Alimentary canal; Ec,
ectoderm; H, heart; N, phagocytic cells;
a Carcinus maenas, which O, ovaries; P, proboscis.
contains an adult Portunion
maenadis (P), is shown in Fig. 93.
 The parasite is of a reddish colour
when alive.

Fig. 90.—Galathea intermedia, with a

Pleurocrypta microbranchiata under
its left branchiostegite (B), × 1. (After
Sars.)
Fig. 91.—Ventral view of male Bopyrus
fougerouxi, × 30. A, 1st and 2nd
antennae; T, 8th (last) thoracic
appendage. (After Bonnier.)
 Fig. 92.—Bopyrus fougerouxi. A, Ventral
view of female carrying a male (M)
between her abdominal appendages, × 8;
B, ventral view of part of head of female,
the maxillipedes and the left mandible
having been removed. A.1, A.2, 1st and
2nd antennae; M, male; Mn, right
mandible; Mx, left maxillipede; O,
oostegite; T, left 4th thoracic appendage
or 3rd leg. (After Bonnier.)

The Entoniscidae pass through a free living Epicaridian and

Cryptoniscus stage, and become adult males in the Bopyrus stage. It
is stated, however, by Giard and Bonnier[109] that these individuals,
which actually function as males, never grow up into adult females,
though all the adult females have
passed through a male stage in
which the male genital ducts are
not formed. The
hermaphroditism, therefore, in
these animals at any rate is
absolutely useless from a
reproductive point of view, and
this justifies our looking for some
other explanation of it, such as
was suggested on p. 105.

Fig. 93.—Cephalothorax of Carcinus

maenas, seen from the ventral side,
containing a parasitic Portunion
maenadis (P), × ½. (After Bonnier.)

Fig. 94.—Portunion maenadis, ♀:—A, Young, × 10; B, older, × 5;

C, adult, before the eggs are laid, × 3. A, 2nd antenna; Ab,
abdomen; B, anterior lobe of brood-pouch; B′, its lateral lobe; H,
head; 1, 2, 1st and 2nd incubatory lamellae (oostegites). (After
Giard and Bonnier.)
 The Bopyrus fixes in the gill-chamber of the host and becomes
converted into the adult female by a series of transformations. As
these changes take place it invaginates the wall of the gill-chamber
and pushes its way into the thoracic cavity of the crab, though it lies
all the time enveloped in the invaginated wall of the gill-chamber,
and not free in the body-cavity of the crab. The transformations
which it undergoes are shown in Fig. 94. The body first assumes a
grub-like appearance (A), and two pairs of incubatory lamellae (1, 2)
grow out from the first and second thoracic segments. In the next
stage (B) these lamellae assume gigantic proportions, and four pairs
of branchiae grow out from the abdominal segments (Ab). In the
final stage (C) the incubatory lamellae have further increased in size,
and constitute the main bulk of the body; the enormous mass of eggs
is passed into the incubatory pouch, and all that remains of the rest
of the body is the small head (H) and the abdomen (Ab), furnished
with its branchiae. Communication with the external world is kept
up through an aperture which leads from the brood-pouch into the
gill-chamber of the host, and through this aperture the young are
hatched out when they are developed sufficiently.
The presence of these parasites, although they are never in actual
contact with the internal organs of the crab, calls forth the same
phenomenon of parasitic castration as was observed in the
Rhizocephala. A remarkable association is also found to exist
between the Entoniscidae and Rhizocephala, of such a kind that, on
the whole, a crab infested with a Rhizocephalan is more likely to
harbour an Entoniscid than one without. The explanation of this
association is probably that a crab with a Sacculina inside it is
prevented from moulting as often as an uninfected crab, and, in
consequence, the larval stages of the Entoniscid in the crab’s gill-
chamber are more safely passed through.

Sub-Order 7. Phreatoicidea.[110]

The members of this sub-order, although agreeing with the

Isopoda in the essentials of their anatomy, resemble the Amphipoda
in being rather laterally compressed, and in having the hand of the
first free thoracic limb enlarged and subchelate. The abdomen is
greatly produced laterally by expansions of the segments. In fact, the
shape of the body and of the limbs is very Amphipodan.—
Phreatoicus from New Zealand, Southern Australia, and Tasmania.
Phreatoicopsis,[111] a very large form from Gippsland, Victoria. Only
one family exists, Phreatoicidae.

Order IV. Amphipoda.

In this order the body is flattened laterally, the heart is anterior in

position, and the branchial organs are attached to the thoracic limbs.
There are three well defined sub-orders, (i.) the Crevettina,
including a vast assemblage of very similar animals, of which the
common Gammarus and Orchestia may serve as examples; (ii.) the
Laemodipoda or Caprellids, and (iii.) the Hyperina.
We cannot do more than touch on the organisation of these sub-
orders.

Sub-Order 1. Crevettina.

In this sub-order only one thoracic segment is fused with the head;
the basal joints of the thoracic limbs are expanded to form broad
lateral plates, and the abdomen is well developed, with six pairs of
pleopods, the last three pairs being always turned backwards, and
stiffened to act as uropods.
This group has numerous fresh-water representatives, e.g.
Gammarus of several species, the blind well-shrimp Niphargus, and
the S. American Hyalella; but the vast majority of the species are
marine, and are found especially in the littoral zone wherever the
rocks are covered with a rich growth of algae, Polyzoa, etc. The
Talitridae or “Sand-hoppers” have deserted the waters and live
entirely in the sand and under rocks on the shore, and one common
European species, Orchestia gammarellus, penetrates far inland,
and may be found in gardens where the soil is moist many miles
from the sea.
The Rev. T. R. R. Stebbing, in his standard work[112] on this group,
recognises forty-one families, and more than 1000 species, so that