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The Ethics of Storytelling
ii

Explorations in Narrative Psychology

Mark Freeman
Series Editor

Books in the Series

Speaking of Violence
Sara Cobb
Narrative Imagination and Everyday Life
Molly Andrews
Narratives of Positive Aging: Seaside Stories
Amia Lieblich
Beyond the Archive: Memory, Narrative, and
the Autobiographical Process
Jens Brockmeier
The Narrative Complexity of Ordinary Life: Tales from the Coffee Shop
William L. Randall
Rethinking Thought: Inside the Minds of Creative Scientists and Artists
Laura Otis
Life and Narrative: The Risks and Responsibilities of Storying Experience
Edited by Brian Schiff, A. Elizabeth McKim, and Sylvie Patron
Not in My Family: German Memory and Responsibility
After the Holocaust
Roger Frie
A New Narrative for Psychology
Brian Schiff
Decolonizing Psychology: Globalization, Social Justice,
and Indian Youth Identities
Sunil Bhatia
Entangled Narratives: Collaborative Storytelling and
the Re-​Imagining of Dementia
Lars- ​Christer Hydén
The Ethics of Storytelling: Narrative Hermeneutics, History,
and the Possible
Hanna Meretoja
The Ethics
of Storytelling
Narrative Hermeneutics, History,
and the Possible

Hanna Meretoja

1
iv

1
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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

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© Oxford University Press 2018

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Library of Congress Cataloging-in-Publication Data

Names: Meretoja, Hanna, 1977– author.
Title: The Ethics of storytelling : narrative hermeneutics, history,
and the possible / Hanna Meretoja.
Description: New York : Oxford University Press, 2018. |
Series: Explorations in narrative psychology |
Includes bibliographical references and index.
Identifiers: LCCN 2017028106 | ISBN 9780190649364 (hardback)
Subjects: LCSH: Storytelling. | Narration (Rhetoric)—Moral and ethical aspects. |
Narration (Rhetoric)—Psychological aspects. | Narration (Rhetoric)—Social aspects. |
Self-perception in literature. | Social perception in literature. | Awareness in literature. |
Imagination in literature. | BISAC: PSYCHOLOGY / Social Psychology. |
PHILOSOPHY / Mind & Body. | LITERARY CRITICISM / Semiotics & Theory.
Classification: LCC PN56.S7357 M47 2017 | DDC 808/.036—dc23
LC record available at https://lccn.loc.gov/2017028106

9 8 7 6 5 4 3 2 1
Printed by Sheridan Books, Inc., United States of America
To Alma and Eliel, my beloved storytelling animals
vi
CONTENTS

Acknowledgments ix

1. Introduction: Toward an Ethics of Storytelling 1

2. Narrative Hermeneutics 43
3. Storytelling and Ethics 89
4. The Uses and Abuses of Narrative for Life: Julia Franck’s
Die Mittagsfrau 149
5. Narrative Ethics of Implication: Günter Grass and Historical
Imagination 179
6. Narrative Dynamics, Perspective-​Taking, and Engagement:
Jonathan Littell’s Les Bienveillantes 217
7. Transforming the Narrative In-​Between: Dialogic Storytelling and
David Grossman 255
8. Conclusion: Struggles over the Possible 299

References 309
Index 333
vi
ACKNOWLEDGMENTS

One of the guiding ideas of this book is that people and narratives become
who and what they are in dialogue with other people and their stories. This
is true of this book as well. It has taken shape in a conversation with innu-
merable people whose stories, thoughts, affection, and support have made
its writing possible. I can here name only some of those to whom I am most
indebted.
The relationship between storytelling and ethics has occupied my mind
for such a long time that it is difficult to say when exactly I began work on
this book. I was reflecting on these issues already when writing my pre-
vious book, The Narrative Turn in Fiction and Theory (Palgrave Macmillan,
2014), and while finishing it, I felt compelled to develop a more systematic
and more broadly interdisciplinary account of narrative hermeneutics and
a hermeneutic narrative ethics. This book seeks to provide such a system-
atic account of the project of narrative hermeneutics that I have begun in
my earlier work and to zoom in on its ethical implications. The feedback
I received on my first book has helped me enormously to give shape to
this book.
I owe a special gratitude to my colleagues who have read and provided
insightful comments on parts of the manuscript: Eneken Laanes, Erin
McGlothlin, Frans Svensson, Henrik Skov Nielsen, Marco Caracciolo,
Maria Mäkelä, Robert Eaglestone, and my colleagues at the Department of
Comparative Literature (University of Turku), in particular Aino Mäkikalli,
Jouni Teittinen, Kaisa Ilmonen, Liisa Steinby, Lotta Kähkönen, Tiina
Käkelä-​Puumala, and Tintti Klapuri. I wrote a first draft of the Grass chap-
ter (Chapter 5) in the spring of 2011 as part of the Academy of Finland
research project Literature and Time: Time and Agency in Modern Literature
(led by Liisa Steinby); the feedback from that research group informs my
analyses of temporality in this book. Members of the research project The
Ethics of Storytelling and the Experience of History in Contemporary Literature
and Visual Arts (Emil Aaltonen Foundation), which I had the honor to lead
x

in 2013–​2016, Kaisa Kaakinen, Ilona Hongisto, Mia Hannula, and Riitta

Jytilä, have commented on earlier drafts of several chapters of the book,
and the group has provided a stimulating environment for developing this
project.
While writing this book, I have had the pleasure to work at both the
University of Turku and the University of Tampere with wonderful col-
leagues with whom I established in 2014 the interdisciplinary research center
Narrare: Centre for Interdisciplinary Narrative Studies (Tampere) and in 2015
SELMA: Centre for the Study of Storytelling, Experientiality and Memory (Turku).
These research communities have provided a supportive environment for my
work. In the latter, Maarit Leskelä-​Kärki and Päivi Kosonen have been particu-
larly inspiring partners of dialogue. I owe a huge debt to my brilliant students
at both Tampere and Turku. A special thanks to Eevastiina Kinnunen, who has
gone through the manuscript with exceptional care and thoroughness.
Conversations with and feedback from my colleagues around the world
have contributed to this book in numerous ways: I want to thank, in par-
ticular, Molly Andrews, Aleida Assmann, Mieke Bal, Sunil Bhatia, Heidi
Bostic, Cassandra Falke, Rita Felski, Roger Frie, Jane Hiddleston, Matti
Hyvärinen, Teemu Ikonen, Stefan Iversen, Kuisma Korhonen, Liesbeth
Korthals Altes, Markku Lehtimäki, Jakob Lothe, Pirjo Lyytikäinen,
Julia Nitz, Sakari Ollitervo, Ann Phoenix, Merja Polvinen, Ann Rigney,
Michael Rothberg, Hannu Salmi, Brian Schiff, Max Silverman, Howard
Sklar, Anneke Sools, Maria Tamboukou, Pekka Tammi, Shane Weller, and
Benjamin Wihstutz. Audiences at numerous conferences and symposia
over the years have also provided valuable feedback. In particular, I want
to express my gratitude for invitations to present my research at Royal
Holloway (University of London, March 2013), The American University of
Paris (where I was a visiting professor in 2013–​2014), and The Centre for
Narrative Research (University of East London, December 2015), to give a
keynote at the Narrative Matters conference in Victoria (June 2016), and to
speak at The Future of Literary Studies conference (Oslo, June 2016).
I am grateful to Matthew James and Avril Tynan for their insightful,
nuanced, and patient linguistic advice. It has been a joy to work with Abby
Gross and Courtney McCarroll of Oxford University Press; their profes-
sionalism, reliability, and support have made the whole publication process
a most enjoyable experience. I also appreciate the helpful comments pro-
vided by several anonymous readers. I am so happy that Bracha L. Ettinger
gave me the permission to use her beautiful picture (Eurydice no. 35) as the
cover of this book.
I am deeply grateful to all of my friends whose stories have inter-
sected with mine, shaping my modes of thought, perception, and affect in

[x] Acknowledgments
innumerable ways. I feel particularly privileged for the in-​depth dialogue
I have been able to engage in with kindred philosophical souls over the
years on the topics of this book. A few of them have been so important for
this project that I want to thank them for giving me much more than just
valuable comments and encouragement. Jens Brockmeier, my fellow nar-
rative hermeneuticist, has commented on a large part of the manuscript
with great insight, generosity, and perceptiveness, from the broad interdis-
ciplinary perspective that is singular to him. I want to thank Anna Reading
for her warm friendship and stimulating discussions; she gave me the most
detailed comments on the Introduction that I have ever received—​in my
favorite spot on the pier of our summer house. I am immensely grateful
to Mark Freeman for supporting this project from early on, for his willing-
ness to include it in his inspiring series, and for generous and thoughtful
comments at various stages of the project. I want to thank Andreea Ritivoi
for just being there and making me feel, by speaking the same philosophi-
cal language, that what I do may actually speak to someone out there. I am
inexpressibly thankful to Colin Davis for his unique friendship, affection,
and unwavering support—​for helping me become more than I would have
been able to without him.
My heartfelt thanks to my parents and siblings for all their love and
support. My deepest gratitude goes to the person with whom I have shared
both my intellectual and non-​intellectual life for more than 23 years.
Valtteri Viljanen has set an example for me with his courage, perseverance,
and sense of humor. This book has benefited enormously from his philo-
sophical perceptiveness and passion, and it is through our daily narrative
dialogue that I have become the thinker, writer, and person I am. Alma
and Eliel have taught me the power of narrative imagination: from them
I have learned that with enough imagination, almost anything is possible.
Ultimately, it is their love, patience, encouragement, and wisdom that has
made this book possible.

I gratefully acknowledge the permission from publishers to draw on the fol-

lowing earlier publications, although most of the material I have used from
them has been heavily revised and reworked. In Chapters 2 and 7, I have
integrated passages from “For Interpretation” (Storyworlds 8 [2]‌, 2016,
pp. 97–​117). The discussion of Nussbaum in Chapter 3 owes something to
my chapter “A Sense of History—​A Sense of the Possible: Nussbaum and
Hermeneutics on the Ethical Potential of Literature” in Values of Literature
(edited by Hanna Meretoja, Saija Isomaa, Pirjo Lyytikäinen, and Kristina
Malmio, Brill Rodopi, 2015, pp. 25–​46), its discussion of the possible con-
tains passages from “Exploring the Possible: Philosophical Reflection,

Acknowledgments [ xi ]
xi

Historical Imagination, and Narrative Agency” (Narrative Works 6, 2016,

pp. 92–​107), and its section on the non-​subsumptive mode of under-
standing draws on “From Appropriation to Dialogic Exploration: A Non-​
Subsumptive Model of Storytelling” in Storytelling and Ethics: Literature,
Visual Arts, and the Power of Narrative (edited by Hanna Meretoja and Colin
Davis, Routledge, 2018). Some key ideas of the chapter are presented in a
very condensed form in “Narrative Hermeneutics and the Ethical Potential
of Literature” in The Future of Literary Studies (edited by Jakob Lothe, Novus
Press, 2017, pp. 147–159). Chapter 4 grew from “On the Use and Abuse
of Narrative for Life: Toward an Ethics of Storytelling,” which appeared
in Narrative and Life: The Risks and Responsibilities of Storying Life (edited
by Brian Schiff, Sylvie Patron, and A. Elizabeth McKim, Oxford University
Press, 2017, pp. 75–​97). Chapter 5 contains, in a modified form, passages
from “An Inquiry into Historical Experience and Its Narration: The Case of
Günter Grass” (Spiel: Siegener Periodicum for International Empiricist Literary
Scholarship special issue, “Towards a Historiographical Narratology,” edited
by Julia Nitz and Sandra Harbart Petrulionis, 30 [1], 2011, pp. 51–​72). An
earlier version of Chapter 6 was published as “History, Fiction and the
Possible: Jonathan Littell’s Les Bienveillantes” (Orbis Litterarum 81 [5],
2016, pp. 371–​404).

[ xii ] Acknowledgments
The Ethics of Storytelling
xvi
CHAPTER 1

Introduction
Toward an Ethics of Storytelling

T hat stories are indispensable for human existence is an idea that reaches
back at least to One Thousand and One Nights: as Scheherazade’s fasci-
nating tales delay and ultimately prevent her murder by King Shahryar,
storytelling becomes, quite literally, an art of survival. At the same time,
entanglement in narratives has notoriously raised suspicion. In the
Western imagination, Don Quixote and Emma Bovary epitomize the dan-
gers of reading too many stories, and the crisis of European humanism,
in the wake of two world wars and the Holocaust, thoroughly problema-
tized the imposition of narrative order on history and our experience of
the world. The protagonist of Jean-​Paul Sartre’s La Nausée (1938, Nausea)
encapsulates this sensibility: “you have to choose: to live or to recount”
(1965, p. 61).1 Over the past few decades, discourse surrounding the ethi-
cal significance of narrative for human existence has gained unprecedented
urgency and intensity. As the debate on the ethics of storytelling has
become one of the liveliest in interdisciplinary narrative studies, positions
have also become increasingly polarized: as theorists of narrative argue
“for” or “against” narrativity, the quarrel tends to be whether narratives
are “good” or “bad” for us.2
As part of the “narrative turn,” the idea that stories are not only indis-
pensable but also inherently beneficial for us has become hugely popular.3
A wide range of thinkers have come to share Paul Ricoeur’s view that only
“a life narrated” can be “a life examined” and hence worth living (1991b,
p. 435). Many contemporary novelists seem to agree: while Paul Auster
2

asserts that “stories are the fundamental food for the soul” (Irwin & Auster,
2013, p. 46), Jeanette Winterson’s narrator compares storytelling to light-​
housekeeping and presents “stories going out over the waves, as markers
and guides and comfort and warning” (2004, p. 41). Yet the strong nar-
rativist position has provoked a fierce counter-​reaction. One of the most
outspoken representatives of the “against narrativity” movement, Galen
Strawson, argues provocatively, “the more you recall, retell, narrate your-
self, the further you risk moving away from accurate self-​understanding,
from the truth of your being” (2004, p. 447).
Against the backdrop of this polarized debate, there is a need for a
theoretical-​analytical framework that allows us to explore the ethical
complexity of the roles that narratives play in our lives. In this book, I set
out to develop such a framework—​one that acknowledges both the eth-
ical potential and the risks of storytelling. My starting point is that a
nuanced analysis of the uses and abuses of narrative for life is possible
only when we are sensitive to the ways in which narratives as practices
of sense-​making are embedded in social, cultural, and historical worlds.
We are always already entangled in webs of narratives. They are integral
to the world that precedes us, and they make it possible for us to develop
into subjects who are capable of narrating their experiences, sharing
them with others, and telling their own versions of the stories they have
inherited. Each cultural and historical world functions as a space of possi-
bilities that encourages certain modes of experience, thought, and action,
and discourages or disallows others, and stories play a constitutive role
in establishing the limits of these worlds—​both enabling experience and
delimiting it.
This book aims to develop an approach that invites analyzing both how
narratives enlarge the dialogic spaces of possibilities in which we act, think,
and reimagine the world together with others, and how they restrain or
impoverish these spaces. Precisely this, I argue, is a crucial but generally
overlooked dimension of the ethics of storytelling: narratives both expand
and diminish our sense of the possible. I call my approach a narrative her-
meneutics because it treats narratives as culturally mediated practices of
(re)interpreting experience, and I will explore its ethical implications.4 It
aims to provide a philosophically rigorous, historically sensitive, and ana-
lytically subtle approach to the ethical stakes of the debate on the narrative
dimension of human existence. On the basis of narrative hermeneutics,
I propose a hermeneutic narrative ethics, which acknowledges that narrative
practices can be oppressive, empowering, or both, and provides resources
for analyzing the different dimensions of the ethical potential and dangers
of storytelling.

[2] The Ethics of Storytelling

 Narrative hermeneutics emphasizes that interpretation does not con-
cern only our engagement with texts; it characterizes our whole being in
the world and is the basic structure of experience, narrative, and memory.
When we go through meaningful experiences, weave them into stories, and
remember them in a certain light and from a certain perspective, we engage
in interpretative processes of sense-​making. It is as “self-​interpreting ani-
mals” (Taylor, 1985) that we narrate our experiences and fashion our lives.
Narratives are interpretative practices through which we make sense of our
lives, and these meaning-​making practices are ethically charged. As Ricoeur
puts it, narratives are “never ethically neutral” (1992, p. 140), and story-
telling can function as “a provocation to be and to act differently” (1988,
p. 249).
I will explore the ethical potential of storytelling with particular atten-
tion to that of narrative fiction. While it has long been taken for granted
that literature is beneficial for us, this is no longer necessarily the case.
Against the backdrop of the current crisis of the humanities, many phi-
losophers, psychologists, and literary scholars have defended the value of
narrative fiction by drawing attention to its cognitive and ethical signif-
icance for our development as human beings, moral agents, and demo-
cratic citizens, particularly insofar as it boosts our capacity for empathetic
perspective-​taking. Martha Nussbaum describes such a capacity in terms
of narrative imagination, which she sees as a major counterforce to the anti-
democratic tendencies in the contemporary world:

[Narrative imagination] means the ability to think what it might be like to be

in the shoes of a person different from oneself, to be an intelligent reader of
that person’s story, and to understand the emotions and wishes and desires that
someone so placed might have. (2010, pp. 95–​96)

Cognitive scientists have recently sought to provide empirical support

for such views by arguing that narrative fiction is more “transformative”
than nonfiction (Djikic et al., 2009) and has a stronger impact on our “social
perception and moral self-​concept” (Hakemulder, 2000). David Comer Kidd
and Emanuele Castano’s (2013) widely reported study asserts that the affec-
tive and cognitive skills involved in understanding that others have beliefs,
desires, and intentions that are different from one’s own—​what cognitive
scientists call “theory of mind”—​are improved by reading literary fiction
compared with reading nonfiction, popular fiction, or nothing at all.5 The
researchers not only claim that after short-​term exposure to fiction, people
do better on tests that measure empathy, social perception, and emotional
intelligence, but also suggest that in addition to temporary enhancement

Introduction [3]
4

of these skills, long-​term effects could be achieved by regular engagement

with literature. Recently, Kidd and Castano (2016) have reached the same
conclusion through a different method, but other researchers have failed to
reproduce their results (Panero et al., 2016).
Irrespective of the short-​term effects of reading fiction, it is far from
evident how long-​term effects could be measured in reliable ways. To me
it seems far more plausible to argue that literature cultivates our ability
to perceive the world from multiple perspectives, or at least increases our
awareness of and sensitivity to such multiplicity—​what I call perspective-​
awareness and perspective-​sensitivity—​than to argue that literature makes
us ethical in the sense of causing us to engage in moral action. As many
critics have observed, there is a significant difference between embracing
the perspectives of others—​or imagining what one might do in hypotheti-
cal scenarios—​and actually carrying out concrete actions in the real world.
Suzanne Keen, for example, argues that “the very fictionality of novels pre-
disposes readers to empathize with characters,” because “fictional worlds
provide safe zones for readers’ feeling empathy without experiencing a
resultant demand on real-​world action” (2007, p. 4). Indeed, there is ample
evidence that reading fiction is no guarantee of ethical action. As George
Steiner famously reminds us, the Holocaust seriously undermined the long
unquestioned belief in the “humanizing force” of literature: “We know now
that a man can read Goethe or Rilke in the evening, that he can play Bach
and Schubert, and go to his day’s work at Auschwitz in the morning” (1967,
p. 15).
However, perspective-​awareness may be a necessary condition for moral
agency, even if it is not a sufficient condition. Several moral philosophers
have argued that imagination is indispensable not only for our cognitive
and emotional development, but also for our ethical development and
agency.6 As moral agents we are capable of initiating new processes in the
world, and in making decisions and embarking on actions we must imag-
ine the myriad potential consequences of these actions—​how they might
affect others and change the world we co-​inhabit. Without such a power of
imagination, our existence as acting subjects would be truncated. A sense
of the possible—​a sense of how things could be otherwise—​is integral to
moral agency and to the ethical imagination of individuals and communi-
ties. It has transformative potential.
I will explore the capacity of narratives to expand our sense of the pos-
sible in relation to five additional aspects of the ethical potential of story-
telling: narrative’s contribution to personal and cultural self-​understanding;
narrative as a non-​subsumptive mode of encountering alterity; storytelling
as a way of shaping the narrative in-​between; narrative as cultivating our

[4] The Ethics of Storytelling

perspective-​awareness and capacity for perspective-​taking; and narrative as a
form of ethical inquiry. On the basis of these six aspects of what narratives
can do to us, ethically speaking, I propose a schematic map for exploring
and evaluating, in a differentiated manner, the ethical potential and prob-
lems of different storytelling practices. As a literary scholar, I mainly test
the model on literary narratives, but it is meant to be usable for the ethical
analysis of any narratives.
In reflecting on the sense of the possible, what interests me are real,
genuine possibilities, in contrast to merely logical possibilities.7 As Gary
Saul Morson writes, “the temporal world consists not just of actualities and
impossibilities but also of a third, in-​between category: real, though unac-
tualized, possibilities” (1998, p. 602). These real possibilities often go unac-
knowledged in actual worlds, and they are linked to our power to imagine
the “what-​ifs” and alternative courses of events that could make the world
utterly different. While many kinds of narratives can stimulate our capac-
ity to think beyond—​and resist—​the actual, this task is particularly impor-
tant for literature and other arts. As Jacques Rancière puts it, aesthetic
acts can function “as configurations of experience that create new modes
of sense perception” (2013, p. 3). In Gilles Deleuze’s terms, storytelling has
power to add to the real, for example by contributing to the invention of
new subjectivities and “a people to come” (2005, pp. 208, 264).8 Through
the exploration of human possibilities, narrative fiction opens up new per-
spectives on history, the everyday, and the yet-​to-​be. We engage with fic-
tional narratives as whole, embodied human beings with our own desires
and anxieties, values and beliefs, memories and fantasies. What is at stake
is not just an escape to the realm of the unreal but an exploration of the
possible. Through reading, we encounter what Ricoeur (1991a, p. 88) calls
imaginative variations of ourselves that allow us to explore—​as individuals
and communities—​who we are in relation to who we could be. Such explo-
ration cultivates our understanding of where we come from, where we are
now, and where we could go. This, in turn, affects who we in fact are.

TOWARD A NARRATIVE HERMENEUTICS

Hermeneutics refers to theoretical reflection on interpretation. It orig-

inates in the study of sacred and legal texts and in its modern form (at
the turn of the nineteenth century) came to signify theoretical reflection
on interpretation and understanding. In the early twentieth century, in
Martin Heidegger’s and Hans-​Georg Gadamer’s work, hermeneutics went
through an “ontological turn”: while nineteenth-​century hermeneutics

Introduction [5]
Another random document with
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its way. In Greenland, B. cordatus is very common in the dog, and probably also
in man, though few cases have been recorded. B. mansoni Cobb. (= B.
liguloides Leuck.) was, till recently, known only in the larval state from China and
Japan. Iijima, however, has found older specimens in the latter country. B.
cristatus Dav. is a species founded somewhat doubtfully on two fragments
found, one in a child, the other in a man, in France.

Occurrence of Cestodes in Domestic Animals.[103]—Among domestic

animals, the dog is, undoubtedly, the most frequently attacked by Taeniae. Six
species of Taenia (T. serrata, marginata, coenurus, echinococcus, krabbei, and
possibly T. serialis), Dipylidium caninum (the commonest form), Mesocestoides
lineatus, and three or four species of Bothriocephalus have been found in the
dog. The table of life-histories (p. 83) shows that sheep, rabbits and other
Rodents serve as the intermediate hosts, in which the cystic stages of the
species of Taenia are found. Hence the prevalence of T. serrata in a given
locality is connected with the abundance there of the rabbit and hare, in which
the larva (Cysticercus pisiformis) occurs. Bothriocephalus cordatus develops
from the young stage present in the fish which the Icelanders give to their dogs.
In Iceland and certain parts of Australia T. echinococcus infests one-third to one-
half the number of dogs examined; a fact connected with the frequency of
Echinococcus in man in these countries.

In sheep the most noteworthy and dangerous parasite is Coenurus cerebralis (or
the cystic stage of the dog-taenia, T. coenurus), which gives rise to the disease
known as "gid" or "staggers." It is found in various parts of the brain or spinal
cord, and the symptoms differ according to the position of the parasite. If this
presses upon one hemisphere the sheep describes circles and finally falls: if on
the optic lobes, the eyes are affected: if the pressure affects the cerebellum the
movements of the sheep are uncertain and incoordinated. Four or six weeks
after the appearance of the symptoms, death results from cerebral paralysis, or
from general debility, and the loss of sheep incurred by this disease (happily less
frequent in England than formerly) has been calculated by Youatt at a million for
France annually; at 35 per cent of the flocks for England in bad seasons; and
about 2 per cent for Germany. Besides sheep, which are most subject to "gid"
during their first year, various ruminants—Goat, Ox, Moufflon, Chamois, Roe,
Antelope, Reindeer, Dromedary—are attacked in the same way. A similar form,
Coenurus serialis Baill., is common in the wild rabbit in this country, and in
Australia in the hare and squirrel. It forms large swellings in the connective
tissue of various parts of the body, but usually does not affect the health of the
host. It is not known in what carnivore Taenia serialis Baill. normally occurs.
Experiments have, however, shown that it develops rapidly in dogs.
The preventive measures which are steadily diminishing the prevalence of the
Cestode parasites in man in some parts of Western Europe cannot be dealt with
here, but it may be noticed that the Jewish observance with regard to swine is
the surest preventive measure against taeniasis and trichinosis. Careful
inspection of meat and general cleanliness, are the leading measures that in
these hygienic matters secure the greatest immunity from disease.

Table of the Life-Histories of the principal Cestodes of Man and the

Domestic Animals.

Cestode. Final host. Larva. Intermediate host.

Rabbit, Hare, Mice
Taenia serrata Cysticercus
Dog (liver and
Goeze pisiformis Zed.
peritoneum)
Monkeys,
Cyst. tenuicollis Ruminants,
T. marginata Batsch Dog, Wolf
Rud. Ungulates (in
peritoneum)
T. saginata Goeze
(= Ox, Giraffe (in
Man Cyst. bovis Cobb.
T. mediocanellata muscles)
Küch.)
Pig, Man,
Cyst. cellulosae
Monkeys, Bear,
Rud. (? Cyst.
T. solium Rud. Man Dog, Cat, Black
acanthotrias
Rat (in various
Weinl.)
organs)
Cat and
other Cyst. fasciolaris Rat, Mouse, Bat
T. crassicollis Rud.
Felidae, Rud. (liver)
Stoat
Brain of Sheep,
Ox, Goat,
Dog, Arctic Coenurus
T. coenurus Küch. Dromedary,
Fox cerebralis Rud.
Camel,
Antelope, Horse
Coenurus serialis Rabbit (connective
T. serialis Baill. ? Dog
Baill. tissue)
T. echinococcus Dog, Dingo, Echinococcus Man, Monkeys,
v. Sieb. Jackal, polymorphus many
Wolf Dies.(incl. Carnivores,
Rodents,
 E. multilocularis Ungulates,
found in Man) Ruminants, and
Marsupials; also
in Turkey and
other birds
Moniezia expansa Sheep, Ox,
Unknown
Rud. Goat, etc.
Thysanosoma Sheep,
Unknown
fimbriata Dies. Cervidae
Stilesia
Sheep Unknown
globipunctata Riv.
Anoplocephala
Horse Unknown
perfoliata Goeze
Dipylidium caninum
Cysticercoid larva Body-cavity of
L. (= Taenia
Man, Dog, (Fig. 43), Trichodectes
cucumerina Bloch
Cat Cryptocystis and Pulex of
= T. elliptica
trichodectis Vill. Dog
Batsch)
Cercocystis Vill.
Hymenolepis [104] (develops
Mouse, Rat Usually absent
murina Duj. in parental
host)
H. nana v. Sieb. Man Unknown
Meal-moth, Asopia
H. diminuta Rud. (= (Pyralis)
Man,
Taenia farinalis; also
Mouse, Cercocystis Vill.
flavopunctata certain
Rat
Weinl.) Orthoptera and
Coleoptera
The Ostracods
Candona
Duck, rostrata and
Drepanidotaenia
Goose, Cercocystis Vill. Cypris
gracilis Zed.
Wild Duck compressa, and
also Cyclops
viridis
Cypris
D. anatina Krabbe Duck " " incongruens,
and also Perch
Cyclops
D. setigera Fröh. Goose " "
brevicaudatus
D. infundibuliformis Common
" " House-fly
Goeze Fowl
 Dicranotaenia Duck " " Cypris ovum
coronula Duj.
Davainea ? Limax cinereus,
Fowl " "
proglottina Dav. L. agrestis
D.
madagascariensis Children Unknown
Dav.
D. friedbergeri
Pheasant Unknown ? Ants
v. Linst.
Mesocestoides
Dog Unknown
lineatus Goeze
Probably first
enters an
Plerocercoid, i.e
Invertebrate
Bothriocephalus Man, Dog, solid, elongate
host, which is
latus Brems. ? Cat larva, with no
eaten by Pike,
bladder
Perch, Trout,
etc.

Fig. 40.—A, Stickleback (Gasterosteus aculeatus) infested by an advanced larva

of Schistocephalus solidus Crepl. B, The larva. All × 1½. (From specimens in
the Cambridge University Museum.)

Structure and Development of Cestoda.[105]—Of the unsegmented Cestodes,

Caryophyllaeus mutabilis, from the intestine of carp and other Cyprinoid fishes,
is the most easily accessible form. Triaenophorus nodulosus, which is very
useful for the study of the excretory system, occurs mature in the pike. In the
body-cavity of the Stickleback (Fig. 40) a large, broad, yellow worm may
sometimes be found, the larva of Schistocephalus solidus Crepl., which occurs
in the intestine of Terns, Storks, Mergansers, and other birds. Species of Ligula
are found in the same birds. The intestine of a Lophius or Cyclopterus ("lump-
fish") contains, usually, the early and intermediate stages of various Cestodes,
while the alimentary canal of Elasmobranchs often contain many peculiar
Tetrarhynchidae and other forms. For the study of development, the Taenia
anatina from the duck may be used. The ripe proglottides are collected, and the
eggs placed with Cypris ovum in an aquarium, with the probability that some of
the embryos will enter the Ostracod, and the peculiar Cysticercoid may be bred.
[106] Cysticercus pisiformis and Coenurus serialis, which occur commonly in
rabbits, are also suitable objects for examination.

A Cestode such as Echinobothrium (Fig. 36) is divisible into head and

proglottides. Moniez has suggested that the head is really the morphologically
hinder end of the body, in which case the formation of proglottides would closely
resemble the mode of segmentation of an Annelid larva. The close similarity,
however, between the Cysticercoid larva (Fig. 43, F) and the Cercaria of a liver-
fluke, seems to show that the anterior end is the same in both cases, and since
it bears the central part of the nervous system, we may reasonably call it the
"head." Moreover the hinder end of a Platyhelminth usually possesses the chief
excretory pore. Another difficulty is the determination of dorsal and ventral
surfaces. Authors are agreed,—on the analogy of Trematodes, in which the
testes are usually dorsal and the ovaries ventral,—that the dorsal and ventral
aspects of a Cestode are determined by the position of these organs, although
the often radially formed "head," the lateral or superficial position of the genital
apertures, and the variability of these features, render it a matter of considerable
doubt whether "dorsal" and "ventral" are more than useful conventional terms.
The suckers and hooks are borne on a muscular cap, the "rostellum," which is
only slightly developed in the Ichthyotaeniae. The body is solid, and is divisible
into an outer muscular coat—enveloped in a (possibly epidermal) investing
membrane—and an inner parenchymatous tissue containing the chief part of the
excretory, nervous, and reproductive systems. One or two pairs of longitudinal
excretory vessels are present, usually connected by transverse ducts and
opening by a single terminal pore. Occasionally a regularly paired arrangement
of lateral or secondary pores is present (Figs. 38 and 41, for.sec). Flame-cells
occur at the end of the fine tubules (Fig. 38), and the whole system is well
developed, but may undergo degenerative changes in the older proglottides.
The central nervous system varies according to the degree of differentiation of
the rostellum; and, owing to the difficulty of staining the nerves and the
contradictory statements of authors, we do not yet possess a fully reliable
account of the nervous system even of the commoner Taeniae. Free nerve
endings and other sensory terminations have been recently stated to exist in the
cuticle of Cestodes and Trematodes. If true, this would tend to show that the
parasitic mode of life of these animals demands a complex nervous system
comparable with that of the Turbellaria.
 Fig. 41.—Diagrammatic transverse section of Schistocephalus solidus Crepl., from
the Wild-duck, illustrative of the Cestodes with uterine aperture (uto). × 12.
cs, Cirrus-sac; for.sec, one of the paired lateral openings of the excretory
vessels; ln, longitudinal nerve; ov, ovary; ovd, oviduct; par.m,
parenchymatous muscles; r.sem, receptaculum seminis; sh.gl, shell-gland; te,
testes; ut, uterus; uto, uterine pore; vag, vagina; vd, vasa deferentia; yd, yolk-
duct; yg, yolk-glands (black); ♂ , male, ♀ , female genital aperture. (After
Riehm.)

The reproductive organs, unlike the preceding systems, are discontinuous from
one proglottis to the next. The male and female organs and their mutual
connexions, especially in the unsegmented Cestodes, may be compared in
detail with those of Trematodes, but the difference between the arrangement of
the generative organs of various Cestodes is very great.[107] The penis (Fig. 41,
cs) is evaginated through the male pore (Fig. 41, ♂ ), and inserted far into the
vagina (♀, vag) of the same or another segment of the tape-worm.

Fig. 42.—A, Free-swimming, six-hooked larva of Bothriocephalus latus Brems.

(the broad tape-worm of Man), still enclosed in a ciliated (possibly cellular)
double membrane or mantle. In this condition it may continue to live in water
for a week or more, but eventually throws off its ciliated coat (as in B) and
commences to creep about vigorously by the aid of its hooks, in search of its
first host, which is at present unknown. (After Schauinsland.) × 600.

From this fact and the anatomical relations of the vagina, it is becoming
increasingly probable that the so-called uterus of Trematodes is an organ
corresponding to the vagina of Cestodes, and not to the uterus of Cestodes. The
latter opens to the exterior in Schistocephalus, Bothriocephalus, and some other
Cestodes of fishes by a special pore (Fig. 41, uto). Through this, some of the
eggs (which in these genera give rise to ciliated larvae) are enabled to escape,
and need not wait for the detachment of the proglottis, as must happen in the
Taeniidae, where the uterus is closed. This uterus, a true physiological one, is
probably the homologue of the "canal of Laurer" ("Laurer-Stieda canal," or
"vagina") of Trematoda. The fertilised ovum and yolk are brought together into
the "ootype," where the shell-gland forms the egg-shell around them (Fig. 41,
sh.gl) and the egg is then passed into the uterus. The ovum segments to form a
minute six-hooked larva, which may (Bothriidae, Fig. 42) or may not (Taeniidae)
be ciliated. Thus in Taenia serrata the proglottides are shed with the faeces of
the host (dog), and they protect the young from the desiccating influence of the
surroundings. If inadvertently eaten by a rabbit along with herbs, the proglottis
and larval envelope are digested, and by its six hooks the tiny larva bores
through the gastric wall into the portal vein, and so into the liver. Here the hooks
are thrown off, and the solid mass of cells becomes vacuolated.

Fig. 43.—Stages in the development of Dipylidium caninum L. (= Taenia elliptica

Batsch, T. cucumerina Bloch), the commonest of the Dog-Taeniae; compare
Fig. 44. A, Six-hooked larva (now often spoken of as an "Onchosphaera"); B,
larva elongating; formation of a central lacuna; C, larva further advanced; D,
distinction between body and tail is visible; E, invagination of the rostellum is
commencing; F, Cysticercoid larva with four suckers, invaginated rostellum,
and excretory vessels. c, Calcareous concretions in cells of the larva; ex.o,
excretory aperture; ex.v, excretory vessels; inv, invagination commencing;
rost, rostellum; sc, suckers. (After Grassi and Rovelli; highly magnified.)

Fig. 44.—Schematic longitudinal sections through the larvae of Dipylidium

caninum L. All these stages are passed in the body-cavity of the Dog-flea
(Pulex serraticeps). (Compare Fig. 43 for further details.) A, Six-hooked larva
with developing rostellum (shaded) and suckers (black). In this species the
invagination (C, invag.) occurs after the formation of these organs, and not,
as in most Taeniae, before it. B, Invagination commencing; the hooks are
developing above the rostellum, while beneath it the nervous system (dotted)
is seen. C, The invagination has now carried the suckers inwards. The tail
has become distinct, and the whole larva at this stage is known as a
Cysticercoid. hk, Larval hooks; invag, mouth of the invagination; n, central
nervous system; rost, rostellum and hooks; sc, suckers, of which only two
 can be seen in a longitudinal section; four are really present. (After Grassi
and Rovelli.)

At one pole an invagination occurs, at the bottom of which the rostellum,

suckers, and hooks are gradually formed, but inside out as compared with the
head of the Taenia serrata. At this stage the larva (Cysticercus pisiformis) has
usually issued from the liver and attached itself to the omentum. The
invagination projects into the cavity of the bladder, within which a watery fluid
accumulates. Thus the "bladder worm" is formed, the head of which is
evaginated if the larva be introduced into the digestive system of a dog. The
bladder and neck of invagination are digested, while the head, protected by
these, remains, and forms the neck, from which the proglottides are afterwards
segmented off. In Taenia (Hymenolepis) murina the whole development may
take place in the parental host, the larva living in the villi, the adults in the cavity
of the same rat's intestine (Grassi). The different forms of Cestode larvae
depend largely upon the presence and degree of development of the caudal
vesicle or bladder, which in Scolex polymorphus (Fig. 38) (the young stage of
Calliobothrium filicolle Zsch.) is practically absent. If the bladder be small, the
larva is known as a Cysticercoid. For example, the common Dipylidium caninum,
which lives in the dog, has such a larva, the development of which is explained
and illustrated by Figs. 43 and 44. The bladder becomes exceeding capacious in
Coenurus and Echinococcus.

Table for the Discrimination of the more usual Cestodes of Man and
Domestic Animals.[108]

I. Scolex in most cases with hooks; uterus with a median and lateral
branches; yolk-glands simple, median; genital pore single; dorsal
excretory vessel narrower than the ventral, without a circular
commissural trunk; eggs without pyriform apparatus (processes of the
ovarian membrane)
Gen. Taenia L. (s. str.)
A. Genital ducts pass on the ventral side of the
nerve and of the two longitudinal excretory
vessels T. crassicollis Rud.
B. Genital ducts pass between the dorsal and ventral longitudinal vessels.
a. Nerve present on dorsal side of genital ducts.
α. Head armed T. solium Rud.
β. Head unarmed T. saginata Goeze.
b. Nerve on ventral side of genital ducts.
Dog-Taeniae[109]
Head armed; genital pore marginal and
 — Single
Many proglottides; strobila several centimetres long; small hooks
with guard.
Bifid hooks, which are
— 230µ-260µ long[110]; genital pore very
distinct T. serrata Goeze.
— 136µ-157µ long; genital pore not very
salient T. serialis Ball.
Entire large hooks, which are
— 180µ-220µ long; length of mature
segments double their width T. marginata Batsch.
— 150µ-170µ long; length of mature
segments treble their width T. coenurus Küch.
3-4 segments; a few mm. long T. echinococcus
v. Sieb.
— Double and bilateral Dipylidium caninum
L.
Head unarmed; two genital pores on ventral Mesocestoides
surface lineatus Goeze.
II. Scolex without hooks; one or two transverse uteri present; one or two
genital pores and yolk-glands, the latter never median; genital ducts
pass on the dorsal side of the nerve; eggs with pyriform apparatus.
A. One transverse uterus present.
a. Uterus with bullate egg-sacs; pyriform apparatus without horns; genital
ducts between dorsal and ventral vessels
Thysanosoma Dies.
α. Head large (1.5 mm.); square lobed testes in median field; posterior
margin of segments fimbriated; genital pore double
T. fimbriata Dies.
β. Head small; no fimbriae; pore rarely double T. giardii Riv.
b. Uterus without saccular dilatations; segments Anoplocephala E.
short, thick, and slightly imbricate Blanch.
Horse-Taeniae.
α. Head very large A. plicata
— No posterior lobes Zed.
— Four posterior lobes A. perfoliata
Goeze.
β. Head small, without posterior lobes A. mamillana Mehl.
B. Two uteri and two genital pores present; horns of pyriform apparatus
well developed; genital ducts pass on the dorsal side of the
longitudinal vessels
Moniezia R. Bl.
a. Interproglottidal glands [111] arranged in linear series (planissima
 group)
M. planissima S. and H. M. benedeni Mz. M. neumani Mz.
b. Interproglottidal glands saccular (expansa group)
M. expansa Rud. M. oblongiceps S. and H. M. trigonophora S. and H.
c. Interproglottidal glands absent (denticulata M. denticulata Rud.
group) M. alba Perr.
C. Uterus single or double, without spore-like egg-sacs; eggs with a single
shell; genital pores irregularly alternate; strobila narrow; testes absent
from median part of the field
Stilesia Raill.
a. A transverse uterus in middle part of median S. centripunctata
field; head 2 mm. diameter Riv.
b. Two lateral uteri in each segment; head less
than 1 mm. in diameter S. globipunctata Riv.
III. Scolex almost invariably provided with hooks; genital pores on left border
of segment; eggs with three shells but no cornua. Segments broader
than long; posterior angles salient.
Hymenolepis Weinl.
a. Scolex with a single series of 24-30 hooks, each 14-18µ long
H. nana v. Sieb. H. murina Duj.
b. Scolex very small, unarmed H. diminuta Rud.
IV. Scolex provided with two elongated muscular pits. Body segmented; three
genital apertures in middle of ventral surface
Bothriocephalus Rud.
Body 2-20 metres in length
B. latus Brems. B. cristatus Dav. (doubtful species). B. cordatus Leuck. B.
mansoni Cobb. (= B. liguloides Leuck.)

Classification of Cestodes.—The following classification, which, so far as the

Taeniidae are concerned, follows that employed by Railliet, Blanchard, and most
recent writers, includes only a few representative genera:—

1. Fam. Cestodariidae Mont. (Monozoa Lang).

Gen. Caryophyllaeus, Archigetes, Gyrocotyle, Amphilina.
2. Fam. Bothriocephalidae.
Sub-Fam. 1. Bothriocephalinae. Gen. Bothriocephalus, Schistocephalus,
Triaenophorus (= Tricuspidaria).
Sub-Fam. 2. Ligulinae. Gen. Ligula.
Sub-Fam. 3. Solenophorinae. Gen. Solenophorus, Duthiersia.
Sub-Fam. 4. Diphyllinae. Gen. Echinobothrium.
3. Fam. Tetrarhynchidae.
Gen. Tetrarhynchus.
4. Fam. Tetraphyllidae.
 Sub-Fam. 1. Phyllobothrinae. Gen. Phyllobothrium, Echeneibothrium,
etc.
Sub-Fam. 2. Phyllacanthinae. Gen. Calliobothrium, Anthobothrium, etc.
5. Fam. Taeniidae.
Sub-Fam. 1. Cystotaeninae. Gen. Taenia s. str.
Sub-Fam. 2. Anoplocephalinae. Gen. Moniezia, Thysanosoma, Stilesia,
Anoplocephala.
Sub-Fam. 3. Cystoidotaeninae. Gen. Dipylidium, Hymenolepis,
Drepanidotaenia, Dicranotaenia, Echinocotyle, Davainea.
Sub-Fam. 4. Mesocestoidinae. Gen. Mesocestoides, Dithyridium.
Sub-Fam. 5. Ichthyotaeninae. Gen. Ichthyotaenia, Corallobothrium.

CHAPTER IV

MESOZOA

DICYEMIDAE—STRUCTURE—REPRODUCTION—OCCURRENCE: ORTHONECTIDAE—
OCCURRENCE—STRUCTURE: TRICHOPLAX: SALINELLA.

The Mesozoa are an obscure group, the position of which in the animal kingdom
is still doubtful. The name Mesozoa was given to the group by its discoverer, E.
van Beneden,[112] as he concluded that they were intermediate between the
Protozoa and the higher Invertebrates. Recent authors, however, have called
attention to the resemblance existing between them and the "sporocysts" of
Trematodes, and though we still are ignorant of certain important points in their
life-histories, the Mesozoa are most conveniently (and probably rightly)
considered as an appendix to the Platyhelminthes.

Fig. 45.—A, B, C, Stages in the development of the vermiform larva in Dicyema

typus van Ben. (After Ed. van Beneden.) cal, "Calotte"; gc, germinal cell; n,
 nucleus of endodermal cell.

The animals composing this group are minute and parasitic, and are composed
of a small number of cells. They may be divided into two families: the
Dicyemidae, which occur exclusively in the kidneys of certain Cephalopods
(cuttle-fish); and the Orthonectidae, which live in the brittle-star Amphiura
squamata, the Nemertine Nemertes lacteus, or the Polyclad Leptoplana
tremellaris. In addition to the undoubted Mesozoa, certain anomalous forms—
Trichoplax adhaerens and Salinella salve—may be referred to this group.

Fig. 46.—Dicyemennea eledones Wag., from the kidney of Eledone moschata. A,

Full-grown Rhombogen with infusoriform embryos (emb); B, one of the latter
developing; C, fully formed; D, calotte, composed of the upper nine cells
shown in the figure. (After Ed. van Beneden and Whitman.) emb, Infusoriform
embryo; g, part of endoderm-cell where formation of these embryos is rapidly
proceeding; n.ect, nucleus of ectoderm-cell; n.end, nucleus of endoderm-cell;
p, "calotte."

Dicyemidae.—If the kidney of Eledone moschata, a Cephalopod common on

our south-western shores, be opened, a number of fine, yellowish, hair-like
filaments may be seen attached at one end to its inner surface, floating in the
fluid contained in the renal cavity. These may be Dicyemennea eledones Wag.,
although another form, Dicyema moschatum Whit., also occurs in the same
host. D. eledones (Fig. 46) is 7 to 9 mm. long, transparent, and is composed of
one large inner cell with a simple nucleus (Fig. 46, n.end), and of an outer layer
of ciliated cells, nine of which form the "calotte" or pole by which the animal is
attached. Within the former (endodermal) cell the formation of urn-shaped
"infusoriform embryos" takes place (B and C), the fate of which is not known, but
they are possibly the males. The individual which produces these larvae is called
a "Rhombogen." Other individuals which produce a more elongated larva
("vermiform larva," Fig. 45) are called "Nematogens," and Whitman has
described a third kind, which produce first infusoriform, and then vermiform,
larvae (Secondary Nematogens).[113]

The occurrence of the known species of Dicyemids (a group which has not been
investigated on our coasts) is as follows:—

Species. Host.
Dicyema typus van Ben. Octopus vulgaris.
D. clausianum van Ben. O. macropus.
D. microcephalum Whit. O. de Filippi.
D. moschatum Whit. Eledone moschata.
D. macrocephalum van Ben. Sepiola rondeletii.
D. truncatum Whit. Rossia macrosoma, Sepia elegans, S.
officinalis.
D. schultzianum van Ben. S. biseralis, Octopus vulgaris.
Dicyemennea eledones Wag. Eledone moschata, E. aldrovandi.
D. mülleri Clap. E. cirrosa.
D. gracile Wag. Sepia officinalis.
Conocyema polymorphum S. officinalis, Octopus vulgaris.
van Ben.

Orthonectida.[114]—Two species of Orthonectids are fairly well known,

Rhopalura giardii Metschn. from Amphiura squamata, and R. intoshii Metschn.
from Nemertes lacteus. The latter appears to be very rare, the former occurring
in 2 to 5 per cent of the number of hosts examined. The parasites occur in a
granular "plasmodium," the nature of which is uncertain. Metschnikoff regards it
as formed by the Orthonectids, and he considers that the cellular envelope, by
which it is sometimes enclosed, is developed from the neighbouring tissue of the
host. These granular, sometimes nucleated, plasmodial masses, which can
perform active amoeboid movements in sea-water, occur attached to the ventral
part of the body-cavity of Amphiura, and between the gut-branches and body-
wall in Nemertes. Should these hosts be infected by great numbers of the
Orthonectids, their sexual organs degenerate (as is the case with pond-snails
attacked by sporocysts[115]), and it is possible that the remains of these organs
may constitute the "plasmodia" (Braun).

Rhopalura giardii is of distinct sexes. Either males or females are found in one
Amphiura. Two kinds of females, flattened unsegmented, and cylindrical
segmented forms, are known. They consist of a ciliated ectodermal layer
enclosing an endodermal mass of eggs, between which is a fibrillar layer usually
considered to be of a muscular nature. The cylindrical female gives rise to eggs
which develop, probably exclusively, into males. The flattened female produces
eggs from which females alone arise, though the origin of the two forms of this
sex is not well ascertained. The males contain spermatozoa which fertilise the
eggs of the cylindrical female, whereas the ova of the flat form probably develop
parthenogenetically.

Fig. 47.—Rhopalura giardii Metschn. (from the brittle-star Amphiura squamata). ♂,

Full-grown male (× 800); ♀ 1, flattened form of female (× 510); ♀ 2, cylindrical
female (× 510). (After Julin.)

Trichoplax.[116]—This anomalous animal has only been found in aquaria,

originally in the marine aquarium at Graz by Schulze. It has the appearance of a
large, flattened, ciliated Amoeba (1.5-3 mm. in diameter), but is distinguished by
its structure. The upper surface is composed of a flattened epithelium. The lower
surface is made up of cylindrical ciliated cells, which pass imperceptibly into the
branched cells, embedded in a hyaline matrix, which compose the middle layer
of the body. No distinct organs, and beyond simple fission, no mode of
reproduction, have been observed. One species, T. adhaerens, is known, but
has never been met with in a free state.

Salinella.[117]—This is another aquarium-animal, found by Frenzel in the

Argentine, in an artificial saline solution with which he filled some aquaria. It
measures .2 mm. in length, and has a somewhat flattened, barrel-shaped
appearance. A single layer of ciliated cells bounds a central cavity opening at
each end. Fission, and conjugation followed by encystment, have been
observed. One form, S. salve, is known from salines taken from Cordova.
 NEMERTINEA

BY

LILIAN SHELDON
Staff Lecturer in Natural Science, Newnham College, Cambridge.

CHAPTER V

NEMERTINEA

INTRODUCTORY—EXTERNAL CHARACTERS—ANATOMY—CLASSIFICATION—
DEVELOPMENT—HABITS—REGENERATION—BREEDING—GEOGRAPHICAL
DISTRIBUTION—LAND, FRESH-WATER, AND PARASITIC FORMS—AFFINITIES

The Nemertinea form a compact group, the affinities of which have not been at
present clearly determined. Several species were mentioned and described in
the works of various naturalists during the latter half of the eighteenth century,
though their anatomy was not understood until considerably later. The first
mention of any member of the group was made by the Rev. W. Borlase in his
Natural History of Cornwall, published in 1758. He gives a short description and
a rough figure of Lineus marinus. From that time the increase in the knowledge
of the group was very gradual. New species were from time to time described,
but few of the descriptions could boast of much completeness, and many
erroneous views were held until comparatively recent years. The group was very
variously classified, but the general arrangement in early times seems to have
been to unite it with the Planarians. Valuable contributions to the history of the
development were made in 1848 and the few subsequent years by Desor,[118]
Gegenbaur,[119] Krohn,[120] and Leuckart and Pagenstecher[121]; and more
recently by Metschnikoff[122] and Salensky.[123]

Nemertines for the most part closely resemble one another in all essential
points, though they differ considerably in size, colour, and external details. They
vary in length from less than an inch to thirty yards, this extreme size being
attained by Lineus marinus.
 Fig. 48.—Lineus marinus Mont., from the living specimen in the coiled condition.
Plymouth. × 1. a, Anterior end; b, posterior end.

Fig. 49.—L. marinus, from the same specimen as Fig. 48, in the expanded
condition. a, Anterior end; b, posterior end.

Nemertines are common on the British coasts; about forty species have been
recorded from this area. On turning over a stone on a sandy or muddy shore in a
pool left by the receding tide, there may often be seen a coiled mass, having the
appearance of a uniform slimy string twisted into a complicated knot. If it be
carefully removed, the ends can generally be made out, one bluntly rounded and
the other slightly tapering (Fig. 48, a and b). Occasionally there may be seen
attached to the blunter end a fine thread, which moves about freely. This thread
may, by an instantaneous movement, be drawn into the body, no trace of its
existence being left except at the tip of the head, where a small pore is visible;
this is the orifice through which it was withdrawn. Shortly afterwards the thread
may be again shot out, the process being instantaneous and often accomplished
with great force. This thread (Fig. 50, p) is the proboscis, a very important and
characteristic organ in Nemertines.
Most Nemertines are marine; they are mostly indifferent to climate and to the
nature of the soil on which they live.

A few forms live on land (e.g. Tetrastemma agricola,[124] Geonemertes

palaensis,[125] and G. chalicophora[126]) or in fresh water (e.g. Tetrastemma
aquarum dulcium[127] and T. lacustre[128]) in various parts of the globe. There
are also parasitic forms; the best known of which is Malacobdella.[129] A pelagic
form, Pelagonemertes,[130] has been described by Moseley.

Fig. 50.—Side view of head of Cerebratulus (Micrura) tristis Hubr., showing the
everted proboscis. Naples. × 2. Drawn from a spirit specimen. c.s, Cephalic
slit; m, mouth; p, proboscis.

External Characters.—A typical Nemertine possesses an elongated worm-like

body (Fig. 49), which is usually thrown into numerous close coils (Fig. 48). In
section it may be either round or more or less flattened, with the lateral edges in
some cases quite thin and almost fin-like. One or two broad, flattened, and leaf-
shaped forms are known, but such a condition is exceptional, and the forms in
which it occurs have probably assumed it owing to the adoption of special
modes of life.

In the ordinary forms the posterior end of the body is pointed either bluntly or
sharply. The head is somewhat broader than the rest of the body, and often
assumes a spatulate form. Eyes (Fig. 51, e) are usually present either in one or
several pairs, or in symmetrically-arranged groups on each side of the head. The
mouth (Fig. 58, m) is situated near the front end of the body on the ventral
surface, and is usually rendered conspicuous by being surrounded by thick
tumid lips. It varies in form from being slit-like to elliptical. At the anterior end of
the body a small terminal pore occurs; this is the external opening of the
proboscis (Fig. 51, p.p).

Nemertines are often very diversely and brilliantly coloured, the hues most
commonly found being white, yellow, green, deep purple, and various shades of
red and pink. The ventral surface is usually paler in colour than the dorsal, and
the latter is often marked by longitudinal and transverse stripes (Fig. 59) in
contrasting colours.

The whole animal is enveloped in a layer of mucus, which sometimes becomes

hardened to form a tube, and this may be still further strengthened by an
admixture of particles of sand or earth.

The body is capable to a great extent of contraction and extension, a Nemertine

many inches long being apt, when irritated or alarmed, to contract itself to the
length of not more than half an inch. Hence, unless the animal is kept and
carefully watched, a very erroneous idea may be conceived as to its size.

Anatomy.—The body-wall consists of several layers (Fig. 52), which in a typical

highly-developed Nemertine are as follows:—

1. An external epidermic layer (ep), consisting of ciliated cells, among which are
placed numerous unicellular glands. These glands probably secrete the mucus
in which the Nemertine is usually enveloped; their contents when in the body are
very highly refracting. The epidermis rests on a basement membrane (b.m).

2. The two or three muscular layers, arranged as either an external circular and
an internal longitudinal, or an inner and an outer circular separated by a
longitudinal layer, or, as in the figure (c.m and l.m), two longitudinal separated by
a circular layer.

Fig. 51.—Amphiporus lactifloreus Johnst., drawn from the living specimen, from
the dorsal surface. Plymouth. × 2. e, Eyes; g, generative organs; n.g, nerve
 ganglion; p.p, proboscis pore; p, proboscis.

3. A fairly thick connective-tissue layer often found between the epidermis and
the muscles, into which latter it gradually merges (s.t).

The Digestive System.—The mouth is placed on the ventral surface near the
anterior end of the body (Figs. 53, 58, m). It leads into a straight oesophagus
(Fig. 53, oes), whence passes off the intestine (int), which is continued as a
straight non-convoluted tube to the anus (a), situated terminally at the posterior
end of the body. The intestine is thrown out throughout the greater part of its
course into paired lateral pouches.

Fig. 52.—Diagrammatic transverse section of a Nemertine (Schizonemertea)

through the middle region of the body. b.m, Basement membrane; c.m,
circular muscle layer; d.b, dorsal blood-vessel; ep, epidermis; g, generative
organs; int, intestine; l.b, lateral blood-vessel; l.m, longitudinal muscle layers;
n.c, lateral nerve-cord; n.l, nerve plexus; p, proboscis; p.s, proboscis sheath;
s.t, subcutaneous layer.

The alimentary canal is lined throughout by a ciliated epithelium. The

oesophagus has, in addition to this layer, an outer thick coat of large granular
cells, which probably have a glandular function.

Proboscis.—The most characteristic organ of the Nemertines is the proboscis

(Figs. 50, 53, 54). For many years its disposition and function were
misunderstood, and it was supposed to be a portion of the digestive system. The
proboscis, which lies dorsal to the alimentary canal, opens at the extreme
anterior end of the body by a small pore (Figs. 51, 53, 58). When retracted it is
sometimes considerably folded, and lies in a long pouch or sheath. To the walls
of this sheath it is attached round its anterior end; and strong muscles unite its
posterior extremity to the sheath a short distance from the posterior end of the
latter.
The proboscis seems to be exclusively a tactile and protective and defensive
organ, for which functions it is eminently fitted by the great ease and rapidity with
which it is everted or thrust out from the body. It consists of two distinct regions
(Fig. 54, g.p and m.p). In the retracted state the anterior part is a hollow tube
with very thick muscular walls made up of several layers. At the base of this part
in many of the Nemertines there is situated a sharp-pointed spine projecting
forward into the lumen, and several smaller stylets situated in a pair of vesicles
close to the base of the central spine. The position of the spines in the everted
proboscis is shown in Fig. 57. The posterior part of the proboscis is also a tube,
but instead of being muscular, its walls are glandular. This posterior glandular
part is never everted.

Fig. 53.—Diagrammatic drawing of a Nemertine from the dorsal surface to show

the position of some of the principal organs. a, Anus; c.s, cephalic slit; g,
generative organs; int, intestine with its lateral diverticula; m, mouth; n.c,
lateral nerve-cord; n.g, nerve ganglion; oes, oesophagus; p, proboscis; p.p,
proboscis pore; p.s, proboscis sheath.

The eversion is effected by a turning inside out of the anterior part of the
proboscis (Fig. 54). The process whereby the proboscis is retracted has been
very aptly compared to the effect which would be produced by the inversion of
the finger of a glove, accomplished by pulling a string attached to its tip on the
inside, the anterior muscular part being comparable to the finger and the
glandular part to the string. It is thus obvious that in the everted condition the