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Capitalism’s Sexual History
ii

Oxford Studies in Gender and International Relations


Series editors: J. Ann Tickner, American University,
and Laura Sjoberg, University of Florida and Royal Holloway,
University of London
Windows of Opportunity: How Women Seize Rewriting the Victim: Dramatization as Research
Peace Negotiations for Political Change in Thailand’s Anti-Trafficking Movement
Miriam J. Anderson Erin M. Kamler
Women as Foreign Policy Leaders: National Equal Opportunity Peacekeeping: Women, Peace,
Security and Gender Politics in Superpower and Security in Post-Conflict States
America Sabrina Karim and Kyle Beardsley
Sylvia Bashevkin Gender, Sex, and the Postnational
Gendered Citizenship: Understanding Gendered Defense: Militarism and Peacekeeping
Violence in Democratic India Annica Kronsell
Natasha Behl The Beauty Trade: Youth, Gender, and Fashion
Gender, Religion, Extremism: Finding Women in Globalization
Anti-Radicalization Angela B. V. McCracken
Katherine E. Brown Global Norms and Local Action: The Campaigns
Enlisting Masculinity: The Construction of Gender against Gender-Based Violence in Africa
in U.S. Military Recruiting Advertising During the Peace A. Medie
All-Volunteer Force Rape Loot Pillage: The Political Economy of Sexual
Melissa T. Brown Violence in Armed Conflict
The Politics of Gender Justice at the International Sara Meger
Criminal Court: Legacies and Legitimacy From Global to Grassroots: The European Union,
Louise Chappell Transnational Advocacy, and Combating Violence
Cosmopolitan Sex Workers: Women and Migration against Women
in a Global City Celeste Montoya
Christine B. N. Chin Who Is Worthy of Protection? Gender-Based
Intelligent Compassion: Feminist Critical Asylum and US Immigration Politics
Methodology in the Women’s International Meghana Nayak
League for Peace and Freedom Revisiting Gendered States: Feminist Imaginings
Catia Cecilia Confortini of the State in International Relations
Complicit Sisters: Gender and Women’s Issues Swati Parashar, J. Ann Tickner, and
across North-South Divides Jacqui True
Sara de Jong Out of Time: The Queer Politics of Postcoloniality
Gender and Private Security in Global Politics Rahul Rao
Maya Eichler Gender, UN Peacebuilding, and the Politics of
This American Moment: A Feminist Christian Space: Locating Legitimacy
Realist Intervention Laura J. Shepherd
Caron E. Gentry A Feminist Voyage through International
Troubling Motherhood: Maternality in Global Relations
Politics J. Ann Tickner
Lucy B. Hall, Anna L. Weissman, The Political Economy of Violence against Women
and Laura J. Shepherd Jacqui True
Breaking the Binaries in Security Studies: A Queer International Relations: Sovereignty,
Gendered Analysis of Women in Combat Sexuality and the Will to Knowledge
Ayelet Harel-Shalev and Shir Daphna-Tekoah Cynthia Weber
Scandalous Economics: Gender and the Politics of Bodies of Violence: Theorizing Embodied Subjects
Financial Crises in International Relations
Aida A. Hozić and Jacqui True Lauren B. Wilcox
Capitalism’s Sexual
History
Nicola J. Smith

1
iv

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 2020

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
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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: Smith, Nicola Jo-Anne, author.
Title: Capitalism’s sexual history / Nicola J. Smith.
Description: New York, NY : Oxford University Press, [2020] |
Series: Oxford studies in gender and international relations |
Includes bibliographical references and index.
Identifiers: LCCN 2020012845 (print) | LCCN 2020012846 (ebook) |
ISBN 9780197530276 (hardback) | ISBN 9780197530290 (epub)
Subjects: LCSH: Prostitution—Great Britain—History. |
Sex—Great Britain—Economic aspects—History. |
Capitalism—Social aspects—Great Britain. | Queer theory. |
Feminist theory. Classification: LCC HQ185 .A5 S65 2020 (print) |
LCC HQ185 .A5 (ebook) | DDC 306. 740941—dc23
LC record available at https://lccn.loc.gov/2020012845
LC ebook record available at https://lccn.loc.gov/2020012846

9 8 7 6 5 4 3 2 1
Paperback printed by LSC Communications, United States of America
Hardback printed by Bridgeport National Bindery, Inc., United States of America
ACKNOWLEDGMENTS

The writing of this book has had a very long history, and I am indebted
to a great many people who have offered me help, advice, and encourage-
ment along the way. I would like to begin by thanking my colleagues in the
Department of Political Science and International Relations and the School
of Government at the University of Birmingham for their individual kind-
ness and collective generosity as well as for the various conversations we
have had about the research and writing process. I would also like to express
my gratitude to Juanita Elias, Penny Griffin, Sarah Kingston, Mary Laing,
Donna Lee, Milly Morris, Lucy Neville, Katy Pilcher, Adrienne Roberts,
Heather Savigny, Dani Tepe-Belfrage, Tiina Vaittinen, Julia Welland, and
Heather Widdows for their intellectual, practical, and moral support for
this project. Special thanks to Tendayi Bloom, Nicola Pratt, and Nick
Wheeler for reading and commenting on the draft materials, and to Lisa
Downing not only for her feedback on the manuscript but also for making
the writing of it a lot less lonely and a lot more enjoyable than it otherwise
would have been.
I owe a huge debt of thanks to Angela Chnapko for making this book
possible, for her enthusiasm for and guidance on the project, and for
leading me through the publication process—it has been such a wonderful
experience to work with her and her colleagues at Oxford University Press.
I am also beholden to J. Ann Tickner and Laura Sjoberg as series editors
for all their help in bringing this book to life, as well as the two anonymous
readers for their invaluable feedback on the manuscript—I hope I have
done justice to their very careful and constructive comments but any errors
and omissions in the final text are of course mine alone.
Sections of text in the Introduction, Chapter 1, and Chapter 5 of this book
draw from earlier chapters published elsewhere. Many thanks to Oxford
University Press and Edward Elgar for granting me permission to repro-
duce material from the following publications: “Toward a Queer Political
vi

Economy of Crisis” by Nicola Smith, in Scandalous Economics: Gender


and the Politics of Financial Crisis, edited by Jacqui True and Aida Hozić
(New York: Oxford University Press, 2016), 231–247; and “Queer Theory
and Feminist Political Economy” by Nicola Smith, in Handbook on the
International Political Economy of Gender, edited by Juanita Elias and
Adrienne Roberts (Cheltenham, UK: Edward Elgar, 2018), 102–112.
Last but definitely not least, I would like to thank my family and
friends—including my brother, Kieron Smith, my beloved friend Cathy
Newman, and “soul restorers” Sophie Khan and Manita Rajgor—for lis-
tening to me, advising me, and cheering me on. I reserve particular thanks
for my lifelong friend Emma Cooper, who has shared the journey of this
book with me from start to finish, and who gives me immeasurable sup-
port for my writing and everything else besides. Above all, I would like to
thank my mum and dad, Lesley and Greg Smith—who are there for me
every day in every way and without whom I could never have completed
this project—and my son, Tom, of whom I could not be more proud.

[ viii ] Acknowledgments
Introduction

W hat does capitalism have to do with sexuality? At one level, the re-
lationship between economic and sexual practices could scarcely be
more contested. When Amnesty International voted to support the de-
criminalization of consensual commercial sex in August 2015, for example,
it did so in response to longstanding activism by sex workers globally. Yet
its proposal was denounced as “unfathomable” in an open letter signed by
hundreds of prominent figures from every region of the world.1 Otherwise
disparate voices from politicians to psychiatrists, religious sisters to
truckers, and university professors to Hollywood stars were united in their
condemnation of the notion that sex could in any way be understood as an
economic activity.
At another level, though, the connections between economic and
sexual life are often rendered invisible. Sexuality—especially but not only
for women—is understood as intimate, sacred, the embodiment of love.
That sexuality exists somewhere beyond economy is believed to be the
natural order of things—an order that, for the most part, can go unno-
ticed but, when transgressed, must be fiercely defended. What Amnesty
International’s decision did was to confront deep-rooted assumptions that
sexuality is inherently separable from capitalism, and that it can be taken
as read that “the inside of a woman’s body is not a workplace.”2
The letter’s use of the term “unfathomable” is telling, for it reveals how
the intended effect of the moral outcry was not just to influence Amnesty
International’s policy position but also to set strict limits on how sexuality
can and cannot be conceptualized. For sex to become work, and so for the
sexual to become economic, was beyond the pale, beyond morality, beyond

Capitalism’s Sexual History. Nicola J. Smith, Oxford University Press (2020). © Oxford University Press.
DOI: 10.1093/oso/9780197530276.001.0001.
2

comprehension. Yet, as this book explores, it is this very move—a move that
removes sexuality from political economy through appeals to morality—
that is, and has long been, extremely convenient for capitalism. It means
that certain tricky questions can be taken off the table, and especially those
regarding the close alignments between normative sexuality and prevailing
economic logics. If the inside of a woman’s body can never be thought of as
a workplace, for instance, then it is easy to conclude that the labor of child-
birth is not really “labor” at all. Similarly, if we protest against paid sex on
the grounds that sexual exploitation for profit can never be tolerated, then
the possibility that capitalism routinely appropriates unpaid sexual labor
for profit can simply slide from view.
The overarching purpose of this book is to challenge these assumptions.
More specifically, it aims to excavate and contest how sexuality has come to
be regarded not only as distinct from political economy but as antithetical
to it. Such an agenda is especially significant given that the field of inquiry
devoted to the study of global capitalism, international political economy
(IPE), is itself structured by the economy/sexuality dichotomy. As decades
of feminist critique have shown, IPE’s disciplinary legacies are not ones of
embodiment, intimacy, and desire.3 Historically, orthodox IPE was founded
on the twin pillars of states and markets—two imaginary spheres that
were understood to touch and, at times, collide but that remained clearly
separable both from each other and from the mess and matter of everyday
life.4 Critical IPE has done much to tackle these dualisms and abstractions,
exploring instead how states and markets are socially embedded and how
it is in everyday life that the oppressions and struggles of global capi-
talism play out.5 Yet critical IPE, too, remains in some important respects
disembodied—both in rendering invisible actually existing human beings
and in neglecting embodied social hierarchies such as gender, race, and sex-
uality.6 Certainly, work on sexuality exists, but it is nevertheless regarded
as marginal at best, and irrelevant at worst, in the vast majority of IPE
scholarship.
Conversely, the prevailing field in sexuality studies, queer theory, has
often come uncoupled from debates about global capitalism. As Judith
Butler notes, queer theory is frequently consigned to the realm of the
“merely cultural”: as only interested in, and applicable to, “matters of cul-
tural recognition” rather than those of political economy.7 This move enables
sexuality to be presented as both non-material and immaterial (i.e., as nei-
ther bound up with, nor relevant to the study of, economic structures).
Although Butler argues against this depiction of queer theory, contending
instead that it potentially has a great deal to say about material (in)justice,
it is also the case that many queer scholars have overlooked—and many

[2] Capitalism’s Sexual History


continue to overlook—exactly these kinds of questions. This is not least
because queer theory has had a rather tetchy relationship with Marxism,
with the former accusing the latter of erasing sexuality and the latter
reproaching the former for ignoring materiality, and this has all too often
played out as a series of dogged battles over meta-theory, the nature of so-
cial “reality,” and the so-called material-discursive divide.8
Thus, rather than challenging the division between the economic and
the sexual, both IPE and queer theory have often reinforced the sense that
this division is somehow natural and neutral as opposed to contingent and
contestable. This lack of engagement between the two fields matters very
much, for it nurtures the illusion that the deployment of sexuality is anom-
alous, not endemic, to capitalism, and that it is on the outside of capitalism
that the inside of women’s bodies can naturally be found.
This is, of course, not to claim that there are no IPE scholars who engage
with sexuality or no queer scholars who are interested in capitalism, for nei-
ther of these things are true.9 Yet, despite the empirical presence of queer
theory in political economy, and vice versa, sexuality is often placed on the
constitutive outside of IPE. This is a function of disciplinary histories, for
IPE was founded on a series of binaries and oppositions that continue to
inform contemporary debates. These include the dualisms of states and
markets, politics and economics, and national and international—all of
which are treated as endogenous to IPE—as well as the boundaries that
constitute the inside and outside of the field itself. Particularly important
has been the separation between the public sphere (e.g., of government
and work) and the private sphere (e.g., of household and family)—a move
that has allowed intimate and domestic life to be bracketed off and treated
as external to the discipline.10 Associated with private/personal space in
this way, sexuality has not even been written out of IPE since it was never
written in to start with. The picture for queer theory is slightly different,
for a major impetus has been to dismantle conceptual binaries and disci-
plinary boundaries rather than to maintain them. However, queer theory’s
own heritages (including its strong tradition in literary and cultural criti-
cism) mean that it, too, has frequently replicated the economy/sexuality
dichotomy. Thus, although intersections between queer theory and polit-
ical economy do exist, the respective fields could nevertheless gain a great
deal from closer dialogue with each other—and a central task of this book
is to foster exactly this kind of engagement.11
This agenda is particularly warranted given that the economy/sexuality
dichotomy does not just play out in disciplinary practice but, as the book
examines, it is woven into the fabric of capitalism itself. Feminist scholars
have illuminated how public space has been historically constituted (and

IntroductIon [3]
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privileged) as the “masculine” sphere of economic production, whereas


private space has been constructed (and devalued) in terms of the “fem-
inine” realm of social reproduction.12 As Silvia Federici writes, capitalism
has needed the creation, and concealment, of a sexual division of labor
that is “above all a power-relation, a division within the workforce, while
being an immense boost to capitalist accumulation.”13 This book focuses
in on the sexual element of this equation by exploring how capitalist de-
velopment has been sexual in the dual sense, for it has relied not only on
the sexual division of labor but also on the cultivation and suppression of
sexuality. Indeed, as the book interrogates, sexuality has played a crucial
role in demarcating the public from the private, the productive from the
reproductive, and the economic from the intimate. Thus, while this volume
aims to bring queer theory more into conversation with political economy,
it does so by drawing on and contributing toward the long-established tra-
dition of feminist political economy—a project that is sometimes sidelined
in IPE and largely ignored in queer theory, and without which no story of
capitalism’s sexual relations could be told.
In order to investigate these themes, the book takes up commercial sex
as a site of particular relevance to the study of capitalism and sexuality.
Indeed, there is arguably no other case in which the economy/sexuality di-
chotomy is both more apparent and more contested, even if debates are
not explicitly articulated as such. Although sex work has received scant at-
tention in queer theory,14 it has attracted considerable interest in feminist
scholarship and activism. For some, commercial sex is synonymous with
sexual exploitation because it entails the sale and purchase of women’s
bodies and, hence, of their sexual selfhood. It is for this reason that aboli-
tionist feminists call for the sex industry to be tolerated no longer, either
legally or culturally.15 For others, commercial sex can constitute a terrain of
agency, subversion, and even radical politics. Some sex-positive feminists,
for instance, have argued that sex work represents not a violation of bodily
integrity but a reflection of an individual’s property in their own body.16
Such accounts are underpinned by the economy/sexuality dichotomy in
that commercial sex is understood in terms of oppression or liberation
that is first and foremost sexual rather than economic in nature.17 Yet the
emphasis on the sexual element of commercial sex has also been disputed,
not least by sex workers themselves in demanding that their sexual labor
be accepted as legitimate work, thereby rejecting conventional separations
between sex and work and, hence, between sexuality and economy.18
This book strongly supports calls by sex workers to recognize and de-
criminalize their sexual labor but, in so doing, it follows the lead of Juno
Mac and Molly Smith in refusing to “prioritize discussion on whether the

[4] Capitalism’s Sexual History


sex industry, or even sex itself, is intrinsically good or bad.”19 Nor does it
engage in philosophical or political deliberation about the essential “truth”
of sex or sex work. As intuitively appealing as such debates are, their strong
tendency is to divert attention away from the material conditions under
which sex workers labor and so to produce distracting and polarizing
effects that, ultimately, capitalism thrives on. As Michel Foucault brings to
light, all of the “garrulous attention which has us in a stew over sexuality,”
and especially the quest for its underlying truth, can be understood as an
apparatus of modern power.20 Indeed, it is precisely this “garrulous atten-
tion” that the book seeks to interrogate, in particular by considering how
the politicization of sexuality as a moral issue both requires and enables its
depoliticization as an economic one. While it was a particularly high-profile
example that garnered widespread interest internationally, the furor over
Amnesty International’s support for the decriminalization of commercial
sex was underpropped by moralizing discourses that were neither distinc-
tive nor new. Indeed, as the book will explore, the sex/work split has been
constituted as much through the proliferation of discourses of sex work
as it has been through the censoring of sex work, with these apparently
incongruous energies working together to foster the illusion that sex work
is fundamentally different from other intimate and economic relations. As
we shall see, capitalism needs and allows sex work to be constructed as an
aberration in order to naturalize the appropriation of unpaid sexual labor
that takes place primarily, if not exclusively, via the institution of marriage.
The case of sex work is thus much more than just an illustrative example
of the economy/sexuality dichotomy in motion, for it is instead a central
mechanism through which this dichotomy has been forged.
It is vital to stress, therefore, that the economy/sexuality dichotomy
is not a simple one, for capitalism does not either deliver sexuality to the
market or insist on a distance between them. Rather, the argument of this
book is that this dichotomy obscures how the two are in fact entwined.
Commercial sex may be condemned and often criminalized, but it also
represents the logical extension of capitalist social relations in which all
aspects of life are increasingly defined in terms of market rationalities.
Accordingly, the book is less interested in the identification of a single sys-
temic logic and more with the possibility of frictions, incongruities, and
what Foucault calls “unstable mixtures.”21 Indeed, it is because of these
deep ambiguities that sex work has become an object of such intense moral
focus, for it has come to be constructed as neither “real” sex nor “real” work
even as debates insist that it is either “really” sex or “really” work. Thus, the
book seeks to overcome some of the dualisms that exist within feminism
itself regarding the relationship between the economic and the sexual,

IntroductIon [5]
6

and that continue to polarize debates about commercial sex. Rather than
contending that the economic and sexual realms either comfortably coexist
or must not adulterate each other, the book sets out to uncover and under-
stand how sexuality is positioned within capitalism while simultaneously
being positioned against it.
In so doing, the volume contributes both to IPE scholarship on capi-
talism and to queer and feminist scholarship on sexuality, first, by devel-
oping and advancing the still-burgeoning field of queer political economy;
second, by demonstrating how capitalist social relations are always already
sexual relations; and, third, by exploring how economy and sexuality have
come to be constructed as not only different but opposite realms. As part
of this agenda, the book also speaks directly to the field of sex work studies
by placing sex work at the crux of its analysis. While there is a dearth of
literature on queer theory and commercial sex, some scholars have begun
to rectify this by asking, for instance, what it means to “be queer in” sex
work in order to center non-normative sexualities in the contemporary sex
industry, and by considering what it means to “do queer to” sex work in
order to examine the interconnections between commercial sex and heter-
onormative power relations.22 Yet queer accounts of sex work have, on the
whole, treated commercial sex as a “merely cultural” issue rather than as
a matter for the theory and practice of political economy.23 The book thus
aims to move this literature forward by highlighting the need for critical
scholarship not only to think queerly about sex work but also to situate
this within wider critiques of capitalism—including through mutually ben-
eficial engagement with feminist political economy.24 By investigating the
constitution and effects of economy/sexuality and sex/work as interrelated
dualisms, the book opens up new space for critical inquiry into the
intersections, tensions, contradictions, and connections that characterize
capitalism’s relationship with sexuality.

HISTORICIZING CAPITALISM AND SEXUALITY

For the reasons outlined, capitalism’s sexual relations are, above all, a ques-
tion of power, and so this book is, above all, a political book. It is important
to be upfront, therefore, that this study is not situated within the disci-
pline of history, nor is it a history book. Nevertheless, the guiding meth-
odology for the research is historical and, more precisely, genealogical in
approach. Genealogy is not conventionally “political” in the sense that it
does not routinely form part of the repertoire of those fields to claim polit-
ical analysis as their domain (political science, international relations, and

[6] Capitalism’s Sexual History


IPE).25 Yet, as Laura Jenkins compellingly argues, genealogy is, at heart, a
politicizing endeavor that can be used to slacken, subvert, and sabotage
perspectives and power relations that are otherwise taken for granted.26
It is, then, a self-consciously political method of inquiry, even if it is not
always acknowledged to be so.
In order to describe and reflect on the research undertaken for this book,
it is helpful to set out what genealogy does and does not entail as a meth-
odology. Specifically, genealogy is a mode of critique that charts the emer-
gence of contemporary power relations to expose how they do not (and
never did) rest on stable foundations but instead are (and always were)
contingent. As such, genealogy is neither the history of politics nor the pol-
itics of history but instead seeks to historicize in order to politicize—that
is, “to locate the historical conditions that allow us to think, speak, and act
as we do now.”27 Most closely associated with Foucault, genealogical studies
are “gray, meticulous, and patiently documentary”28 but they are not “pos-
itivistic returns to a form of science that is more attentive and more accu-
rate.”29 On the contrary, claims to objective truths—including appeals to
scientific legitimacy—are exactly what genealogy sets out to investigate.
Consequently, genealogy is not fastened to a specific set of procedures or
techniques but is best understood as characterizing those forms of inquiry
that use the past to critique the contemporary moment (“the history of the
present”30).
In practicing historicization, genealogy resists rather than pursues the
kind of all-embracing historical project where the identification of “root
or singular causes”31 is the stated or unstated goal. Accordingly, this book
does not attempt to lay bare the causal processes through which history
unfolds as a “unified developing totality”32 or to offer a “linear narrative
that reveals our progressive drive towards enlightenment.”33 Nor do I ap-
proach either capitalism or sexuality as a “pregiven, material reality”34
that determines or causes the other. Instead, I am interested in the messy
entanglements through which capitalism and sexuality constitute each
other, and I do so by offering a history that is necessarily partial and spe-
cific. A key limitation of the research, therefore, is that it does not (intend
to) provide the history of capitalism, sexuality, or sex work writ large. Here
I draw inspiration from other genealogical studies such as Jemima Repo’s
The Biopolitics of Gender, which offers a highly specific account of the “birth”
of gender identity in the clinic in postwar America rather than a history of
gender in general.35 Just as the titles of Foucault’s The History of Sexuality
and Repo’s The Biopolitics of Gender rather teasingly imply all-encompassing
histories and then go on to dismantle exactly this kind of agenda, the title
of this book, Capitalism’s Sexual History, is also meant in this spirit.36 To

IntroductIon [7]
8

openly acknowledge the incomplete and bounded nature of the research


is not to abandon commitments to scholarly rigor but, quite the opposite,
is to recognize—as both queer and feminist scholars urge us to do37— that
being partial (in both senses of the word) is the “precondition of a politi-
cally engaged critique.”38 To embrace specificity can be an act of politiciza-
tion, since it exposes not only how there is never just one perspective but
also how the pretense of singularity is “always intensely political.”39
The book therefore does not try to paint a general picture of capitalist
development but instead chronicles a particular (and, perhaps, peculiar)
sexual history: that of the British. I should immediately note that, in
choosing a major European nation as the primary site of my study, I am
keenly aware of the acute danger of further buttressing the systematic eu-
rocentrism that bedevils both IPE and queer theory. It is for this reason
that the feminist literature on global sexual economies has rightly concen-
trated on sex workers in and from the Global South so as to challenge the
eurocentrism of Western colonial imaginaries.40 Comparatively little atten-
tion has been devoted to the expansion and diversification of the sex trade
in the postindustrial economies of the Global North,41 and understand-
ably so. Yet, at times, scholarship on gender and sexuality may reinforce
such imaginaries even as it works to disrupt them.42 The literature on sex
tourism and sex trafficking, for instance, can sometimes construct women
in the Global South as “people of the body,”43 which perpetuates the mind/
body dualism on which the “fantasy”44 of white Western rationality is built.
Seen in this light, it becomes important not only to recognize the agency
of sex workers in and from postcolonial contexts45 but also to critique the
discourses that map sexual economies onto particular terrains (and bodies)
and not others. Indeed, as the book will consider at various junctures, anti-
immigration agendas are positively reliant on such discourses. Britain is
a prime example of this, for the othering of migrant workers is caught up
with the “fetishization”46 of migrant women’s bodies and this has, in turn,
proved highly expedient for capitalism.
A further limitation of the book is that it tells the story of Britain through
that of England specifically; it does so because, although “Britishness is not
one thing and has never been one thing,”47 it has nevertheless been his-
torically constituted through conflations with both Englishness and white-
ness.48 There is some controversy as to whether England was the birthplace
of capitalism: while some claim that “without English capitalism there
would probably have been no capitalist system of any kind,”49 others justi-
fiably contest the eurocentrism of such accounts by tracing the ancestries
of Western modernity (the supposed “European miracle”) to non-European
societies.50 Yet there can be little doubt that England performed a leading

[8] Capitalism’s Sexual History


part in establishing capitalism’s dominance globally, not only because of
the Industrial Revolution but also because of the sheer enormity of the
British Empire (such that, at its height, the sun was said never to set upon
it). The tale of England’s political, economic, and cultural ascendancy and
subsequent decline has been told any number of times before,51 and so
I will not re-rehearse that here. Instead, the book focuses on the very par-
ticular story of sexuality as one that, as discussed, has attracted a great
deal of interest in queer theory but has largely been discounted in IPE, in-
cluding vis-à-vis the British case.52 By no stretch of the imagination is the
economy/sexuality dichotomy quintessentially British—as illustrated by
the open letter to Amnesty International mentioned earlier—but Britain’s
history is nevertheless implicated directly in its emergence, and it is this
story that the book seeks to recount.
In keeping with the genealogical orientation of the research, I offer a
“history of the discourses”53 of sexuality in Britain to render visible how—
as much as the economy/sexuality dichotomy might appear to be a natural
fact and so beyond dispute—it has instead been carved out of history. It
is worth reiterating that the research does not proceed from the positivist
premise that the purpose and proper practice of social inquiry is to discover
and record the nature of social reality by collecting data on, and reporting
findings about, that nature (“The truth about sex work in Britain is . . . ”).54
Instead, I conceive of reality as constituted in and through discourses as
“practices that systematically form the objects of which they speak.”55 I un-
derstand discourses not as neutral descriptors of exogenous forces that
lie underneath social relations (e.g., objective interests) but as mutually
constitutive of the power relations that govern what can and cannot be
thought, said, and done in particular social contexts.56 I therefore share
Laura Shepherd’s view that there is “nothing more fundamental to poli-
tics than the construction of the conceptual apparatus that structures
knowledge in any given society”; indeed, by analyzing systems of truth and
knowledge production, “we are essentially examining the production of
possibility.”57
The analysis that follows draws from extensive original research into a
wide variety of primary sources, including policy, legal, medical, psychi-
atric, religious, literary, philosophical, and journalistic texts, as well as
theoretical and historical studies by queer, feminist, and political economy
scholars.58 The primary sources were curated through a three-year process
of in-depth archival research into multiple online repositories such as the
National Archives, Hansard, and the British Newspaper Archive. I chose
to consult a diverse range of sources (rather than, say, policy documents
alone) as this approach was most consistent with my genealogical interest

IntroductIon [9]
01

in the “massive, circulating, and discontinuous forms”59 that meaning


production can take. Indeed, I was surprised and fascinated by the sheer
eccentricity of constructions of sex work that would, at times, appear in
discursive sites that might otherwise seem completely unrelated. Yet, if
discourses comprise “a series of discontinuous segments whose tactical
function is neither uniform nor stable,”60 then their articulation in diver-
gent sites can itself be read as evidence that they are internal rather than
incidental to the exercise of modern power under capitalism. Sadly, it is not
feasible to reproduce text from the vast bulk of the materials consulted,
so I have selected quotations that I believe best exemplify the multiform
ways in which commercial sex has been discursively constructed in Britain
over time. In quoting these documents, my intention is not to disregard
the politico-economic context in which such texts are situated but, on
the contrary, to bring together “text” and “context” through the analysis
of discourse.61 Thus, rather than conceiving of discourses in terms of in-
dividualistic intentionalism, I approach individuals, intentions, ideas, and
institutions as co-constitutive of the “wider discursive terrain”62 in which
they are situated.

THE BOOK AHEAD

At the start of this Introduction, I asked what capitalism has to do with


sexuality, and this is the central research question that animates the book
as a whole. The first chapter addresses this question theoretically by devel-
oping a queer political economy lens to conceptualize capitalism’s sexual
relations. Arguing that queer political economy is best approached as a
historicizing endeavor, the chapter combines insights from two histor-
ical projects that are seldom read alongside each other—Michel Foucault’s
The History of Sexuality63 and Silvia Federici’s Caliban and the Witch— to
offer a new framework for the study of sexuality in/and political economy.
By mapping the manifold and shifting ways in which sexuality has been
produced and repressed, we can unearth how capitalism profits not only
from the creation of the economy/sexuality dichotomy but also from its
mystification.
The remaining four chapters attend to these themes empirically through
analysis of the history of sex work in Britain, with the structure and argu-
ment of each chapter organized around the charting of multifarious his-
torical threads in line with my genealogical approach. Chapter 2 begins
this journey in medieval64 England by showing how the sexual division of
labor already rested on distinctions between legitimate and illegitimate

[ 10 ] Capitalism’s Sexual History


sex. Charting the rise of capitalism in early modern England, the chapter
explores how such distinctions were redefined by a new dichotomy—that
between sex and work—that made possible the reconfiguration of the
sexual division of labor into an instrument for capitalist appropriation.
Chapter 3 turns to the Victorian period to consider the making of commer-
cial sex into the “great social evil” of the modern age, and how the prolifer-
ation of discourses on sex work helped to naturalize the public/private split
that underpinned the sexual division of labor. Chapter 4 examines how
the economy/sexuality dichotomy was further normalized in the twentieth
century—even as the private sphere was increasingly governed by market
logics—and how sex work acted as the pivot on which this dichotomy
hinged. Chapter 5 considers the evolutions and effects of these discursive
legacies in the twenty-first century. It explores how the pathologization
and criminalization of commercial sex is intertwined with the ongoing pri-
vatization of social reproduction and, with it, the privatization of wealth
that continues apace under neoliberal capitalism.
The book concludes by reflecting on how queer, feminist, and leftist pol-
itics might oppose the economy/sexuality dichotomy in solidarity with the
sex workers’ movement, since sex workers’ struggles cannot be detached
from wider collective struggles against the extraction of unpaid sexual
labor and the denigration of feminized work. Indeed, in refusing to accept
that sex and work are innately antithetical, the sex workers’ movement
exposes how capitalism “makes use”65 of sexuality not as an aberration or
abomination but as a matter of course, just as it has done for centuries.
Before proceeding, I want to make a few final points. First, the book
hopes to be of use and interest to other scholars seeking to develop the
field of queer political economy but in no way does it claim or aspire to
represent “the” queer political economy perspective or approach. Queer
theory is a terrain, not a template, and so (to quote Jasbir Puar) “there is
no exact recipe for a queer endeavor, no a priori system that taxonomizes
the linkages, disruptions, and contradictions into a tidy vessel.”66 For ex-
ample, my interest in “unstable mixtures” might be understood as compat-
ible with Cynthia Weber’s deployment of “queer logics” (i.e., plural logics
of and/or) to expose and contest either/or dualisms such as order/anarchy
and normal/perverse67 (and to which we might add economy/sexuality
and sex/work). Yet, as I will outline in due course, my analysis also differs
from queer accounts such as Weber’s in that it is not motivated by an on-
tological commitment to queerness as fluidity, whether or not this refers
to fluid identities or the fluidity of identity tout court. This is not to dis-
count the indispensable insights of such work but rather to recognize that
queer theory’s “celebrated energies of motility and resignification” can also

IntroductIon [ 11 ]
21

be mobilized differently.68 In particular, and following Lisa Downing and


Robert Gillett, I approach queer inquiry not as the study of fluidity but as
a historical project69 and, as noted earlier, this takes the form of a “history
of the discourses” of sexuality in this volume. As my historical framework
is set out in the next chapter, suffice it to say that it is meant to contribute
to the development of queer political economy as a field of debate rather
than to provide a blueprint for a discrete position called “queer political
economy.”70
Second, the book does not conflate the history of sexuality with the
history of homosexuality, although queer theory has centered stories of
same-sex desire to counter how such desire has been discarded from and
throughout history.71 While not wishing to undermine this crucial project
in any way, I agree with Cathy J. Cohen that it is also beneficial for queer
(and feminist) scholarship to consider the “varying degrees and multiple
sites of power distributed within all categories of sexuality, including the
normative category of heterosexuality.”72 If homosexuality has been fig-
ured as both deviant and normal,73 then so have particular configurations
of heterosexuality been marked not only as natural and privileged but also
as abnormal and immoral.74 Nor should it be forgotten that the female re-
productive body has long acted as an archetype for other constructions
of the monstrous in Western discourse.75 As will be discussed at various
points in the book, the repression and production of homosexuality has
been intimately connected to the repression and production of sex work,
even if these linkages have been hidden from view. This discursive split
between homosexuality and sex work continues today, hence why feminist
theory often neglects the former and queer theory the latter.76 The book
seeks to remedy this by showing how the histories of sex work, sexuality,
and the sexual division of labor are indivisibly entangled, and how this has
involved both the specific targeting of women’s bodies77 for surveillance
and control (which queer theory could do more to confront) and the multi-
plication of and differentiation between sexualities (which, likewise, fem-
inist political economy could do more to confront). Indeed, the regulation
of sex work has been integral to the naturalization of gender difference,
to the normalization of particular expressions of heterosexual desire, and
to the reproduction of the heteronormative family on which capitalism
continues to depend.78
Lastly, a note on terminology. As Foucault was well aware, to write
about discourses of sexuality is inevitably to participate in their production
and not just their analysis. Choosing the “right” words can therefore be a
fraught and, frankly, problematic process—especially when certain words
are, to modern ears, downright offensive (e.g., “whore,” “sodomite”). Yet

[ 12 ] Capitalism’s Sexual History


the aim of this book is not to use modern-day morality to judge the past
but, in contrast, to use the past to denaturalize modern-day morality. For
example, medieval writers referred not to “sex workers” or “prostitutes”
but to “whores” and “common women” and, while these older terms and
concepts may seem distasteful today, I use them to demonstrate how
the logics they represent cannot be assumed to have simply fallen away.
Although the verb “prostitute” began to be used in English around 1530
and the label “sex worker” was coined in the 1980s,79 this does not mean
that the concept of whoredom vanished (and women can still find them-
selves being identified as “whores”).80 Thus, to insist on the exclusive use
of contemporary, politically acceptable language runs the risk not only of
anachronism but of losing sight of the very resonances and lineages that
ought to be investigated.
Equally, though, to use only historically precise terms throws up
problems of its own. This is not least because, by eschewing modern-day
concepts, we can conjure up an image of the past that is entirely distant
and unfamiliar—and yet the present has been forged out of the past. Sex
was certainly sold in the Middle Ages, even if the people who did so were
not called “sex workers,” and even if whoredom included other forms of
illicit sex.81 To insist on purity of terminology (e.g., by refusing to refer to
medieval sex workers as anything other than whores or, more accurately
still, meretrices) can create the misleading impression of essential differ-
ence between then and now, thereby ascribing false unity and coherence
to discourses that were (and are) contested. Moreover, the historian Ruth
Mazo Karras cautions that “if we analyze all our modern categories out of
existence, we are left without a language to talk about the past.”82 For ex-
ample, Jemima Repo observes that it was not until the twentieth century
that “gender” was habitually used to name “the sexual order of things,” and
so she recommends that feminists use terms such as “sex” and “sexual dif-
ference” instead.83 Yet, given that the concept of sexual difference—as in
two innately different, or opposite, sexes—can itself be read as a modern
invention,84 it is arguably no less of an instrument of modern power than
is gender. In order to get around these issues—if somewhat imperfectly—
I use a combination of contemporary terms and older ones (depending
on the time period) throughout the book. When talking about medieval
England, for instance, I refer to “sex workers” to describe people who sold
sex while also discussing the concept of “whoredom”; likewise, I use both
“sex worker” and “prostitute” when moving into the early modern period.
This is consistent with my overall approach of tracing “the erratic and dis-
continuous process whereby the past became the present”85 to politicize
how capitalism’s history is, at one and the same time, a history of sexuality.

IntroductIon [ 13 ]
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CHAPTER 1

Queer Political Economy

Once again we get another postcard from the Topsy-Turvy land where current lib-
eral intellectual life resides . . . Identity politics, fed with weak, nebulous ideas by the
likes of Foucault and Butler, has killed opposition to real concentrated power—stone
dead . . . I despair whenever I am reminded of the current direction intellectual life
has taken.

T his is an extract from a comment left by a fellow academic on a blog


post I had written about queer theory. I get his point. It captures very
well how queer theory is often treated as a frivolous indulgence rather than
as relevant to “real” issues of capitalist power relations. And it is indeed the
case that queer scholars have often remained silent on matters of political
economy to the extent that, for some, it is high time to go post-queer. In
this chapter I tackle such claims head on. Noting that the question of post-
queer is an awkward one for international political economy (IPE) (since
a field cannot move on from something that it never moved through),
I argue that queer theory has much to offer the study of global capitalism.
Yet queer scholars also need to engage more directly with feminist polit-
ical economy so as to learn from, and contribute toward, this longstanding
tradition. In order to draw out and build on these intersections, I turn to
the work of Michel Foucault and Silvia Federici, whose projects are widely
regarded as landmarks in queer theory and feminist political economy re-
spectively. By bringing these projects together, we can assemble a queer
political economy that interrogates both the linkages and the separations
between the economic and sexual spheres, and that does so in explicitly
historicizing and feminist terms.

Capitalism’s Sexual History. Nicola J. Smith, Oxford University Press (2020). © Oxford University Press.
DOI: 10.1093/oso/9780197530276.001.0001.
61

VERY QUEER PROBLEMS

Is queer history? For a number of commentators, it ought to be. In After


Queer Theory: The Limits of Sexual Politics, James Penney contends that queer
theory has “run its course, its project made obsolete by the full elaboration
of its own logic.” Queer approaches, he suggests, are “intellectually dead
discourses” due to their failure to articulate “any program of thoroughgoing
social change.”1 Similarly, in Post-Queer Politics, David V. Ruffolo argues that
queer theory’s “incessant investment in identity politics” means that it has
“reached a political peak.” He therefore highlights the need to move be-
yond queer in order to overcome its intellectual and political limitations.2
Such claims are not new but form part of queer’s own history. As Michael
O’Rourke notes, queer studies has repeatedly been dismissed as “always
already dead, buried, over, finished . . . Almost since it began we have been
hearing about the death(s) of Queer Theory.”3
In the field of IPE, the question of post-queer is especially pertinent for,
as Sara Ahmed writes, proclamations that we are “over” certain forms of
critique “create the impression that we are over what is being critiqued.”4
To go post-queer implies having first gone queer: it suggests that queer
theory’s key contributions have been recognized, responded to, and folded
into a particular field of inquiry. Yet in IPE queer theory has been, and for
the most part remains, conspicuous by its absence.5 In part, this is because
IPE has traditionally focused on the “upper circuits” of capitalist relations
such as trade and investment flows rather than the “lower circuits” such as
domestic labor and migration.6 But, as discussed in the Introduction, it is
also due to an ongoing tendency to position intimate and sexual relations
on the outside rather than the inside of global capitalism and, hence, be-
yond the proper analysis of IPE. Queer theory is not the stuff of textbooks,
core modules, and mainstream journals in IPE, and this both reflects and
sustains a wider lack of interest in sexuality. It is not possible, then, for IPE
to be over queer theory, as it has never been under it in the first place.
This oversight is problematic both intellectually and politically. Simply
put, if IPE is to analyze and contest the structural hierarchies of global capi-
talism, then it cannot afford to overlook what Judith Butler has termed the
“sexual order of political economy.”7 This means doing justice to—and not
ignoring—the contributions of queer theory as the predominant (if not
the only) field devoted to the study of sexuality.8 Although queer theory is
not exclusively concerned with sexuality—and it is by no means a synonym
for lesbian and gay studies9—sexuality is nevertheless an enduring preoc-
cupation. One of queer theory’s major contributions has been to challenge
dominant assumptions that sexuality is somehow natural, whether rooted

[ 16 ] Capitalism’s Sexual History


in biology or individual psychology. Queer scholarship has highlighted
that sexuality is not a “thing” that lies beyond or below social relations,
quietly doing its work and untroubled by power. Instead, sexuality is deeply
implicated in the constitution of modern power and warrants analysis
accordingly.10
Nor, for that matter, can queer theory afford to neglect political economy.
Indeed, a major charge made against queer theory in post-queer accounts
is precisely that it fails to advance a political economy agenda. Penney, for
instance, states that “sexual politics must be tied essentially and decisively
to an analysis of basic economic conditions” and that queer studies, in con-
trast, has been “alarmingly distanced from the critique of capitalism.” More
than this, queer theory is a “symptom of capitalist social relations” due to
its narrow focus on the subversion of identity.11 Likewise, Ruffolo contends
that queer theory’s preoccupation with subjectivity and the queer/heter-
onormative dichotomy are “unproductive considering the contemporary
complexities of neoliberal capitalism and globalization,” hence the need for
post-queer theorizing.12
Such critiques are not without foundation, for identity is indeed a cen-
tral concern for many queer scholars. Among other agendas, they have
explored how sexual and/or gender identities are constructed as mono-
lithic and static rather than as multifaceted and fluid. In so doing, they
have conceptualized queerness as “the messiness of identity, the fact that
desire and thus desiring subjects cannot be placed into discrete identity
categories.”13 Indeed, Cynthia Weber suggests that one “cannot claim to
be doing queer work” if one has “no genuine interest in those who refuse/
fail to signify monolithically in terms of sexes, genders, and sexualities.”14
Scholarship in this vein has been immensely valuable in challenging the
fixity and duality of social categories, thereby disturbing the epistemolog-
ical and methodological foundations of a great deal of social research.15
Yet some have questioned whether, despite the explicit refusal of fixed
identities, an implicit binary is reproduced in accounts that define “queer”
subjectivities in opposition to “straight” ones, and that this may actually
essentialize rather than destabilize difference.16
It is by no means the case, however, that queer is synonymous with the
fluidity of identity.17 On the contrary: what constitutes the proper domain
of queer inquiry is itself highly fluid. Despite its name, queer theory is not
a single theory or approach but rather represents an ever-shifting terrain
of debate. It is significant that—when Eve Kosofsky Sedgwick famously
described queer as an “open mesh of possibilities, overlaps, dissonances
and resonances, lapses and excesses of meaning”—she added that this
was only “one of the things” to which queer can refer and that some of

Q u e e r P ol I t I c a l e c o n o m y [ 17 ]
81

the “most exciting” queer work was not pursuing this agenda.18 Queerness
has variously been read as performativity, anti-normativity, abjection, dis-
ability, death, unreason, failure, kinship, and futurity, to name just a few
examples.19 Queer theory, therefore, has no “fixed referent”20 but rather
comprises multitudinous (and sometimes conflicting) traditions. Indeed,
as Kath Browne and Catherine J. Nash point out, the policing of queer
boundaries is at odds with commitments to queerness as fluid and man-
ifold.21 It is for this reason that queer theory is best understood as a verb
rather than a noun,22 for queer theorizing can include “any form of research
positioned within conceptual frameworks that highlight the instability of
taken-for-granted meanings and resulting power relations.”23
Nor is queer theory necessarily incompatible with political economy, es-
pecially given its multiple trajectories (it is, after all, a vast field that reaches
across the humanities and social sciences). To be sure, the history of queer
is not one in which debates about global capitalism have dominated. Yet,
over recent years, queer scholars and activists have been taking up what
Lisa Duggan and Richard Kim term a “new queer agenda.” This places eco-
nomic and social justice at the heart of the fight for sexual justice, so that
sexual struggles are explicitly defined as socioeconomic struggles, and vice
versa.24 Such a project challenges the partitioning off of sexuality from so-
called political economy issues such as low and unpaid labor, precarious
employment, access to welfare services, homelessness, incarceration, and
health care. Instead, sexuality is seen to intersect with other deep struc-
tural inequalities such as gender, race, class, dis/ability, and territory to
produce poverty, violence, and discrimination worldwide.25 For example,
Duggan is strongly critical of the equal marriage agenda on the grounds
that it reduces queer politics to the campaign for formal, legal rights rather
than for economic and social justice. This diverts attention and resources
away from struggles against poverty, inequality, and material oppression—
issues that impact significantly on queer communities and yet have come
to be defined as something other than “queer concerns.”26

MATTERS OF LIFE AND DEATH

Increasingly, then, queer scholars are making connections between polit-


ical economy and sexuality, arguing that just as global capitalism cannot
be understood without reference to the sexual sphere, then so must sex-
uality be expanded to include the analysis of capitalist power relations.
Such connections are often articulated in terms of a more open engage-
ment with Marxist theory and even as marking the emergence of “queer

[ 18 ] Capitalism’s Sexual History


Marxism.”27 Yet they also owe a clear debt to long histories of scholar-
ship in feminist political economy—a body of research that is ostensibly
overlooked in queer theory, although queer/feminist intersections in po-
litical economy have long existed via the work of J. K. Gibson-Graham,
V. Spike Peterson, and Rosemary Hennessy, among others.28 For, although
much leftist social theory has treated questions of sexuality as “footnotes”
rather than “starting points,”29 it is precisely the relations between gender,
sexuality, and capitalism that, for decades, feminist political economists
have taken as their point of departure. As Judith Butler acknowledges, so-
cialist feminists of the 1970s and 1980s “sought to establish the sphere of
sexual reproduction as part of the material conditions of life, a proper and
constitutive feature of political economy.” In so doing, they highlighted the
importance of the regulation of sexuality in the reproduction of gender
norms and the sexual division of labor, so that sexuality was understood
as “systematically tied to the mode of production” rather than as somehow
standing apart.30
Feminist political economists have taken this agenda forward by
exploring how gender and sexuality are internal to the mechanics of global
capitalism.31 As part of this project, they have explored how capitalist rela-
tions are made possible by the sexual division of labor—a division that is re-
flected most transparently in persistent wage inequalities between women
and men, and that is further exacerbated by the shift toward low-paid, tem-
porary, part-time, and flexible work across the world due to processes of
neoliberal capitalist restructuring.32 Yet such developments in the public
sphere of formal, paid work are only part of the story, for feminists also in-
sist that the functioning of the “productive economy” is in itself dependent
on that of the “reproductive economy.”33 By this they mean that systems of
regulated production and exchange are sustained by unpaid and informal
labor—such as domestic, caring, and sexual labor—so that economic pro-
duction forms part of broader and deeper processes of social reproduction.
Social reproduction refers to the practices and processes that are involved
in the production of people and populations over time and that unfold
at all levels of social existence, from the minutiae of everyday life to the
large-scale structures of global political economy.34 Important to this is an
emphasis on sexual reproduction in (literally) re/making human life, but
the term also encompasses the reproduction of the labor force and of the
“institutions, processes, and social relations associated with the creation
and maintenance of communities—and, upon which, ultimately all pro-
duction and exchange rests.”35
Although feminists contend that economic production is both part
of, and contingent on, social reproduction, they also interrogate how the

Q u e e r P ol I t I c a l e c o n o m y [ 19 ]
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derived. Many questions relating to the affinities of, or the origin of organs in, the
Annelids, resolve themselves into similar questions about the Turbellaria. For these
reasons, this group is here dealt with at greater length than the others, the interest
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The history of our knowledge of the Cestodes dates back to ancient times, as the
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first figure in Trembley's memoir on Hydra (1744).[3] The whole subject of the
increase in our knowledge of parasitic Platyhelminthes is dealt with in the standard
work, The Parasites of Man, by Leuckart,[4] and a complete list of references in
zoological literature to Cestodes and Trematodes is to be found in Bronn's
Thierreich.[5] O. F. Müller[6] and Ehrenberg founded our knowledge of the
Turbellaria, but for a long time the group remained in a most neglected condition. In
this country Montagu, G. Johnston, and in Ireland, William Thompson, discovered
several marine species, one of which, Planocera folium (from Berwick), has not
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power of regenerating lost parts. The credit of assigning the correct interpretation
to most of the various organs of fresh-water Planarians belongs to von Baer[9] and
Dugès,[10] while Mertens[11] effected a similar service for the marine forms, or
Polyclads. The minute Rhabdocoels were first successfully investigated and
classified by Oscar Schmidt.[12] The great work on this group is, however, the
monograph by von Graff.[13] A similarly comprehensive and indispensable treatise
by Lang, on the Polycladida,[14] contains references to all previous publications on
the group, among which the papers by Quatrefages, Johannes Müller, Keferstein,
Minot, and Hallez stand out conspicuously. Moseley's work[15] on the Land
Planarians of Ceylon is undoubtedly the most revolutionary paper referring to this
group, and the best contribution towards elucidating the structure of the Tricladida
at a time when the subject was very obscure. A monograph on Land Planarians is
being prepared by von Graff.

The Turbellaria are divided into: (1) Polycladida, marine forms with multiple
intestinal branches; (2) Tricladida, marine, fresh-water, and terrestrial Planarians
with three main intestinal branches; (3) the Rhabdocoelida, as varied in habit as
the Triclads, but possessing a straight and simple or slightly lobed, intestine. A
detailed description of an example of the Polyclads, and then a comparative
account of each division, will now be given.

Fig. 1.—Leptoplana tremellaris O. F. M. Seen from the dorsal surface. The


alimentary canal runs down the middle line and sends branches to the margin
of the body. × 6.

Turbellaria. I. Polycladida.

Description of Leptoplana tremellaris.

Appearance and Habits.—An account of the Polyclad Turbellaria may be fitly


prefaced by a description of a very common representative, Leptoplana tremellaris,
so called on account of the thin, flat body which executes when disturbed,
quivering or tremulous swimming movements.

Like all Polyclads, Leptoplana is marine. It is probably found on all European


shores, northwards to Greenland and southwards to the Red Sea, while vertically it
ranges from the littoral zone down to fifty fathoms. There is, however, an
apparently well-marked difference between the littoral specimens, which vary from
three-quarters to one inch in length, are brownish in colour and firm in consistency,
and the more delicate examples half an inch long, white with a brown tinge, which
occur in deeper water.

Fig. 2.—Leptoplana tremellaris. Three-quarters view from the ventral surface. The
pharynx (ph) is widely protruded through the month (mo) as in the act of
attacking prey. br, Brain with nerves, close to which are the four groups of eyes;
mg, stomach; mgc, "marginal groove"; pe, penis; sc, sucker; ut, uterus; vd, vasa
deferentia; ♀ , female genital aperture surrounded by the shell-gland; ♂ , male
aperture. (Semi-diagrammatic, and × 6.)

At low water Leptoplana may be found buried in mud or on the under surface of
stones, in pools where darkness and dampness may be ensured till the return of
the tide. It is, however, by no means easy to detect and remove it from the
encrusting Polyzoa, Ascidians, or Sponges with which it is usually associated. The
flat, soft, unsegmented body is so closely appressed to the substratum that its
presence is usually only betrayed by its movement, an even gliding motion of the
mobile body, which suggested the apt name "la pellicule animée" to Dicquemare.
The creeping surface is called ventral, the upper one dorsal, and as the broader
end of the body always goes first, it is anterior as opposed to the more pointed
posterior extremity. With a lens the characters shown in Figs. 1 and 2 may be
observed. The eyes are seen as black dots near the anterior end, and are placed
at the sides of a clear oval space, the brain. Along the transparent margin of the
body, the ends of the intestinal branches may be seen. These ramify from a lobed
stomach or main-gut, and should the specimen be mature, the "uterus" loaded with
eggs forms a dark margin round the latter (Figs. 1 and 2, ut). The ventral surface is
whitish, and through it the "pharynx," a frilled protrusible structure, may be dimly
observed. The "mouth,"[16] through which the pharynx at the time of feeding is
thrust out (Fig. 2, mo), is almost in the centre of the ventral surface. Behind this, a
white, V-shaped mark (vd) indicates the ducts of the male reproductive organs, and
still further back is the irregular opaque mark of the "shell-gland," by which the egg-
shells are formed (Fig. 2, ♀).
Fig. 3.—Leptoplana tremellaris in the act of swimming. A, Seen from the right side
during the downward stroke (the resemblance to a skate is striking); B, from
above, showing the upward stroke and longitudinal undulations of the swimming
lobes; C, side view during the upward stroke; D, transverse sections of the body
during the strokes. × 5.

Leptoplana employs two kinds of movement, creeping and swimming. Creeping is


a uniform gliding movement, caused by the cilia of the ventral surface, aided
perhaps by the longitudinal muscular layers of this surface, and is effected on the
under side of the "surface-film" of water almost as well as on a solid substratum.
Swimming is a more rapid and elegant movement, employed when alarmed or in
pursuit of prey. The expanded fore-parts of the body act as lobes, which are
flapped rapidly up over the body and then down beneath it, undulations running
rapidly down them from before backwards. The action in fact is somewhat similar
to that by which a skate swims, a resemblance pointed out long ago by Dugès[17]
(Fig. 3).

We have few direct observations on the nature of the food of Leptoplana, or the
exact mode by which it is obtained. Dalyell,[18] who observed this species very
carefully, noticed that it was nocturnal and fed upon a Nereis, becoming greatly
distended and of a green colour after the meal, but pale after a long fast.
Keferstein[19] noticed a specimen in the act of devouring a Lumbriconereis longer
than itself, and also found the radulae of Chiton and Taenioglossate Molluscs in the
intestine. That such an apparently weak and defenceless animal does overpower
large and healthy Annelids and Mollusca, has not hitherto been definitely proved.
Weak or diseased examples may be chiefly selected. The flexible Leptoplana
adheres firmly to its prey, and the rapid action of the salivary glands of its mobile
pharynx quickly softens and disintegrates the internal parts of the victim. The food
passes into the stomach (Fig. 2, mg), and is there digested. It is then transferred to
the lateral branches of the intestine, and, after all the nutritious matters have been
absorbed, the faeces are ejected with a sudden contraction of the whole body
through the pharynx into the water.

Leptoplana probably does not live more than a year. In the spring or summer,
batches of eggs are laid and fixed to algae or stones by one individual, after having
been fertilised by another. Young Leptoplana hatch out in two to three weeks, and
lead a pelagic existence till they are three or four millimetres in length. In late
summer, numbers of such immature examples may be found among sea-weeds
and Corallina in tide pools. In the succeeding spring they develop first the male and
then the female reproductive organs.

Fig. 4.—Portion of a transverse section of Leptoplana tremellaris in the hinder part of


the body. × 100. bm, Basement (skeletal) membrane; cil, cilia; d.m, diagonal
muscles; d.v.m, dorso-ventral muscles; ep, epidermis; f.p, food particles; l.g,
lateral intestinal branches cut across; l.m ext, external, and l.m int, internal
longitudinal muscle layers; m.c, glandular (mucous) cells; md, their ducts; N,
longitudinal nerve; Nu, nuclei of the intestinal epithelium; ov, ovary; ovd,
oviduct; par, cells of the parenchyma; r.d, vasa deferentia, with spermatozoa;
rm, circular musculature; rh, rhabdites; sh, cells of the shell-gland; te, testes; ve,
vasa efferentia; y.c, "yellow cells." (After Lang.)

Anatomy of Leptoplana tremellaris.—Leptoplana may be divided into


corresponding halves only by a median vertical longitudinal plane. The body and all
the systems of organs are strictly bilaterally symmetrical. Excepting the cavities of
the organs themselves, the body is solid. A connective "parenchyma" (Fig. 4, par)
knits the various internal organs together, while it allows free play of one part on
another. These organs are enclosed in a muscular body-wall, clothed externally by
the ciliated epidermis, which is separated from the underlying musculature by a
strong membrane (Fig. 4, bm), the only skeletal element in the body.

Body-Wall.—The epidermis (Fig. 4, ep) is composed of a single layer of ciliated


cells, containing small, highly refractive, pointed rods or "rhabdites" (rh), and gives
rise to deeply-placed mucous cells (m.c), which are glandular and pour out on the
surface of the body a fluid in which the cilia vibrate. The tenacious hold on a stone
which Leptoplana exerts if suddenly disturbed, or when grasping its prey, is
probably due to the increased glutinous secretion of these glands, aided perhaps
by rhabdites, which on such occasions are shot out in great numbers. The
basement membrane is an elastic skeletal membrane composed of stellate cells
embedded in a firm matrix. It serves chiefly for the origin and insertion of the dorso-
ventral muscles (d.v.m). Under the basement membrane lies a very thin layer of
transverse muscular fibres (Fig. 4, rm), which are, however, apparently absent on
the ventral surface. Then follows a stout layer of longitudinal fibres (l.m ext), and
beneath this a diagonal layer (d.m), the fibres of which intersect along the median
line in such a way that the inner fibres of one side become the outer diagonal fibres
of the other. Lastly, within this again, on the ventral surface, is a second stout
longitudinal layer (l.m int). The sucker (sc, Figs. 2 and 5) is a modification of the
body-wall at that point. In addition to the dorso-ventral muscles, there exists a
complex visceral musculature regulating the movements of the pharynx, intestine,
and copulatory organs.

Parenchyma.—The spaces between the main organs of the body are filled by a
tissue containing various kinds of cells, salivary glands, shell-glands, and prostate
glands. Besides these, however, we find a vacuolated, nucleated, thick-walled
network, and to this the word parenchyma is properly applied. Besides its
connective function, the parenchyma confers that elasticity on the body which
Leptoplana possesses in such a high degree. Pigment cells are found in the
parenchyma in many Polyclads.

Digestive System.—The general arrangement of this system may be seen in Figs.


2, 5, and 7; and may be compared, especially when the pharynx is protruded, as in
Fig. 2, with the gastral system of a Medusa. The "mouth" (there is no anus) is
placed almost in the centre of the ventral surface. It leads (Fig. 7, B, phs) into a
chamber (the peripharyngeal space) divided into an upper and a lower division by
the insertion of a muscular collar-fold (the pharynx, ph), which may be protruded,
its free lips advancing, through the mouth (Fig. 2), and is then capable of enclosing
by its mobile frilled margin, prey as large as Leptoplana itself. The upper division of
the chamber communicates by a hole in its roof[20] (the true mouth, Figs. 5 and 7,
g.m) with the cavity of the main-gut or stomach (m.g), which runs almost the length
of the body in the middle line, forwards over the brain (Fig. 5, up). Seven pairs of
lateral gut-branches convey the digested food to the various organs, not directly
however, but only after the food mixed with sea-water has been repeatedly driven
by peristalsis first towards the blind end of the gut-branches and then back towards
the stomach. Respiration is probably largely effected by this means. The epithelium
of the intestine (Fig. 4, l.g) of a starving specimen is composed of separate
flagellated cells frequently containing "yellow cells."[21] After a meal, however, the
cell outlines are invisible. Gregarines, encysted Cercariae, and Orthonectida[22]
occur parasitically in the gut-branches.
An excretory system of "flame-cells" and fine vessels has hitherto been seen only
by Schultze[23] in this species, which will not, however, resist intact the
compression necessary to enable the details to be determined. They are probably
similar to those of Thysanozoon described on p. 25.

Nervous System.—The brain, which is enclosed in a tough capsule (Fig. 5, br), is


placed in front of the pharynx, but some distance behind the anterior margin of the
body. It is of an oval shape, subdivided superficially into right and left halves by a
shallow depression, and is provided in front with a pair of granular-looking
appendages, composed of ganglion-cells from which numerous sensory nerves
arise, supplying the eyes and anterior region. Posteriorly the brain gives rise to a
chiefly motor, nervous sheath (Fig. 5, nn), which invests the body just within the
musculature. This sheath is thickened along two ventral lines (Fig. 5, ln) and two
lateral lines (n.s), but is very slightly developed on the dorsal surface. Ganglion-
cells occur on the course of the nerves, and are particularly large at the point of
origin of the great motor nerves.
Fig. 5.—Diagrammatic view of the structure of Leptoplana tremellaris
as a type of the Polycladida. The body is cut across the middle to
show the relative position of organs in transverse section. In the
posterior half the alimentary canal has been bisected and
removed from the left side, to exhibit the deeply placed nervous
sheath (nn) and the male reproductive organs. br, Brain; dp,
"diaphragm"; e, cerebral group of eyes; et, tentacular eye-group;
gr, marginal groove; gm, true mouth; lg, lateral gut-branch; ln,
longitudinal nerve stem; m, external mouth; mg, mg', main-gut,
whole, and bisected; n, sensory nerve supplying the eyes; nn,
nervous network lying on the ventral musculature; n.s, lateral
nerve; od, oviduct; ov, ovary; pe, penis (in section); ph, pharynx;
pr, prostate or "granule gland"; sc, sucker; sg, shell-gland; te,
testes; up, anterior unpaired gut-branch; ut, uterus; va, vagina (in
section); vd, vas deferens; ve, vasa efferentia; ♂ , male genital
pore; ♀, female pore.

Sense Organs.—Leptoplana possesses eyes, stiff tactile, marginal


cilia, and possibly a sense organ in the "marginal groove." The eyes,
which are easily seen as collections of black dots lying at the sides
of the brain, may be divided into two paired groups: (1) cerebral eyes
(Fig. 5, e), and (2) tentacle eyes (et), which indicate the position of a
pair of tentacles in allied forms (Fig. 8, A, t and B). Each ocellus
consists of a capsule placed at right angles to the surface of the
body in the parenchyma, below the dorsal muscles, and with its
convex face outwards. It is a single cell in which pigment granules
have accumulated. The light, however, can only reach the refractive
rods, which lie within it, obliquely at their outer ends. These rods are
in connexion with the retinal cells, and thus communicate by the
optic nerve with the brain. The cerebral eyes are really paired, and
are directed some upwards, some sideways, some downwards.

The "marginal groove" is a shallow depression of the epidermis (Fig.


5, gr) lined by cilia, and containing the ducts of very numerous gland-
cells. It runs almost parallel to the anterior margin of the body, a
short distance from it, but we have no observations on its functions.

Fig. 6.—Diagram of an eye of Leptoplana from the tentacle group. ×


600. (After Lang.)

Reproductive Organs.—Leptoplana is hermaphrodite, and, as in


most hermaphrodites, the reproductive organs are complicated. The
male organs are the first to ripen, but this does not appear to prevent
an overlapping of the periods of maturity of the male and female
products, so that when the eggs are being laid, the male organs are,
apparently, still in a functional state. The principal parts are seen in
Fig. 5. The very numerous testes (te) are placed ventrally, and are
connected with fine vasa efferentia (ve), which form a delicate
network opening at various points into the two vasa deferentia (vd).
These tubes, especially when distended with spermatozoa, may
easily be seen (Fig. 2, vd) converging at the base of the penis, and
connected posteriorly by a loop that runs behind the female genital
pore (Fig. 5). The penis (pe) is pyriform and muscular, and is divided
into two chambers, a large upper one for the spermatozoa, and a
smaller lower one for the secretion of a special "prostate" gland. The
apex of the penis is eversible and not merely protrusible, being
turned inside out when evaginated. The ovaries (Fig. 5, ov) are
numerous and somewhat spherical. They are dorsally placed, but
when fully developed extend deeply wherever they can find room to
do so, and they not only furnish the ova, but elaborate food-yolk in
the ova, as there are no special yolk-glands. The slender oviducts
(od) open at several points into the "uterus" (ut) (a misnomer, as no
development takes place within it), which encircles the pharynx, and
opens by a single duct into the vagina (va). Here the ova are
probably fertilised, and one by one invested by the shell-gland (sg)
with a secretion which hardens and forms a resistant shell. They are
then laid in plate-like masses which are attached to stones or shells.
The development is a direct one, and the young Leptoplana, which
hatches in about three weeks, has the outline of a spherical triangle,
and possesses most of the organs of the adult. After leading a
floating life for a few weeks it probably attains maturity in about nine
months.

Classification, Habits, and Structure of the Polycladida.

The Polyclads were so called by Lang on account of the numerous


primary branches of their intestine. They are free-living, purely
marine Platyhelminthes, possessing multiple ovaries, distinct male
and female genital pores (Digonopora), but no yolk-glands. The eggs
are small, and in many cases give rise to a distinct larval form,
known as "Müller's larva" (Fig. 12). The Polyclads, with one
exception,[24] fall into two sub-groups, Acotylea and Cotylea:—

Character. Acotylea. Cotylea.


Sucker A sucker absent.[25] A sucker always present
(Figs. 8, D, s; 7, A, sc).
Mouth In the middle, or In the middle, or in front
behind the middle, of of the middle, of the
the ventral surface. ventral surface.
Pharynx More or less intricately Rarely folded. Usually
folded. cylindrical or trumpet-
shaped.
Tentacles A pair of dorsal A pair of marginal
tentacles usually tentacles (except in
present. Anonymus).
Development Usually direct. Larva Müller's larva present.
when present, not a Metamorphosis,
typical Müller's larva. however, extremely
slight.

Fig. 8 shows that, starting with a member (A, D) of each division, in


which the mouth is almost in the middle of the ventral surface, and
the brain and sense organs somewhat remote from the anterior end,
we find in the Acotylea a series leading to an elongated form
(Cestoplanidae), in which the mouth, pharynx, and genital pores are
far back near the hinder end of the body; while in the Cotylea the
series leads similarly to the elongated Prosthiostomatidae, in which,
however, the pharynx and external apertures are in the front part of
the body. This view of the morphology of the Polyclads is due to
Lang, and is based on the assumption that the more radially-
constructed forms (Fig. 8, A, D) are the primitive ones.

Fig. 7.—Diagrammatic vertical longitudinal sections: A, Of


Prosthiostomum (type of Cotylea); B, of Leptoplana; C, of
Cestoplana (types of Acotylea). (After Lang.) These figures
illustrate the changes which follow the shifting of the mouth from a
central position (B) to either end of the body. br, Brain; dphm,
"diaphragm"; gm, true mouth; lg, openings of lateral gut-branches;
m, mouth; mg, main-gut or stomach; mgbr, median gut-branch;
ph, pharynx; ph.m, aperture in pharyngeal fold; phs,
peripharyngeal sheath; sc, sucker; ♂, male, and ♀, female, genital
aperture.

Fig. 8.—Chief forms of Polycladida: A-C, Acotylea; D-F, Cotylea. A,


Planocera graffii Lang, nat. size; B, Stylochoplana maculata
Stimps, × 7; C, Cestoplana rubrocincta Lang, × 4⁄3; D, Anonymus
virilis Lang, × 3, ventral surface; E, Thysanozoon brocchii Grube,
nat. size; the head is thrown back and the pharynx (ph) is
protruded. F, Prosthiostomum siphunculus Lang, × 3. Br, Brain;
CG, cerebral eye group; DM, true mouth; Ey, marginal eyes; m,
mouth; MG, main-gut or stomach; P, dorsal papillae; Ph, pharynx;
s, sucker (ventral); T, tentacles; UP, dorsal median gut-branch. ♂,
male, and ♀, female, genital aperture, except in D, where ♂ refers
to the multiple penes. (After Lang and Schmidt.)

Classification of Polycladida.

ACOTYLEA.
Family. Genus. British
Representatives.
Planocera (Fig. 8,
Planocera folium
A).
Grube. Berwick-
Imogine.
on-Tweed.
Planoceridae. Conoceros.
Stylochoplana
With dorsal tentacles. Stylochus.
maculata Quatref.
Mouth sub-central. Stylochoplana
Among brown
(Fig. 8, B).
weeds in
Diplonchus.
Laminarian zone.
Planctoplana.
Leptoplana
tremellaris O. F.
Müll.
Discocelis.
L. fallax Quatref.
Leptoplanidae. Cryptocelis.
Plymouth.
Without dorsal Leptoplana.
L. droebachensis
tentacles. Penis Trigonoporus.
Oe. Plymouth
directed backwards. ?Polypostia (see
Sound.
p. 27).
L. atomata O. F.
Müll. Doubtful
species.
Cestoplana (Fig. 8,
Cestoplanidae. C).
No tentacles. Body In Mediterranean
elongated. Penis and on French
directed forwards. side of the
Channel.
Enantiidae.
No sucker. No
tentacles. Main-gut Enantia.
very short. External Adriatic Sea.
apertures as in
Euryleptidae.

COTYLEA.
Anonymidae. Anonymus (Fig. 8,
Mouth central. No D).
tentacles. With two Naples (two
rows of penes. specimens).
Pseudoceridae. Thysanozoon (Fig.
Marginal tentacles 8, E).
folded. Mouth in Pseudoceros.
anterior half. Yungia.
Prostheceraeus
vittatus Mont. On
west coast.
P. argus Quatref.
Guernsey.
Cycloporus
Euryleptidae. papillosus Lang.
Tentacles usually Prostheceraeus. On Ascidians in
present and pointed, Cycloporus. 2-30 fms.
or represented by Eurylepta. Eurylepta cornuta
two groups of eyes. Oligocladus. O.F. Müll. On
Mouth close to Stylostomum. sponges and
anterior end. Aceros. shells, 2-10 fms.
Pharynx cylindrical. Oligocladus
sanguinolentus
Quatref.
O. auritus Clap.
Doubtful.
Stylostomum
variabile Lang.
Prosthiostomatidae.
Tentacles absent.
Body elongated.
Prosthiostomum
Pharynx long,
(Fig. 8, F).
cylindrical. Penis
with accessory
muscular vesicles.
Appearance and Size of Polyclad Turbellaria.—Polyclads are
almost unique amongst animals in possessing a broad and thin,
delicate body that glides like a living pellicle over stones and weeds,
moulding itself on to any inequalities of the surface over which it is
travelling, yet so fragile that a touch of the finger will rend its tissues
and often cause its speedy dissolution. The dorsal surface in a few
forms is raised into fine processes (Planocera villosa), or into hollow
papillae (Thysanozoon brocchii), and in very rare cases may be
armed with spines (Acanthozoon armatum,[26] Enantia spinifera); in
others, again, nettle-cells (nematocysts) are found (Stylochoplana
tarda, Anonymus virilis). Some Polyclads, especially the pelagic
forms, are almost transparent; in others, the colour may be an
intense orange or velvety black, and is then due to peculiar deposits
in the epidermal cells. Between these two extremes the colour is
dependent upon the blending of two sources, the pigment of the
body itself and the tint of the food. Thus a starved Leptoplana is
almost or quite white, a specimen fed on vascular tissue reddish.
Many forms are coloured in such a way as to make their detection
exceedingly difficult, but this is probably not merely due, as Dalyell
supposed, to the substratum furnishing them with food and thus
colouring them sympathetically, but is probably a result of natural
selection.

The largest Polyclad, the bulkiest Turbellarian, is Leptoplana gigas


(6 inches long and 4 in breadth), taken by Schmarda, free-
swimming, off the coast of Ceylon. The largest European form is
Pseudoceros maximus, 3½ inches in length and stoutly built. A
British species, Prostheceraeus vittatus, attains a length of from 2 to
3 inches. These large forms, especially the Pseudoceridae (pre-
eminently the family of big Polyclads), are brightly coloured, and
usually possess good swimming powers, since, being broad and flat,
they are certainly not well adapted for creeping rapidly, and this is
well shown by the way these Polyclads take to swimming when in
pursuit of prey at night. The size of any individual is determined,
amongst other factors, by the period at which maturity sets in, after
which probably no increase takes place. Polyclads apparently live
about twelve months, and mature specimens of the same species
vary from ½ inch to 2½ inches in length (Thysanozoon brocchii),
showing that growth is, under favourable conditions, very rapid.

Habits of Polyclad Turbellaria.—Polyclads are exclusively marine,


and for the most part littoral, animals. Moreover, there is no evidence
of their occurrence in those inland seas where certain marine
animals (including one or two species of otherwise characteristically
marine Rhabdocoelida, p. 46) have persisted under changed
conditions. From half-tide mark down to 50 fathoms, some Polyclads
probably occur on all coasts, but as to their relative abundance in
different seas we have very little accurate information. The southern
seas of Europe possess more individuals and species than the
northern, and probably the maximum development of the group
takes place on the coasts and coral islands of the tropics.[27] No
Polyclads have been taken below 60 fathoms; but their delicacy and
inconspicuousness render this negative evidence of little value. Six
truly pelagic forms, however, are known,[28] and these are interesting
on account of their wide distribution (three occurring in the Atlantic,
Pacific, and Indian oceans), and also from the distinct modifications
they have undergone in relation to their pelagic existence.

Whatever may be the interpretations of the fact, Polyclads are


notoriously difficult to detect, and this fact doubtless explains the
scanty references to them by the older naturalists who collected
even in tropical seas. Lang, who worked seven years at Naples,
added to the Mediterranean fauna as many Polyclads as were
previously known for all Europe, in spite of the assiduous labours of
his predecessors, Delle Chiaje and Quatrefages. Again Hallez,
collecting at Wimereux at low-water, obtained some twenty
specimens of Leptoplana tremellaris in an hour, while some other
collectors working by his side could only find two or three. Yet, even
making allowance for the difficulty of finding Polyclads, few of them
appear to be abundant.

Leptoplana tremellaris is frequently associated with colonies of


Botryllus, and if separated soon perishes, whereas the free-living
individuals are distinctly hardy (Hallez). A closely allied but possibly
distinct form lives upon the surface of the Polyzoon Schizoporella, on
the French side of the Channel, and cannot long endure separation
from its natural habitat, to which it is adaptively coloured. A striking
case of protective mimicry is exhibited by Cycloporus papillosus, on
the British coasts. This species, eminently variable in colour and in
the presence or absence of dorsal papillae, is usually a quarter of an
inch in length and of a firm consistency. Fixed by its sucker to
Polyclinid and other Ascidians, Cycloporus appears part and parcel
of the substratum, an interesting parallel to Lamellaria perspicua,[29]
though we are not justified in calling the Polyclad parasitic. Indeed,
though a few cases of association between Polyclads and large
Gasteropods, Holothurians, and Echinids are known,[30] there is only
one case, that of Planocera inquilina,[31] in the branchial chamber of
the Gasteropod Sycotypus canaliculatus, which would seem to bear
the interpretation of parasitism. The jet-black Pseudoceros velutinus
and the orange Yungia aurantiaca of the Mediterranean, are large
conspicuous forms with no attempt at concealment, but their taste,
which is not known, may protect them. Other habits, curiously
analogous with devices employed by Nudibranch Mollusca (compare
Thysanozoon brocchii with Aeolis papillosa), emphasise the
conclusion that the struggle for existence in the littoral zone has
adapted almost each Polyclad to its particular habitat.

As regards the vertical distribution of this group on the British coasts,


Leptoplana tremellaris has an extensive range, and appears to come
from deeper to shallower water to breed.[32] In the upper part of the
Laminarian zone, Cycloporus papillosus, and, among brown weeds,
Stylochoplana maculata are found. At and below lowest water-mark
Prostheceraeus vittatus, P. argus, and Eurylepta cornuta occur.
Stylostomum variabile and Oligocladus sanguinolentus, though
occasionally found between tide-marks, especially in the Channel
Islands, are characteristic, along with Leptoplana droebachensis and
L. fallax, of dredge material from 10 to 20 fathoms.
Locomotion.—Locomotion is generally performed by Polyclads at
night when in search of food, and two methods, creeping and
swimming, are usually employed—creeping by the cilia, aided
possibly, as in the case of some Gasteropod Mollusca, by the
longitudinal muscles of the ventral surface; and swimming, by
undulations of the expanded margins of the body. In the former case
the cilia work in a glandular secretion which bathes the body, and
enables them to effect their purpose equally well on different
substrata. The anterior region is generally lifted up, exploring the
surroundings by the aid of the tentacles, which are here usually
present. The rest of the body is closely appressed to the ground.

Swimming is particularly well performed by the Pseudoceridae,


certain species of Prostheceraeus, the large Planoceridae, some
Stylochoplana, Discocelis, and Leptoplana, and in the same manner
as in Leptoplana tremellaris (p. 9). In Cryptocelis, Leptoplana alcinoi,
and L. pallida, however, the whole body executes serpentine
movements like an active leech (e.g. Nephelis); a cross section of
the body would thus present the same appearance during the whole
movement. Many Polyclads, notably Anonymus (Lang), if irritated,
spread out in all directions, becoming exceeding thin and
transparent.

Fig. 9.—Discocelis lichenoides Mert. (after Mertens), creeping on the


inner side of a glass vessel by means of the lobes of the extended
and exceedingly mobile pharynx (ph). These lobes also serve to
enclose Crustacea (a), and one lobe may then be withdrawn
independently of the rest, back into the body (b). The brain (br)
and shell-gland (sg) are shown by transparency.
Discocelis lichenoides, Planocera graffii, and Anonymus virilis have
peculiar modes of progression. The first, according to Mertens, will
climb up the sides of a vessel by means of the expanded lobes of
the pharynx (Fig. 9, ph), a habit of considerable interest, since we
know that certain Ctenophores—Lampetia, for instance—progress
when not swimming on the expanded lobes of their "stomach."[33]
Planocera and Anonymus creep by extending parts of the anterior
margin and dragging the rest of the body behind. In consequence,
the brain and dorsal tentacles may come to lie actually behind the
middle of the body, and thus no definite anterior end or "head"
advances first. Along with this curious habit it may be noticed (Lang)
that the radial symmetry of the body is well marked; but even without
accepting this author's suggestion of the concurrent development of
a "head" with locomotion in a definite direction, the facts, whether
these two forms are primitive or not, are highly interesting.

Food.—Though we are probably right in calling Polyclads a


carnivorous group, the food of very few forms has been ascertained.
Those which possess a large frilled pharynx (most Acotylea)
probably enclose and digest large, and, it may be, powerful prey, as
appears to be the case in Leptoplana tremellaris. Cryptocelis alba
has been seen by Lang with the pharynx so distended, owing to a
large Drepanophorus (Nemertine) which it contained, as to resemble
a yolk-sac projecting from the under surface of an embryo. The
Cotylea such as Thysanozoon, with a bell- or trumpet-shaped
pharynx, are fond of fixing this to the side of the aquarium, but
whether they thus obtain minute organisms is not clear.
Prosthiostomum shoots out its long pharynx with great vehemence
(Fig. 8, F) and snaps up small Annelids by its aid (Lang). Those
Polyclads which, as Cycloporus and others, are definitely associated
with other organisms are not certainly known to feed upon the latter,
though "Planaria velellae" has been seen by Lesson[34] devouring
the fleshy parts of its host. The salivary glands which open on the
lips and the inner surface of the pharynx powerfully disintegrate the
flesh of the prey. Digestion takes place in the main-gut, and the
circulation of the food is accomplished by the sphinctral musculature
of the intestinal branches (conf. Leptoplana, p. 13).

Fig. 10.—Diagram of the musculature, causing peristaltic movements


of the intestinal branches of Polyclads. (After Lang.)

A distinct vent or anus is always absent. After a meal the faecal


matter collects in the main-gut, and is discharged violently by the
pharynx into the water. In a few species, however, the intestinal
branches open to the exterior (Lang). Yungia aurantiaca, a large and
abundant Neapolitan form, possesses such openings over the
greater part of the dorsal surface; Cycloporus papillosus has
marginal pores; Oligocladus sanguinolentus apparently possesses
an opening at the posterior end of the main-gut; and Thysanozoon
brocchii frequently rends at this point, in consequence of the
accumulation of food.

Respiration.—The oxygen of the atmosphere dissolved in the sea-


water is, in default of a special circulatory fluid, brought to the tissues
of Polyclads in two ways. The ciliated epidermis provides a constant
change of the surrounding water, by which the superficial organs
may obtain their supply; and the peristaltic movements of the
digestive system, aided by the cilia of the endoderm cells, ensure a
rough circulation of the sea-water, which enters along with the food,
to the internal organs. The papillae of Thysanozoon brocchii,
containing outgrowths of the intestinal branches, are possibly so
much additional respiratory surface, although still larger forms (other
Pseudoceridae) are devoid of such outgrowths.

Excretion.—The excretory system of only one Polyclad


(Thysanozoon brocchii) is accurately known. Lang, by compressing
light-coloured specimens, found the three parts of the system known

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