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Parasitism
The Diversity and Ecology of Animal Parasites
S E C O N D E D I T IO N

Reflecting the enormous advances made in the field over the past 10 years, this text synthesizes
the latest developments in the ecology and evolution of animal parasites against a backdrop
of parallel advances in parasite systematics, biodiversity, and life cycles. It has been thoroughly
revised to meet the needs of a new generation of parasitology students, whether their interest
is in ecology, conservation biology, evolution, immunology, or health sciences.
Balancing traditional approaches in parasitology with modern studies in parasite ecology
and evolution, the authors present basic ecological principles as a unifying framework to help
students understand the complex phenomenon of parasitism. Richly illustrated with over
250 figures, the text is accompanied by case study boxes designed to help students appreciate
the complexity and diversity of parasites and the scientists who study them. This unique
approach, which is presented clearly and with a minimum of jargon and mathematical detail,
encourages students to think generally and conceptually about parasites and parasitism.

Timothy M. Goater is Professor and former Chair in the Biology Department at Vancouver Island
University, British Columbia, Canada. During the past 20 years he has taught courses in
introductory biology, parasitology, ecological parasitology, invertebrate zoology, and ento-
mology. His research interests focus on the population and community ecology of parasites.

Cameron P. Goater is Associate Professor and former Chair in the Department of Biological
Sciences at the University of Lethbridge, Alberta, Canada. His parasitological research roots are
in the community ecology of helminths of waterfowl on the Canadian prairies, and over the
past 15 years he has taught courses in introductory biology, invertebrate biology, field biology,
and symbiotic interactions. His current research interests are in the experimental ecology of
helminth-host interactions.

Gerald W. Esch is Charles M. Allen Professor of Biology at Wake Forest University, North
Carolina, USA, where he has taught for 47 years. He is widely regarded to be one of the
world’s leading ecological parasitologists, and served as Editor of the Journal of Parasitology for
19 years.
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C:/ITOOLS/WMS/CUP-NEW/4272879/WORKINGFOLDER/GOAT/9780521190282TTL.3D iii [3–3] 8.8.2013 9:20PM

Parasitism
The Diversity and Ecology
of Animal Parasites
SECOND EDITION

TIMOTHY M. GOATER
Vancouver Island University, British Columbia, Canada

C A M E R O N P. G O A T E R
University of Lethbridge, Alberta, Canada

GERALD W. ESCH
Wake Forest University, North Carolina, USA
C:/ITOOLS/WMS/CUP-NEW/4272879/WORKINGFOLDER/GOAT/9780521190282IMP.3D iv [4–4] 8.8.2013 9:44PM

University Printing House, Cambridge CB2 8BS, United Kingdom

Published in the United States of America by Cambridge University Press, New York

Cambridge University Press is part of the University of Cambridge.

It furthers the University’s mission by disseminating knowledge in the pursuit of
education, learning, and research at the highest international levels of excellence.

www.cambridge.org
Information on this title: http://www.cambridge.org/9780521190282
© Cambridge University Press 2001, 2014
This publication is in copyright. Subject to statutory exception
and to the provisions of relevant collective licensing agreements,
no reproduction of any part may take place without the written
permission of Cambridge University Press.
First published 2001
Second edition 2014
Printed and bound in the United Kingdom by the MPG Books Group
A catalog record for this publication is available from the British Library
Library of Congress Cataloging in Publication data
Goater, Timothy M., 1959–
Parasitism : the diversity and ecology of animal parasites / Timothy M. Goater, Cameron
P. Goater, Gerald W. Esch. – Second edition.
pages cm.
First published as: Parasitism / Albert O. Bush . . . [and others], 2001.
Includes bibliographical references and index.
ISBN 978-0-521-19028-2 (hardback) – ISBN 978-0-521-12205-4 (paperback)
1. Parasites – Textbooks. 2. Parasitism – Textbooks. 3. Parasites – Ecology – Textbooks.
4. Biodiversity – Textbooks. 5. Parasitology – Textbooks. I. Goater, Cameron P.
II. Esch, Gerald W. III. Parasitism. IV. Title.
QL757.P287 2013
578.6′5–dc23
2013016194
ISBN 978-0-521-19028-2 Hardback
ISBN 978-0-521-12205-4 Paperback
Additional resources for this publication at http://www.cambridge.org/parasitism
Cambridge University Press has no responsibility for the persistence or accuracy of
URLs for external or third-party internet websites referred to in this publication,
and does not guarantee that any content on such websites is, or will remain,
accurate or appropriate.
C:/ITOOLS/WMS/CUP-NEW/4272879/WORKINGFOLDER/GOAT/9780521190282DED.3D v [5–6] 8.8.2013 9:48PM

We dedicate this book to our students,

past, present, and future.
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CONTENTS

List of Boxes page ix 5.5 Phylogenetic considerations

Foreword by John C. Holmes xi and classification 106
Preface and Acknowledgments xiii
6 Platyhelminthes: the flatworms 113
1 Introduction 1 6.1 General considerations 113
1.1 Encounters with parasites 1 6.2 Temnocephalidea 114
1.2 Scope 2 6.3 Udonellidea 115
1.3 Terminology 5 6.4 Aspidobothrea 115
1.4 Overview 10 6.5 Digenea 116
6.6 Monogenea 146
2 Immunological aspects of parasitism 16 6.7 Gyrocotylidea 152
2.1 General considerations 16 6.8 Amphilinidea 154
2.2 Vertebrate immunity 21 6.9 Eucestoda 154
2.3 Invertebrate immunity 31 6.10 Phylogenetic relationships and
2.4 Ecological immunology 34 taxonomic classification 171

3 Protista: the unicellular eukaryotes 40 7 Acanthocephala: the thorny-headed

3.1 General considerations 40 worms 179
3.2 Form and function 41 7.1 General considerations 179
3.3 Biodiversity and life-cycle variation 43 7.2 Form and function 179
3.4 Phylogenetic relationships and 7.3 Nutrient uptake and metabolism 186
classification 81 7.4 Development and general life cycle 188
7.5 Biodiversity and life-cycle variation 191
4 Microsporida: the intracellular, 7.6 Phylogenetic relationships and
classification 192
spore-orming fungi 86
4.1 General considerations 86
4.2 Form and function 86 8 Nematoda: the roundworms 199
4.3 Development and general life cycle 88 8.1 General considerations 199
4.4 Biodiversity and life-cycle variation 88 8.2 Form and function 199
4.5 Phylogenetic relationships and 8.3 Nutrient uptake and metabolism 208
classification 93 8.4 Development and general life cycle 209
8.5 Biodiversity and life-cycle variation 211
5 Myxozoa: the spore-forming cnidarians 96 8.6 Phylogenetic relationships and
classification 237
5.1 General considerations 96
5.2 Form and function 97
5.3 Development and general life cycle 97 9 Nematomorpha: the hairworms 244
5.4 Biodiversity and life-cycle 9.1 General considerations 244
variation 100 9.2 Form and function 244
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viii Contents

9.3 Development and general life cycle 246 14 Parasite biogeography and
9.4 Biodiversity and ecology 247 phylogeography 379
9.5 Phylogenetic relationships and 14.1 General considerations 379
classification 249 14.2 Historical biogeography 380
14.3 Ecological biogeography 386
10 Pentastomida: the tongue worms 252 14.4 Applied aspects of parasite biogeography
10.1 General considerations 252 and phylogeography 389
10.2 Form and function 252
10.3 Nutrient uptake and metabolism 256 15 Effects of parasites on their hosts: from
10.4 Development and general life cycle 256 individuals to ecosystems 396
10.5 Biodiversity and life-cycle variation 258 15.1 General considerations 396
10.6 Phylogenetic relationships and 15.2 Effects of parasites on host individuals 396
classification 261 15.3 Effects of parasites on host
populations 411
11 Arthropoda: the joint-legged animals 263 15.4 Effects of parasites on host communities
11.1 General considerations 263 and ecosystems 422
11.2 Crustacea 264
11.3 Chelicerata 289 16 Evolution of host–parasite interactions 432
11.4 Hexapoda 308 16.1 General considerations 432
11.5 Phylogenetic relationships and 16.2 Parasite-mediated natural selection and
classification 328 evolution 432
16.3 Genetic structure of parasite
12 Parasite population ecology 335 populations 441
12.1 General considerations 335 16.4 Introduction to host–parasite
12.2 Terminology and general approaches 336 coevolution 446
12.3 Introduction to parasite population
ecology 337 17 Environmental parasitology: parasites
as bioindicators of ecosystem health 459
13 Parasite community ecology 356 17.1 General considerations 459
13.1 General considerations 356 17.2 Parasites as effect indicators of pollutant
13.2 Introduction to parasite community stress 460
ecology 357 17.3 Parasites as environmental sentinels 470
13.3 The structure of parasite infracommun-
ities: restricted niches 361
Glossary 477
13.4 The structure of parasite communities:
species richness 368 Index 489
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LIST OF BOXES

1.1 A brief historical perspective of para- 7.4 Phylogeny of the Acanthocephala and
sitology: pioneering scientists and their the evolution of parasitism 195
ground-breaking parasitological 8.1 Evolution of nematode life-cycle plas-
discoveries page 4 ticity: developmental switching in
1.2 Parasite systematics: a phylogenetics Strongyloides spp. 213
primer 11 8.2 Entomopathogenic nematodes and their
2.1 Immunology: a terminology primer 18 microbial symbionts: tapping their
3.1 A plastid in apicomplexans: a promise chemotherapeutic potential 216
for new drug therapies? 43 8.3 Transmammary transmission in
3.2 Protists as hosts for microbial hookworms 225
symbionts 44 8.4 Wolbachia spp.: endosymbiotic bacteria
3.3 Giardiasis: epidemiology and patho- in filarid nematodes 234
genesis of beaver fever 48 8.5 Classification of the Nematoda 239
3.4 Pathogenesis, epidemiology, and diag- 10.1 Pentastomids: masters of
nosis of Chagas’ disease 53 immunoevasion 254
3.5 Relapse and recrudescence in malaria 68 10.2 Classification of the Pentastomida 258
3.6 Diagnosis of human malaria 74 10.3 Fossil pentastomes from the Cambrian
3.7 Classification of the Protista 79 era suggest an alternate phylogenetic
4.1 Nosema ceranae: an emerging pathogen hypothesis 259
of European honey bees 89 11.1 Astonishing metamorphosis in pen-
4.2 Classification of the Microsporida 94 nellid copepods and rhizocephalan
5.1 Pathogenesis and epizootiology of barnacles 280
whirling disease caused by Myxobolus 11.2 On acorns, ticks, and mice: the epi-
cerebralis 101 demiology of Lyme disease 297
5.2 Proliferative kidney disease and solving 11.3 A mite-y ecological and economic
the PKX organism mystery: an orphan problem: Varroa destructor of
parasite of bryozoans finds a taxonomic honey bees 303
home in the Myxozoa 106 11.4 Fly maggots as agents of lethal
5.3 Classification of the Myxozoa 108 parasitism 322
6.1 ‘Getting in’: host location, 11.5 Parasite’s web of death! Manipulation
recognition, and penetration by of spider web-building behavior by an
trematode miracidia 123 insect parasitoid 326
6.2 ‘Getting out’: the enigma of egg release 11.6 Classification of the Arthropoda 329
in the human schistosomes 139 13.1 Host immunity and parasite infracom-
6.3 Classification of the Platyhelminthes 162 munity structure: new evidence for an
7.1 Acanthocephalans: masters of pheno- old hypothesis nts’: parasite-induced
typic manipulation 180 fruit mimicry in Neotropical rainforests 366
7.2 Sexual selection in the Acanthocephala 188 15.1 ‘Berry ants’: Parasite-induced fruit
7.3 Classification of the Acanthocephala 194 mimicry in neotropical rainforests 407
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x List of Boxes

15.2 The politics of parasitism: sea lice, 16.2 Parasites, sex, and the Red Queen:
aquaculture, and the decline of salmon castrating trematodes maintain
populations in coastal British Columbia 412 sexual reproduction in New Zealand
15.3 Avian malaria determines the distribu- snails 449
tions, biodiversity, and community 17.1 Ribeiroia and the complexity of
structure of Hawaii’s native birds 423 amphibian deformities: a multidiscipli-
16.1 Parasite-mediated natural selection on nary approach to understanding the
the human HbS gene 433 possible link to eutrophication 466
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FOREWORD

The ability of parasites to cause disease has always further study in other biological specialties. Our
been an important reason to study them, and the courses, books, readings, and other materials used in
teaching of parasitology has almost always been our classes should be chosen to expose those students
stimulated by conditions conducive to disease, such as to the usefulness of parasites in investigations in their
war or climate change. Currently, zoonotic diseases chosen fields.
emerging from altered ecosystems, or carried by This book is the best I have seen for that purpose.
arthropod vectors spreading their ranges due to cli- The authors have provided a wide-ranging review of
mate changes, supply that stimulation. However, most the diversity of parasites, emphasizing those which
of us who teach, or have taught, parasitology have provide examples of the insights provided by the use of
chosen that topic because of the fascinating life cycles the new techniques or examples of how parasites can
of many parasites and their complex interactions with provide new and exciting insights into other aspects of
their hosts. Much of that fascination stemmed from biology. One of the best features of this book is that it
learning how parasites can affect the population emphasizes the complexity of host–parasite systems,
dynamics of their hosts, or the behavior of the hosts, or with full recognition that most of the outcomes are
even the evolution of their hosts. In addition, that markedly dependent on the conditions in which that
fascination was based on how much parasites could system is embedded. This emphasis on complexity
tell us about the life of their hosts, such as their diet, starts with a chapter on immunity, which is the best
travels, or evolution. Or even of the earth itself – some and most succinct coverage I have ever seen of those
of the earliest evidence for continental drift was the aspects of immunity that are important in host–
similarity in parasites of amphibians in Africa and parasite interactions. This emphasis is most apparent
South America. Examples of all of these influences are in the most integrative chapters – those on the influ-
provided in this book. ence of parasites on their hosts, and parasite evolu-
Many of the systems that parasitologists have used tionary ecology.
to show these fascinating features have become rela- This is the book I would have loved to have
tively easy to study due to new techniques, such as been available when I was teaching. But, of course, it
those in genomics and proteomics, which have pro- could not have been written then. Most of the more
vided new and more powerful ways to study system- provocative insights, and especially the evidence for
atics, evolution, and host–parasite relationships. This complexity and conditional outcomes of host–parasite
has attracted the attention of biologists with a wide encounters have come in the past two decades since
variety of backgrounds, so that much of the very I retired. The field of parasitology has become
interesting work done on host–parasite systems increasingly fascinating, and its implications for
recently has been done by those trained in other spe- other fields of biology more significant, in those two
cialties, such as ecology, behavior, neurophysiology, decades. Enjoy this book, as I have, and see where it
and evolutionary biology. Very few of the students in leads you.
senior-level parasitology courses will go on for further
study in parasitology, but many more will go on for John C. Holmes
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C:/ITOOLS/WMS/CUP-NEW/4273551/WORKINGFOLDER/GOAT/9780521190282PRF.3D xiii [13–16] 9.8.2013 11:13AM

PREFACE AND ACKNOWLEDGMENTS

In this second edition, we stay true to the philosophical reflecting developments in their systematics, and their
approach that was adopted in the first. Thus, we con- value as models in parasite ecology and evolution. By
tinue to see a need for a single text with dual focus on necessity, the ‘phylogenetic relationships and classifi-
the diversity and ecology/evolution of parasites. At the cation’ sections for all of the diversity chapters have
core, we feel that an ideal strategy for senior under- been updated, adopting the most current molecular-
graduate and beginning graduate students to under- based taxonomic schemes. The protist chapter in
stand and appreciate breakthroughs in parasite particular has been completely revised from the first
ecology is through a solid understanding of parallel edition, reflecting the monumental changes in protist
advances in parasite diversity, life-cycle variation, systematics. New text boxes that highlight key areas of
systematics, and functional morphology. By way of development, and the scientists behind them, are
example, we suggest that an understanding of the role integrated into each of these chapters. New life cycle
of falciparum malaria in determining the worldwide diagrams and dozens of new photographs and micro-
distribution of the human sickle-cell gene, and thus graphs have also been incorporated. A color plate
the role of parasites in mediating natural selection section has been added, showcasing dramatic photo-
(Chapter 16), comes from an understanding of life- graphs of parasites in or on their hosts.
cycle variation, functional morphology, and biodi- Armed with a solid background in parasite biodi-
versity of the apicomplexans (Chapter 3). Likewise, real versity, systematics, and functional biology,
understanding of the evidence in support of the para- Chapters 12–17 cover advances in the ecology and
site hypothesis for the evolution and maintenance of evolution of parasites. The titles and content of these
sexual reproduction in molluscs (Chapter 16) comes chapters have been completely revised from the first
from a detailed understanding of variation in life edition, reflecting in part, the interests and
cycles and life histories of the platyhelminths backgrounds of the new authors. Yet the substantial
(Chapter 6). This dual focus, under one cover, is the revisions also reflect the pace of development in
hallmark of this text. methodologies and in overall approaches that have
Our aim is to provide students with a synthetic matured the field over the past decade. While some of
understanding of the biodiversity, ecology, and evo- these developments have confirmed earlier ideas,
lution of animal parasites. Thus, throughout most of others have revolutionized our understanding of even
the text, we unabashedly take a parasite-centered view the most fundamental aspects of the parasitic way of
of the phenomenon of parasitism. Yet, we also aim to life. Thus, the incorporation of new model host–
provide insights on the nature of the host–parasite parasite interactions that are amenable to manipula-
interaction itself. It is for this reason that following a tion in the laboratory and field have provided key
brief introductory chapter, we provide an overview of insights into how parasite populations are regulated
vertebrate and invertebrate immunity, and the new and how they are distributed among hosts in space and
discipline of ecological immunology. We turn again time (Chapter 12). Studies at the community level
and again to the importance of fundamental immu- (Chapter 13) have also benefited from rigorous empir-
nological principles throughout the text. ical approaches involving key model systems where
There are now nine biodiversity chapters the composition of component species can be manip-
(Chapters 3–11). We have added chapters on the ulated. In Chapter 14, we see how advances in molec-
Myxozoa, Microsporida, and Nematomorpha, ular biology, genomics, and remote sensing have
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xiv Preface and acknowledgments

transformed our understanding of parasite biogeog- photographs, and micrographs for the new edition.
raphy and phylogeography. Coverage in Chapter 15 is Their generous contributions are acknowledged in the
focused on the diverse manner in which parasites can figure captions. John Sullivan is especially thanked for
affect the biology of their hosts, whether it is at the sharing several of his photographs from his excellent
level of the host individual, or on the structure of entire parasitological resource, A Color Atlas of Parasitology.
host ecosystems. Again, key advances stemming from Vancouver Island University is thanked for provid-
empirical, hypothesis-testing approaches involving ing Tim Goater the sabbatical and professional devel-
selected model systems have markedly advanced our opment funds that enabled this revision to take shape.
understanding of the magnitude of these effects, and Special thanks also to Mike Steele, David Marcogliese,
their underlying mechanisms. The focus in Chapter 16 and Herman Eure for providing office space during his
takes the next logical step, covering the manner in sabbatical, as well as Eric Demers, Larissa Nelson,
which parasites affect the evolutionary and coevolu- Wendy Simms, and Jane Watson for their enthusiastic
tionary trajectory of their hosts. We conclude the text encouragement throughout the project. Likewise, Cam
by summarizing the nature of the parasite/human/ thanks Dean Chris Nicol and Chair Brent Selinger for
habitat interface, and how the multidisciplinary field moral support and teaching relief during the peak
of environmental parasitology (Chapter 17) can assist phases of this revision, and colleagues Doug Bray,
in interpreting the nature of host–parasite interactions Doug Colwell, Andy Hurly, Joe Rasmussen, and Brian
in the face of anthropogenic change. Wisenden for their constant support. Cam also extends
As with all projects of this scope, this book is a thanks to Barb Johnson and staff at Waterton Lakes
collaborative effort. We extend sincere thanks to the National Park for access to their cabin during key
authors of the first edition, Al Bush, Jackie Fernández, writing phases. Most sincere thanks also to Lori Goater
and Dick Seed for their initial vision and dedication. for her monumental patience and support and to Ben
Several of their line drawings and photographs, and Ali for frequently reminding their dad, and their
incorporating the image-editing skills of Maggie Bush, uncle, that parasite ecologists come in all ages.
have been retained here. Numerous colleagues offered Our primary editor, Katrina Halliday and her assis-
valuable suggestions on specific sections/chapters, tant, Megan Waddington at Cambridge University
especially Carter Atkinson, Mark Blaxter, Katharina Press are thanked for all of their help addressing our
Dittmar, Eric Hoberg, Jens Høeg, Kayla King, David many queries and, especially for their patience and
Marcogliese, Jim Mertins, Beth Okamura, George devotion to seeing this project to its completion.
Poinar, John Webster, Chris Whipps, and Stephen
Yanoviak. We also appreciate the insightful comments Timothy M. Goater
John Holmes provided for several chapters. Several of Cameron P. Goater
our former students, especially Martin Anglestad,
Mellisa Beck, Aaron Jex, Chelsea Matisz, Phillip
Morrison, Vanessa Phillips, Brad van Paridon, and About this edition
Chris Whipps helped to review and edit chapters. Their
perspectives helped clarify and focus our efforts. The first Edition of our book was published in 2001. Al
This revision contains many new drawings, as well Bush, Jackie Fernández, and Dick Seed were
as new photographs and micrographs. Bill Pennell co-authors, along with myself. Sadly, Al died in 2006.
spent many hours of his retirement taking several new Further, Jackie stepped aside to raise two sons and Dick
photographs, as well as editing countless others. Doug retired from his faculty position at the University of
Bray and Brad van Paridon took several of the new North Carolina-Chapel Hill. As the only original
scanning electron micrographs. We thank our col- author that was still active professionally, it became
leagues for contributing extensive new data figures, my responsibility to recruit new co-authors. Given the
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Preface and acknowledgments xv

overall theme and target audience of the text, my Throughout the writing of both editions, I maintained
choice fell upon brothers, Tim and Cam Goater. Tim my duties as Editor of the Journal of Parasitology, as
was a former Ph.D. student of mine at Wake Forest well as my teaching. Mrs. Vickie Hennings, my Editorial
University, and Cam was a former Ph.D. student with Assistant for the Journal, continued her responsibilities
Clive Kennedy at the University of Exeter. Both while I was occasionally subsumed by the book. Cindy
brothers were mentored by Al Bush at Brandon Davis and Zella Johnson, both long-term secretaries for
University in Manitoba, Canada. Tim and Cam have the Department of Biology at Wake Forest University,
extensive experience teaching senior undergraduate are thanked for their help as well. I especially express
courses in parasitology and ecology, and Cam extends my appreciation to Ann for being such a marvelous
his teaching perspectives to the mentoring of graduate ‘listener’ and for her constant support from the book’s
students. Both have diverse and complementary inception.
research backgrounds that, together, span most areas
of modern parasitology. Gerald W. Esch
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C:/ITOOLS/WMS/CUP-NEW/4264071/WORKINGFOLDER/GOAT/9780521190282C01.3D 1 [1–15] 6.8.2013 8:15AM

1 Introduction

1.1 Encounters with parasites For the pet and livestock owner, parasite encounters
may have occurred when a veterinarian requested a
On a fateful spring day in a small northern Canadian fecal sample for diagnosis of eggs/larval stages of
town in the 1970s, two of the authors (the two that are intestinal worms. Perhaps, as a hunter or a fisherman,
related) of this text came upon a sickly red fox. you have queried the identity of that animal wriggling
Following some foolhardy thinking, they handled the in wild game meat or fish. In recent years, these com-
fox and carried it home. A few days later, health mon or at least dramatic parasites of humans, their
officials diagnosed the fox with rabies. To avoid the livestock, or their pets have been made famous in the
fatal consequences of the disease, the brothers required popular media (e.g., Zimmer, 2000a; 2000b), even
daily intramuscular injections of the prophylactic drug including in situ video footage on You-Tube and on
that was used at the time. We recall the episode with prime-time television shows.
memories of pain, dismay from parents, and ruthless As undergraduate students, your first encounters
teasing from our friends. And so goes our introduction with parasites and with the phenomenon of parasitism
to the world of parasites. So too goes our introduction likely occurred in your introductory courses. At each of
to the phenomenon of parasitism. Readers might our universities, majors in many of the life sciences
envision two teenagers discussing how their predica- require an introductory course that describes the
ment arose: How did that fox get infected? Why was diversity and unity of the Tree of Life. In a course such
the fox population, but not the racoon population, so as this, it would be impossible for instructors to sample
heavily infected that year? How does the virus migrate that diversity without covering examples of parasites,
from the site of a wound, to the brain, to saliva? How, although coverage is likely restricted to key human
and why, does it transform a normally secretive and parasites – a protist, a fluke, a cestode, and so on.
nocturnal animal into one that is aggressive and Likewise, our majors are required to take an introduc-
diurnal? There are obvious parallels between these tory course that covers basic principles of ecology and
early queries and modern questions associated with evolution. One encouraging sign of the expanding
host specificity, parasite site selection, the geograph- reach of studies on the phenomenon of parasitism is its
ical mosaic of coevolution, and mechanisms of alter- increased coverage in mainstream ecology and evolu-
ations in host behavior. tion texts (e.g., Begon et al., 2006; Freeman & Herron,
We hope that your introduction to parasites was (is) 2007). Nonetheless, time constraints in a single-
not as dramatic, or as dangerous, as it was for two of semester introductory course likely limit coverage of
us! Indeed, for many, initial exposure to the concept of examples involving parasites.
parasitism likely originated from media reports that In the chapters that follow, our coverage assumes
describe human mortality and morbidity caused by that you have encountered parasites, both anecdotally
diseases such as malaria, or other parasitic diseases and academically. Thus, we assume that senior stu-
that are so common in developing countries. Or, per- dents in the life or medical sciences have an appreci-
haps you have heard about certain parasites that are ation for basic principles of classification and
transmitted via ingestion of untreated water, or phylogeny and an appreciation for variation in the life
swimming in it, or from eating poorly cooked meat. cycles and general biology of a few animal parasites.
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2 Introduction

We also assume a general understanding of basic histories, and in their strategies for nutrient acquisi-
concepts of ecology and of the fundamental and uni- tion. Our intent through this section is to provide
fying nature of evolutionary processes. Although we insight on ‘the art of being a parasite’, a phrase
do not emphasize the mathematical underpinnings of coined by Claude Combes (2005) to describe the man-
host–parasite interactions, we do assume that senior ner in which parasites of all types solve the unifying
students have a numeracy background consistent with problems of entering a host (‘getting in, or on’),
introductory courses in calculus, linear algebra, and/or remaining in a host (‘staying in’), and reproducing
statistics. We do not assume a strong background in (‘getting out’). Our taxonomic scope is broad, with
immunology or pathology. emphasis on the traditional protists and ‘worms’ but
also on lesser-known groups such as the microspori-
dians, myxozoans, hairworms, and pentastomes.
1.2 Scope Much of our coverage through this section distils
material that is covered in parasitology texts
Our first aim is to provide students with an apprecia- (e.g., Noble et al., 1989; Kearn, 1998; Roberts &
tion for the biodiversity of animal parasites. From the Janovy, 2009). However, relative to these excellent
perspective of understanding our planet’s biodiversity, texts, we restrict our taxonomic scope to key families
and understanding factors leading to its loss, an or orders within each group, and we emphasize those
appreciation for the diversity of parasites is important. groups that provide models for enquiries on the ecol-
Parasitism is recognized as the most common strategy ogy and evolution of parasitism that we cover in later
used by animals to obtain nutrients (Price, 1980; de chapters.
Meeus et al., 1998; de Meeus & Renaud, 2002), ubiq- Our second aim is to develop in students an appre-
uitous across the Tree of Life. Poulin & Morand (2004) ciation for the phenomenon of parasitism. And from
consider that there have been at least 60 independent our perspective, we view the core of the phenomenon
evolutionary transitions from free-living to obligately to be ecological in nature. Thus, whether interest is in
parasitic animals. Estimates of the overall biodiversity understanding the innumerable rates that define the
of parasites vary depending on how inclusive we outcome of host–parasite relationships (e.g., rates of
define ‘parasite,’ but approximately 30–50% of exposure to infective stages, rates of within-host
described animal species are parasitic at some stage migration, rates of parasite-induced host mortality,
during their life cycle (Price, 1980; Poulin & Morand, rates of dispersal, and so on), or in the dynamics of the
2004). Given that virtually all metazoan species are molecular exchange that occurs at the host–parasite
infected with at least one species of parasite (most interface, or in the global distribution of parasites,
species contain many more), that all viruses and many basic ecological principles can be applied to help
prokaryotes and fungi are parasitic, and that we focus our thinking about host–parasite interactions. It
underestimate the biodiversity of groups such as is this perspective that lies at the roots of ‘parasite
nematodes and mites (see Chapters 8 and 11), these ecology’ as a subdiscipline within the ecological sci-
rough estimates are undoubtedly low. Clearly, knowl- ences. These roots were developed and formalized
edge of parasite biodiversity equates to knowledge of some 30–40 years ago following the coincident pub-
key branches of the Tree of Life. lications of seminal works by empirical field biolo-
The biodiversity section of the text (Chapters 3–11) gists (Kennedy, 1975; Price, 1980) and quantitative
provides an overview of the main taxa of protist and ecologists (Crofton, 1971; Anderson & May, 1979).
metazoan parasites. Our focus is on characterization of The dynamic tension between their alternative per-
key features that define each group, followed by cov- spectives continues to richly define the direction of a
erage of how natural selection has shaped variation in field that is now seeing an unprecedented level of
their morphologies, in their life cycles and life activity.
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Scope 3

Over the 10 years since the first edition of our text wide range of host–parasite interactions. Our
was published, key advances have been made in vir- overall approach through this section is empirical,
tually all areas of parasite ecology and evolution. rather than conceptual. Thus, we develop our argu-
These include the epidemiology of wildlife disease ments based primarily upon observations from field-
(Hudson et al., 2001), parasite phylogeny and phylo- based and laboratory-based experiments, although
genetics (Brooks & McLennan, 2002), parasites and we incorporate key results from field surveys of
host behavior (Moore, 2002), parasite biodiversity particular hosts when warranted. Although we
(Poulin & Morand, 2004), evolutionary ecology (Frank, cover the mathematical and conceptual framework
2002; Poulin, 2007; Thomas et al., 2009; Schmid- of certain areas of enquiry, our perspective is
Hempel, 2011), and parasite biogeography (Morand & empirical and rests strongly on the background that
Krasnov, 2010). Two texts that synthesize general we developed in the biodiversity section of the text.
advances in parasite ecology and evolution, one Readers seeking advances in more quantitative
from an empirical standpoint (Combes, 2001) and one aspects of parasite ecology and epidemiology should
from a conceptual standpoint (Poulin, 2007), are consult Hudson et al. (2001), Ebert (2005), or
especially notable. Over the past 10 years or so, the Schmid-Hempel (2011).
new subdisciplines of ecological immunology, land- Despite the enormous strides made in methods and
scape epidemiology, emerging diseases, and environ- approaches, modern studies in parasite biodiversity
mental parasitology have blossomed as exciting ‘hot and ecology continue to be influenced by traditional
topics.’ This surge in interest is partly due to the approaches in parasitology, in which parasites of
explosion in the use of modern molecular methods, humans and their domesticated animals have
enabling advances in our understanding of parasite played a key role. Throughout the biodiversity section
biodiversity, phylogenetics, population genetics, and of the text, we retain some of that traditional cover-
host–parasite coevolution that would have been age. We do so because the history of discovery in
unthinkable even 10 years ago. Yet, the pace of parasitology provides the roots of current enquiry,
advance is also due to the rapid increase in the use of and is itself a fascinating story of human endeavour
experimental model systems to test key hypotheses (Box 1.1). For further account of key historical
regarding the ecology of host–parasite interactions. developments in parasitology, readers are directed
While traditional model systems involving laboratory to Esch (2007). We also retain some emphasis on
rats and mice as hosts continue to provide important select human parasites because the discovery in
insights, major recent advances have arisen from these groups has provided unmatched opportunities
models involving parasites of hosts such as stickle- for increased understanding of ecological and
backs, guppies, water fleas, songbirds, and wild small evolutionary phenomena. For instance, results of
mammals. Indeed, we view the multidisciplinarity studies on the interaction between falciparum
arising between parasitologists and ecologists, so long malaria and the gene responsible for sickle-cell
called for by the fathers of parasite ecology, that is anemia provide one of the best examples of
perhaps most responsible for the unparalleled advan- parasite-mediated natural selection (Chapter 16).
ces we are currently witnessing in the field (review in Yet, this example stems from years of dedicated
Poulin, 2007). effort that enabled detection of the single amino acid
Our aim in this section of the text is to provide substitution that alters the structure of the hemoglo-
an overview of modern parasite ecology, evolution, bin molecule. We cover similar examples throughout
and coevolution. In this edition, we update our ear- the text, not necessarily in the context of human
lier treatment by taking into account results origi- disease, but in the context of central questions
nating from modern advances in molecular regarding the ecology and evolution of host–parasite
methodologies and from experimental models on a interactions.
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4 Introduction

Box 1.1 A brief historical perspective of parasitology: pioneering scientists

and their ground-breaking parasitological discoveries

Sometime around 1500 BC, an Egyptian physician assembled a large body of medical infor-
mation regarding the diagnosis and treatment of diseases known to occur at the time. Written in
hieroglyphics on papyrus and sealed in a tomb, it was discovered in 1872. It was translated by
Georg Ebers in 1873, becoming known as the Ebers’ Papyrus among Egyptologists. This volume
became an invaluable source that documented the medical profession that existed in the ancient
world.
Based on these writings, we now know that early Egyptian physicians were aware of at least
two parasitic helminths infecting humans. One of these was a nematode, probably Ascaris. The
recommended treatment for infection by this apparently common worm included turpentine and
goose fat! The second parasite was a tapeworm, most likely Taenia saginata, for which a special
poultice applied to the abdomen was the recommended treatment. Whereas the digenean,
Schistosoma haematobium, was not described per se, the bloody urine produced by this parasite
was a well-known symptom. Moreover, eggs of this worm have since been identified in
mummies from the thirteenth century BC (Grove, 1990). It is also possible that the hookworm
nematode, Ancylostoma duodenale, was present based on descriptions in the Ebers’ Papyrus of a
‘deathly pallor’ in some patients, a condition that may have been caused by hookworm-induced
anemia.
Concurrently, another group of ancients was acquainted with a number of helminth parasites
in the Nile Valley. Thus, for example, consider Numbers 21:6–9, which refers to ‘the Fiery
Serpent,’ now recognized as the nematode Dracunculus medinensis. When the Israelites mis-
behaved during their trek out of Egypt, they were directed by God, through Moses, to “make a
serpent of brass and put it upon a pole.” And, “when he beheld the serpent of brass, he lived.” This
treatment is still used today, that is, to remove the large female nematode from its subcutaneous
site of infection, and then to slowly twist the parasite on a stick, until it is removed intact. Many
feel the Hebrew law against eating the flesh of an ‘unclean’ animal, e.g., a pig, can be traced to
the nematode Trichinella spiralis or the cestode Taenia solium. On the other hand, the Talmud (a
sacred Jewish book), written in AD 390, referenced the hydatid cysts of the tapeworm
Echinococcus granulosus, indicating that they were not fatal.
Periodic fevers due to malaria were mentioned in Chinese writings from around 2700 BC and
in every civilization since. Hippocrates (460 BC–377 BC) provided the earliest detailed descrip-
tion of these periodic fevers. Both Hippocrates and Aristotle (383–322 BC) were aware of ‘worms’
and refer to cucumber and melon seeds in the ‘dung’ of humans. Both references are probably to
the gravid proglottids of Taenia saginata. Galen (AD 130–200) referred to the intestinal phases of
what were probably Ascaris lumbricoides and Enterobius vermicularis, saying that the former
worms preferred the upper portion of the gut whereas the latter were closer to the anus. He found
that tapeworms, on the other hand, were found throughout the length of the intestine. These
observations, so long ago, may be the first reference to the site specificity exhibited by parasites.
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Terminology 5

Box 1.1 (continued)

The earliest use of the microscope, by Antony von Leeuwenhoek in the seventeenth century,
provided a phenomenal breakthrough for the biological sciences and parasitology. He actually
observed, and described, the unicellular protozoan parasite Giardia lamblia, apparently from his
own feces! Also in the seventeenth century, several scientists prepared detailed drawings of a
number of parasitic helminths. One father of parasitology was Francesco Redi (1626–1697), who
not only determined that mites could make one itch, but apparently was also an inveterate collector,
describing some 108 species of parasites. Perhaps Redi’s greatest contribution was that he showed
that parasites produce eggs, dispelling the widespread myth that parasites developed through
spontaneous generation. The idea of spontaneous generation persisted for many years, however,
and it took Louis Pasteur’s now classic experiments in nineteenth-century Paris to quash the notion.
The late nineteenth and early twentieth centuries were times of major discoveries dealing with
some of the protist and helminth scourges of humans, including Wuchereria bancrofti as the
causative agent for elephantiasis and tsetse flies as the vectors for African trypanosomiasis.
Ronald Ross, while working in India in 1897, demonstrated that mosquitoes vectored
Plasmodium, winning the Nobel Prize for physiology in 1902. At the turn of the century, Paul
Erlich described the first chemotherapeutic agents for African trypanosomiasis and syphilis.
With this discovery, he hypothesized that organic molecules with selective toxicity to parasitic
organisms would be found. For this, he is considered the father of modern chemotherapy.
Between 1907 and 1912, Carlos Chagas determined the identity of trypanosomes that cause
Chagas’ disease and worked out the parasite’s life cycle in the reduviid bug vector. In the early
1880s, Algernon Thomas and Rudolph Leuckart independently completed stages in the life cycle
of the liver fluke, Fasciola hepatica, including detailed descriptions of the swimming behaviors
of the ‘embryos’ that hatched from eggs, their penetration into snails, and their subsequent
intramolluscan development. Thomas and Leuckart and others will be remembered for their
many contributions (review in Esch, 2007), and paving the way for all those who resolved so
many other parasitological mysteries.

1.3 Terminology discussion, we might offer a classic dictionary

definition. Webster’s Third New International
We often start our courses with a request for students Dictionary of the English Language defines ‘parasite’
to define ‘parasite.’ This is always an interesting and as follows:
engaging exercise. Often, the discussion rapidly
An organism living in or on another living organism
deteriorates into a mix of vague terminology, exam-
obtaining from it part or all of its organic nutrient, and
ples, and counter-examples. Are mosquitoes and
commonly exhibiting some degree of adaptive structural
vampire bats parasites? Are leeches parasites? Is my
modification – such an organism that causes some degree
brother a parasite? Is a fetus a parasite? To direct the
of real damage to its host.
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6 Introduction

It is here where some students may voice discomfort,

especially those with an ecological background. In this
SYMBIOSIS
characteristic definition, vague and unquantifiable
terms such as ‘part,’ ‘some,’ and ‘damage’ are prominent.
In our courses, we do not offer a solution to the funda-

Dir
raction

ec
mental vagueness that characterizes the definition of

ion

t tro
parasite. Nor do we do so in this text (for complete

t
rac

ph
ophic inte
discussions, see Zelmer, 1998; Combes, 2001). In his

ic
inte

int
influential text on coevolution, Thompson (1994)

era
hic
trop

c
emphasizes that all definitions dealing with interspecific

Indirect tr

tio
n
No
interactions are necessarily vague. From our perspective,

Benefit
Harm
we consider that a parasite has a metabolic commitment
to its host, has evolved morphological and physiological
adaptations to living in, or on it, and has the potential to re
Pho sy mensalis ploitation tualism
om Ex Mu
decrease host fitness. As we have indicated previously,

m
C
our focus is on the familiar parasitic protists, worms, and
arthropods, although we extend our coverage to include
lesser-known taxa because they provide splendid mod-

Se
els in parasite ecology. The extent to which groups such d
ille

ldo
as phytophagous insects, molecular parasites, blood- sk
ay

m
l w

kil
sucking leeches and flies, and brood parasites (e.g., A

led
cuckoos) apply to our coverage of animal parasitism,
Many hosts

Many hosts
One host

One host
provides an excellent topic for discussion in our classes,
but they lie outside the scope of this text.
Parasitism is one of at least four complex symbiotic dat
Pre or rasitoid opredato Parasite
relationships. Symbiosis, a term coined by de Bary in Pa icr
M

r
1879, literally means ‘living together of differently
named organisms.’ It describes the relationship in which
a symbiont lives in, or on, another living host. Symbiotic
interactions, or ‘symbioses’, include a tremendous vari- Fig. 1.1 ‘Parasitism’s place’ within the context of symbiotic
ety of intimate partnerships in nature. In the broadest relationships. This is one way of looking at parasitism and
sense, there is no implication with respect to the length or it is based, initially, on trophic relationships, followed by
outcome of the association, nor does it imply physio- ‘harm,’ and finally, quantity of hosts involved. The final
logical dependence or benefit or harm to the symbionts criterion, number of hosts attacked, is meaningful only if
restricted to a single life history stage. For example, adult
involved in the partnership. Given such a broad defini-
parasitoids may parasitize many host individuals but their
tion of symbiosis, a functional separation can be made in larvae live in, and consume, only a single individual.
relation to the feeding biology of one or both of the Likewise, a typical helminth parasite may have both
symbiotic partners, as well as the degree of host exploi- intermediate and definitive hosts, but each life history
tation. Thus, categories of symbiosis relate to trophic stage will infect only a single host individual. These
categories are arbitrary and, often, there is considerable
relationships, and if and how energy is transferred
overlap between many of the relationships. (Figure
between the partners. Such categories are best viewed as courtesy of Al Bush.)
a continuum with overlapping boundaries (Fig. 1.1).
If there is no trophic interaction involved in the sym-
biotic interaction, then the relationship is called
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Terminology 7

‘phoresy’ (Fig. 1.1). In this case, the symbiont (=phoront)

merely travels with its host; there is no metabolic com-
mitment by either partner. Protists or fungal microbes
that are mechanically carried by insects are examples of
phoretic associations. Similarly, even though whale and
turtle barnacles are often described as ectoparasites, there
is no metabolic commitment. Functionally, they are
phoronts. Phoresy grades into ‘commensalism’, a sym-
biotic interaction that implies a trophic relationship
between the partners (Fig. 1.1). Commensalism means
‘eating at the same table.’ Here the benefit gained is
unidirectional. The smaller commensal partner typically
benefits via food transfer and increased dispersal oppor- Fig. 1.2 Scanning electron micrograph of the mutualistic
tunities, while the host is neither harmed nor benefited. hypermastigote flagellate Trichonympha campanula from
When sharks feed on large prey, they scatter fragments of the intestine of a termite. Another, much smaller flagellate
food that are made available to remoras. Yet, some Streblomastix sp. (arrows) is also present. (Micrograph
courtesy of Ron Hathaway.)
remoras also feed on ectoparasites of their shark hosts,
implying an indirect metabolic linkage. Commensalism
therefore grades into ‘mutualism’ in many cases Several major categories of this kind of exploitation
(Fig. 1.1). Many mites are commensals, hitching a ride can be recognized, based primarily on the number of
and sharing food with hosts as diverse as insects and hosts attacked by the symbiont and the subsequent
molluscs to birds and mammals. fate of the organism assaulted (Fig. 1.1). If more than
When there is a direct transfer of energy between the one organism is attacked, but typically is not killed,
partners, the interaction may be either mutualistic or then the aggressor is called a ‘micropredator’.
exploitative (Fig. 1.1). Obligate mutualists are metabol- Hematophagous organisms such as mosquitoes, and
ically dependent on one another. A classic example of some leeches and biting flies, for example, are con-
an obligatory mutualism is the diverse microfauna of sidered micropredators, taking frequent blood meals
protists and prokaryotes in the intestines of wood- from several hosts. Some micropredators are often
eating termites. A single species of flagellated protist, considered as ectoparasites, e.g., leeches. If more than
Trichonympha campanula (Fig. 1.2), may account for up one organism (considered as prey) is attacked and
to one-third of the biomass of an individual termite. always killed, then the aggressor is considered a pred-
These flagellates produce enzymes that digest cellulose, ator. If only one specific host is attacked and is almost
enabling the host to survive on a diet of wood. The always killed, then the aggressor is usually referred to
mutualistic relationship between ruminant mammals as a ‘parasitoid’, most of which are wasps and flies.
and the ciliated protists and microbes in their stomach is If only one host is attacked, but typically is not killed
similar. The biochemical complexity of these, and many outright, the aggressor is a parasite (Fig. 1.1).
other mutualistic associations found throughout nature, ‘Endoparasites’ include those that are confined within
is the product of a long coevolutionary history between the host’s body. They include the protists, microsporidian
the partners. Such coevolved mutualisms are regarded and myxozoans, as well as the ‘worm’ parasites such as
as being creative forces in the adaptive radiations of flukes, tapeworms, acanthocephalans, and nematodes. A
many taxa (Thompson, 1994; Price, 1996). variety of holdfast adaptations often serve to anchor
In many exploitative interactions, however, benefit these endoparasites to specific sites within their specific
is unidirectional and, moreover, some form of disad- hosts. The holdfasts of elasmobranch cestodes, for
vantage, or harm, is the outcome for the other partner. example, are often exquisitely adapted to match the
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8 Introduction

Fig. 1.3 Scanning electron micrographs illustrating the elaborate holdfasts of host-specific tapeworms of elasmobranchs.
(A) Scolex of the trypanorhynch cestode Paragrillotia similis from the spiral intestine of the Atlantic nurse shark
Ginglymostoma cirratum; (B) scolex of the rhinebothriidean cestode Rhinebothrium megacanthophallus from the spiral intestine
of the freshwater whipray Himantura polylepis. (Micrographs courtesy of Janine Caira (A) and Claire Healy (B).)

microstructure of the intestines of their specific elasmo-

branch hosts (Fig. 1.3). Parasites found on the surface of
the host’s body are called ‘ectoparasites’. Most parasitic
arthropods and monogeneans are ectoparasitic. There are
also some parasites that are classified as ‘mesoparasites’
(Kabata, 1979). The pennellid copepods, for example, are
endoparasitic in the sense that they have elaborate
holdfasts that extend deeply into their host’s tissues.
However, their highly modified trunk regions and egg
sacs extend outside the host (Fig. 1.4; Color plate
Figs. 4.2, 4.3).
Anderson and May (1979) went further, highlight- Fig. 1.4 Host and site specificity exhibited by the
ing key differences within groups of parasitic organ- mesoparasitic copepod Phrixocephalus cincinnatus attached
isms. ‘Macroparasites’ are large (usually visible to the to the eye of an arrowtooth flounder Atheresthes stomias.
eye), have generation times approximating those of A metamorphosed female develops an elaborate holdfast,
penetrating deeply into the eye of the specific fish host, while
their hosts, generate a low-to-moderate immune
the egg sacs and trunk region extend out of the eye.
response, and the pathology they cause to their hosts is (Photograph courtesy of Dane Stabel.)
tied to the numbers of parasites present. These are
typically the classical ‘worms’ (trematodes, cestodes, Varroa destructor, provides another example (Fig. 1.6).
and nematodes) and the arthropods, such as copepods, ‘Microparasites’ are much smaller (typically micro-
fleas, lice, and mites. They can be endoparasitic or scopic), have generation times much shorter than their
ectoparasitic. The nematode, Heligmosomoides polyg- hosts, are capable of asexual replication within their
rus (Fig. 1.5), is an example of an endoparasitic mac- hosts, and typically induce strong acquired immunity
roparasite infecting mice. The ectoparasitic mite, in recovered and re-exposed hosts. They can be
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Terminology 9

Fig. 1.7 Scanning electron micrograph of the trophozoites of

the flagellated protist Giardia muris attached to the villi of
the small intestine of an experimentally infected mouse. This
microparasite reproduces asexually via binary fission.
(Micrograph courtesy of Břetislav Koudela.)
Fig. 1.5 Scanning electron micrograph of the nematode
Heligmosomoides polygrus from the intestine of a mouse.
This macroparasite-host system is a widely used model in
untreated drinking water can lead to massive numbers
experimental parasitology. (Micrograph courtesy of Doug
Colwell.)
of feeding stages in the intestine of a range of verte-
brate hosts.
Parasites can have parasites too! The parasites living
in/on other parasites are called ‘hyperparasites’.
Parasite biodiversity will increase exponentially when
we fully understand how common hyperparasitism is
in nature. The sea louse, Lepeophtheirus salmonis, for
example, is a common skin ectoparasite of salmonid
species. A monogenean fluke, Udonella caligorum is a
hyperparasite of the egg sacs of sea lice. In addition,
microsporidians such as Desmozoon leopeophtherii,
have recently been described as intracellular hyper-
parasites of L. salmonis.
The organism in, or on, which a parasite reaches
Fig. 1.6 Female of the ectoparasitic mite Varroa destructor
sexual maturity is the ‘definitive host’. Many parasites
attached to the abdomen of a developing honey bee.
(Photograph courtesy of Scott Bauer, USDA Agricultural have a simple, direct life cycle, requiring only one host
Research Service, Bugwood.org.) for transmission to occur. All monogeneans, and many
nematode and arthropod parasites, have direct life
ectoparasitic or endoparasitic. They are typically cycles. Many animal parasites, however, have obligate
intracellular, i.e., adapted to recognize, penetrate, and ‘intermediate hosts’ in which the parasites undergo
reproduce within host cells, or they may exploit developmental and morphological changes.
extracellular tissues, or both. Eukaryotic micropara- Intermediate hosts may be the prey of the predatory
sites include protozoans, microsporidians, and myxo- definitive host in the life cycles of parasites. Thus,
zoans. In the case of the protist, Giardia spp., parasites with intermediate hosts in their complex life
(Fig. 1.7), ingestion of a single cyst originating from cycles are often transmitted trophically to definitive
C:/ITOOLS/WMS/CUP-NEW/4264071/WORKINGFOLDER/GOAT/9780521190282C01.3D 10 [1–15] 6.8.2013 8:16AM

10 Introduction

hosts via food web interactions. Life cycles in which Trichinella spiralis, and the human blood fluke,
more than one host are required are referred to as Schistosoma japonicum. Ecologically, reservoir hosts
indirect life cycles. Many parasites have remarkably are similar to paratenic hosts since they may greatly
complex life cycles with several hosts and both free- increase transmission rates, and also help prevent local
living and obligate parasitic larval stages. extinction of the parasite. The potential for controlling
Some protozoans and filarial nematodes have ‘vectors’ zoonotic diseases in humans is greatly complicated by
as hosts. Vectors are micropredators that transmit infec- the presence of these reservoir hosts. Furthermore, res-
tive stages from one host to another. A vector may be an ervoir hosts greatly complicate the zoonotic parasite’s
intermediate or a definitive host, depending on whether ‘epidemiology’. Epidemiology is the study of all the
the sexual phase of the parasite’s life cycle occurs in it or many complex, inter-related ecological factors respon-
not. For example, the insect vectors for species of sible for the transmission and distribution of a human
Plasmodium, the causative agents of malaria, are certain disease. A related term is ‘epizootiology’. Epizootiology
species of female mosquitoes that actively inoculate usually refers to the factors involved in the transmission
infective stages of the parasite into the next host during and distribution of non-human parasites, often in
their blood meals. Sexual reproduction occurs within the reference to epizootics. The epidemiology of human
stomach of the mosquito; consequently, mosquitoes are parasites and the epizootiology of parasites of fishery,
the definitive hosts for the parasite. veterinary, and wildlife importance will be stressed
A number of parasites may use hosts in which there throughout the upcoming chapters.
is no development and that are not always obligatory
for the completion of a parasite’s life cycle. These are
called ‘paratenic’ or ‘transport hosts’. Such hosts are 1.4 Overview
most frequently used to bridge an ecological, or tro-
phic, gap. For example, adults of the fluke, Halipegus All eukaryotes have the capacity to recognize invading
occidualis, live under the tongue of green frogs (Color cells or organisms as non-self. It follows that a fun-
plate Fig. 1.1). Snails in the genus Helisoma are obli- damental feature of the parasitic life style lies in the
gate first intermediate hosts, whereas aquatic micro- host’s ability to defend itself against a limitless diver-
crustaceans such as ostracods are obligate second sity of invaders. It is the strength and duration of host
intermediate hosts. But green frogs do not normally defenses that defines such features of the host–parasite
consume these small crustaceans. It turns out that relationship as specificity, parasite-induced pathology
various species of odonate (dragonflies and damsel- and phenotype alteration, site selection, parasite–
flies) prey upon ostracods, thus acting as paratenic mediated natural selection, and coevolution. Given the
hosts for this trematode. Thus, frogs are exposed to fundamental importance of host defenses to the ecol-
Halipegus larvae when they prey on nymphs and ogy and evolution of parasitism, we provide an intro-
metamorphosed odonates. In this four-host life cycle, duction to the massive field of immunology in
the parasite exploits two predator–prey interactions to Chapter 2. Our coverage is not meant to replace an
enhance transmission (see Fig. 6.23). introductory course in immunology. Instead, our aim
A number of animals are normal hosts for parasites is to cover the basics of innate and adaptive immunity
that may also infect humans. These are called ‘reservoir in both invertebrates and vertebrates, and also provide
hosts’. These non-human hosts act as reservoirs of an introduction to the interdisciplinary fields of
infection for certain parasites. Diseases of animals that immunoparasitology and ecological immunology.
are transmissible to humans are called zoonotic dis- The following nine chapters cover the biodiversity
eases, or ‘zoonoses’. Thus, giardiasis, trichinellosis, and of animal parasites. In each of these chapters we take a
schistosomiasis are examples of zoonoses. Similarly, parasite-centered approach to describe functional
rats are important reservoir hosts for the nematode, morphology, life-cycle variation, and biodiversity. Our
C:/ITOOLS/WMS/CUP-NEW/4264071/WORKINGFOLDER/GOAT/9780521190282C01.3D 11 [1–15] 6.8.2013 8:16AM

Overview 11

coverage within these chapters is selective, and not of Life. These new tools have, in many cases, trans-
meant to parallel the coverage of this material in formed traditional views of parasite biodiversity and
modern parasitology texts. Our aim is to provide have led to a re-interpretation of the course of evolu-
examples that are representative of key groups, espe- tion in several of the main groups. These new phylo-
cially those that provide models that are used to ask genetic schemes also have implications for how we
leading ecological or evolutionary questions. To con- diagnose and treat some key parasites of humans.
clude each of these chapters, we cover phylogenetic Thus, we emphasize modern phylogenetic classifica-
classification. Our aim is to summarize the tremendous tions of the main groups of animal parasites because
advances that have been made over the past decade in they provide an evolutionary context for the chapters
parasite ‘systematics’, paralleling the increased use of that follow, and because they have led to new insights
molecular classification schemes across the entire Tree into the evolution of parasitism. Box 1.2 provides a

Box 1.2 Parasite systematics: a phylogenetics primer

Deciphering the evolutionary relationships of parasitic animals can be a daunting, yet exciting
challenge. Consider briefly three enigmatic taxa that have long intrigued invertebrate biologists
and parasitologists. The pentastomids (Chapter 10) are primarily obligate parasites of the lungs
of reptiles. These parasites have been shown to be most closely related to a group of arthropods
known as branchiuran crustaceans. A crustacean in a snake’s lung seems bizarre to say the least!
Or, consider the Myxozoa described in Chapter 5. Previously classified with single-celled
organisms, it turns out that these multicellular fish endoparasites share evolutionary affinities
with cnidarians. This makes the cnidarians far more diverse, both in terms of morphology and
habitat exploitation, then was ever appreciated. Perhaps equally remarkable is that the endo-
parasitic thorny-headed worms (Acanthocephala) are evolutionarily linked to certain free-living
rotifers (see Box 7.4). To fully understand how biologists have made such monumental changes
to the Tree of Life you need to master some of the terminology of the modern systematist.
The focus of phylogenetics research is ‘homology’. Morphological features that share a common
evolutionary origin are said to be homologous. Thus, any differences in homologous features
among different parasite groups reflect descent from ancestors with modification (Pechenik, 2010).
Homologous features can then be used in construction of phylogenetic trees and taxonomic
classifications. Unfortunately, homologous features are not always easily recognized.
Morphologies, for example, can independently evolve from very different ancestors, to give a close
resemblance by convergence. These morphologies may appear to be homologous, but they are not.
Another problematic issue concerns the direction of evolutionary change. Which of two homol-
ogous characters is the original, or ancestral, state, and which represents the more derived state?
This question can be particularly difficult to decipher for parasites. We will see that much of the
debate concerning the systematics of a given group of parasites centers on these issues.
Cladistics is a widely used approach to deduce evolutionary relationships. The terminology in
this field of systematics is complex. We introduce here as few terms as possible in order to help
you understand the basics. Cladistics use so-called ‘synapomorphies’ (Greek, synshared,
C:/ITOOLS/WMS/CUP-NEW/4264071/WORKINGFOLDER/GOAT/9780521190282C01.3D 12 [1–15] 6.8.2013 8:16AM

12 Introduction

Box 1.2 (continued)

aposeparate/derived, morphēform) to establish evolutionary relationships. These are shared,

evolutionarily novel features derived from a common ancestor in which the characters origi-
nated. These evolutionary novelties are used to construct a ‘cladogram’ that depicts evolutionary
relationships (Fig. 1.8). A cladogram is a representation of branching sequences that are
characterized by unique changes in key morphological or molecular characteristics (=synapo-
morphies). A cladogram shows the least complex, most parsimonius way of explaining the
evolutionary history of the groups. A ‘clade’ is a group of organisms that includes the most
recent ancestor of all its members and importantly, all descendents of that ancestor. By this
definition, a valid clade forms a ‘monophyletic’ taxon (Fig. 1.8a). Synapomorphies are vital
because they identify monophyletic groups. If species and groups are classified in a way that
reflects evolutionary history, only monophyletic taxa should be given names. Determining
monophyly, then, is a primary goal for systematists. Monophyly can be elusive, and instead,
there may be inconsistencies between taxonomic schemes and estimates of phylogeny. A taxon
is ‘paraphyletic’ if it contains the most recent common ancestor of all members of the group and
some, but not all, of its descendents (Fig. 1.8b). Finally, a ‘polyphyletic’ taxon is an incorrect
grouping containing species that descended from two or more ancestors (Fig. 1.8c). Thus,
members of polyphyletic groups do not share the same immediate ancestor. It is important to
recognize that these inconsistencies are an artefact of a naming scheme that is an inaccurate
estimate of phylogeny, or vice versa. Both cladists and evolutionary taxonomists reject poly-
phyletic groups in taxonomic classifications because of this mixed ancestry. It is for this reason
that some formerly recognized taxa are no longer considered in classification schemes or they
are reclassified to generate a monophyletic group. For example, the ‘old’ protist phylum
Sarcomastigophora, comprised of amoeboid and flagellated unicellular animals is no longer
considered valid because many phylogenetic studies have shown that it is a polyphyletic taxon
(see Box 3.7).
A B C D X A B C D X A B C D X Cladists denote the common descent
of different taxa by identifying ‘sister
up

up

up
ro

ro

ro
tg

tg

tg

taxa’ or sister groups on cladograms.

ou

ou

ou

a) Monophyletic b) Paraphyletic c) Polyphyletic Sister taxa are each other’s closest rela-
Fig. 1.8 Cladograms showing the relationships of four taxa: A, B, tives; they arise from the same branch-
C, and D. (a) depicts a monophyletic group or clade, defined by a ing point (node) on a cladogram because
shared, derived trait or synapomorphy; note that taxa CD and
ABCD are also monophyletic; (b) 1 of 4 possible paraphyletic they are derived from the same ancestor.
groups is illustrated (others are ABC, ABD, ACD); (c) 1 of 4 Cladists also identify an ‘outgroup’, a
possible polyphyletic groups is shown (others are AC, AD, BC). closely related taxon that lies outside the
The outgroup (X) is phylogenetically close but not within the
taxa being studied (Fig 1.8). The out-
taxa being studied. (Figure courtesy of Danielle Morrison.)
group’s characteristics are assumed to
represent the ancestral condition.
C:/ITOOLS/WMS/CUP-NEW/4264071/WORKINGFOLDER/GOAT/9780521190282C01.3D 13 [1–15] 6.8.2013 8:16AM

Overview 13

Box 1.2 (continued)

The best phylogenetics studies consider dozens of different characters to infer evolu-
tionary relationships. The field of phylogenetic systematics has been revolutionized by
accessing the genetic code of life via the nucleotides in DNA and RNA, and comparing
these gene sequences (especially 18S ribosomal RNA and mitochondrial DNA) among taxa.
Indeed, throughout the upcoming chapters we will see that it is this molecular revolution that
has greatly altered our taxonomic classification system of parasitic animals. Moreover,
molecular data are helping us to more fully understand the origins and evolution of para-
sitism, and the interrelationships within the various animal parasite taxa (e.g., see Figs. 5.11,
7.12, 8.26).
While a primary focus of many phylogenetics studies is to help resolve parasite taxonomic
issues, they are also widely used for several other reasons, including analyzing the geographical
distributions of hosts and their parasites, and how distributions have changed over time. The
research field uniting phylogeny with geography, as well as evolution and geology, is called
‘phylogeography’ (Chapter 14). In addition, phylogenetics is widely used in studies of host–
parasite ‘coevolution’ and ‘cospeciation’ (Chapter 16).

primer for the two approaches, classical taxonomy characterize the structure of multi-species parasite
(or evolutionary systematics) and phylogenetic sys- assemblages and then emphasize how modern
tematics (or ‘cladistics’), that are so often used to empirical approaches have uncovered some unex-
test hypotheses regarding the origins of parasitism pected underlying mechanisms. We conclude this
and the evolutionary relationships between and chapter by describing comparative patterns of para-
among taxa. site biodiversity among host species, ending with a
The next six chapters summarize advances in discussion of the approaches that field-based para-
parasite ecology and evolution, drawing upon sitologists utilize to infer causation. In Chapter 14,
results from modern empirical studies on a wide we extend the hierarchy to include aspects of para-
range of model host–parasite interactions. This sec- site biogeography and phylogeography. Our
tion is structured hierarchically, beginning with emphasis in this chapter is on how tools drawn from
coverage of parasite population biology and epi- molecular biology, genomics, and remote sensing
demiology in Chapter 12. Following a section on are revolutionizing our understanding of the geo-
terminology, we briefly introduce the mathematical graphical distribution of parasites, ranging from
foundation upon which these subdisciplines rest. We continent-wide scales, down to a few meters.
then shift to a brief introduction of how parasite In the final three chapters, we shift our focus
populations are distributed within their host popu- away from the parasites themselves, onto the nature
lations, and how parasite population sizes are of the host–parasite interaction and the complex
determined and regulated. We proceed along this interplay that exists between the two partners over
natural hierarchy to synthesize advances in parasite ecological and evolutionary timescales. Given the
community ecology in Chapter 13. Here, we first exploitative nature of the relationship, the concept of
Discovering Diverse Content Through
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Jje forseid wonde, forsope, was fully cured in pe forseid maner.
Witte pou 36 pat pis anoyntmewt is best to al spotte} or filpe} of pe
skyn which giffep oute watre and makep redne}, for it * " lekej "
overlined. "smallach " overlined. c Obliterated.
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man was smyten on his legge vpon J>e shynbone, A

patient Treatment of an Inflamed Leg. 51 driej) mich and dobe away
rednes in Query place of be body, out-tak in be ei^en. Bot witte bou
bat after be puttyng to of be oynt- [leaf 155, 4 ment fat is )ws made
— Recipe sape, sulphwr & arsenec — ber appered a blak litel cruste
to be biknes of a seme of a scho, ]>at was hard for to parte with be
forseid corrosiue} for it was mich ritted. To which I putte aboue 8 a
cautery, i.[e.] a bry?myng iren, bat be pacient almost The cautery
feled it no^t. After J>e cauteriyng forsobe, I putte to J»e schauyng
of larde, as it is seid aboue, in sewyng al pings vnto ]>e ende. 12
12. [A] but neberlesse be skyn was nojt cleuen l alsone after be ieg
and an ulcer smytyng. Afterward, forsobe, be J>rid day it bolned
formed. and bigan to ake. berfor he went to a man bat haunted 16
or vsed sich cure vnto be tyme bat ber come in his legge ane hole,
rounde and depe, and ful of blak filth in maner of brent flesch ;
whome whan he come to me I heled hym bus. ffirst I wasched be
wounde vrith hote 20 wyne, or water in which was decocte )>e
croppej or be iuyse of plantayne or sich, or in vryne. Afterward I
putte to ane emplastre made of iuyse of playntayne, Ardeme's of
rubarb, of smalach, of hony, and whete or rie mele & 24 white of
eyren y-medled togidre ; or ane emplastre bat is called
sangwiboetes. Afterward, be place sumwhat mollified, I putte to
poudre Creoferoboron, -with J>e andstimumedicine of arsenec y-
medlet ; aboue be poudre stupe^ powder. 28 or carp ; aboue al-
togidre, forsobe, )>e emplastre of apii, mugwort, walwort seid afore.
After" J>e puttyng to forsobe of bis poudre, I did pe cure in al bings
-with lard & wt'tA oper bings, as it is seid aboue, vnto be 32
clensyng of be wounde. Afterward, forsojj, -with vnguento viridi &
vnguento albo and carp I wrojt in maner as it is seid aboue vnto be
ende. Afterward, forsob, when ber growed or wex any superflue
flesch in be wounde, I He treated 36 wztAstode it or mette it vrith
poudre of creoferoboron or flesh, of litarge vnto be fulle curyng of
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52 Two cases of Swollen Leg. Wounds with swords and

axes must be treated like other injuries. Bruises from the kick of a
horse or from stones should first be scarified. A patient's leg swelled
suddenly on a Christmasday. Arderne fomented it, and by cockcrow
the patient was relieved. The juice of marigold is very useful in
inflammation of the breasts and in whitlow. [leaf 156] A prescription
for tartar water. cure pam wztA vnguento albo, as it is seid aboue.
fforsop if pe wonde be in pe leg of swerd or ax or sich oper, be it
cured as ben oper woundes. If any man, forsop, be smyten in any
party of pe legge 4 violently and without \vondyng of pe skynne, as
fallep oft-tyme of )>e smytyng of ane horse fote, or of a stone or of
sich oper, fan is it gode sone in pe bigynnyng for to garse J>e place
y-smyten and for to draw out blode 8 per-of, and afterward for to
putte to emplastre^ repressyng )>e akyng and bolnyng. [A] man in
pe day of pe natiuite of our lorde sodenly had his legge gretly
bolned fro pe kne to pe ankle}, 12 with redenes and gret brennyng,
so pat he my^t no^t stand. I (smeared ?)a pe legge on ych side,
and epithimated wt'tA pe iuyse of solseqwz, i.[e.] marigold, and a
litil vinegre putte perto,1 and made paui a litel leuke b ; be which 16
y-do, lynnen elopes wette in pe same iuyse I laide warme aboute his
legge, and when pe elope was dronken of pe iuse I laid hym in his
couche ; And for certayn afor pe cok kraw pe akyng and pe
brennyng was cesed and 20 pe pacient rested wele. And witAin pre
daies witAout any oper medicyne he was perfitely cured, whar-of
many men wondred. Also for certayn pe iuyse of solsiquii, marigold,
epithimated bi it-self or \vith vinegre destroyep 24 meruelowsly
apostemej in pe pappes of wymmen, and pe felon,0 and pe
carbuncle and ^ekyng,2 and rednes, and blone}, and brennyng pat
comep of pe forsaid pings. \_A-j chanon was on a tyme seke, and
when he bigan to 28 wex hole par was made a grete gedryng to-
gidre of humowrs descendyng doune in his legge. After a tyme,
forsop, per wex puscele} brovnysch and clayisch.3 He, forsop, putte
pat he schuld dry pe pusche} watre of 32 tartar pws y-made :
~Recipe tartari Bb i or ij, and putte it in ane newe erpen potte, and,
pe moupe of pe potte stopped witA clay, putte it in a strong fire and
lat it be per a ni^t and a day or more if pou witf. Afterward 36 tak
pat tartar and hyng it in some place in a lynnen a Obliterated. b "
warme " overlincd. 0 " antrace " overlined. 1 quern curavi cum succo
solsequii addito parum de aceto et cum dicto succo tepido
epithimiavi tibiam suam undique. [MS. Digby 161. leaf 22, back.] 2
Pruritus. 3 pustnlse fuscse et latae.
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Treatment of a Mormale or Ulcerated Leg. 53 sacce or

pokette, and vnder it putte a brasen vessel to receyue fe watre fat
distillef droppyngly to fe maner Thetreatof lye out of f e sak ; f is
watre is seid for to dry pusche} ulcerated 4 wele, bot nef erlesse it
availed no}t to hym. At f e last, canon!* forsof, fer grow in fat party
of fe legge a large wounde, And about fe ankles f re or four smale
wounde} to fe brede of ane halfpeny. And fe legge semed of 8 }elow
colour medled \\iih rednes fro fe calf to fe ankele}, And fe skynne
kast euermore out many skale}. When, forsof, he had vsed a
certayne tyme lede or puluer incarnatyue and sawe fat it availed
hym nofing, fan 12 he vsed a long tyme ane entrete fat is called
entractuw Arderne i^Quse titu- nigruw,1 blak entrete, which is made
of white lede and the"caseas Dyueiyn. rede and comon oile and
tartarye «fcc. ; bot nef erlesse he inflamed ,, ,, . , sore on the
perceyued none amendyng ferol, ior it come to a mor- ieg. 1 6 male
; f e which, when I had sene it, I aff ermed it to be a mormale. And
I did sich a cure to it : f is is f e cure to fe mormale — ffirst sewe fe
pacient legge strongly His treatsere'^ihiam Wit^ a tynne clobe 2 ;
After wasche Wele fat legge SO bandaging foititer et 20 sewed vfith
hote watre, after fat fe pacient may suffre. foment!- '° lum stricte
And so after be waschyng lat it lye by a natwrel day, * ms> in panno
lineo. fat is ane hole day & a ni3t, kepyng fe legge fro aier and fro
cold, be second day, forsof, remoue fe clofe 24 and muwdifie fe
wouwde or fe wouwdes if fai be many, and putte in euery wounde a
litel pece of lynne clofe then cold . , compresses, moisted in cold
watre. Afterward putte of f e oyntement » Postea of dyuylyne in be
circuite of f e wouwde 3 aboue f e hole pone de isto unguento 28
skynne so fat it touche no waie? fe Avounde? wttft-in. & in circnitu J
' ' vuineris. couer it \vith a lynne clofe y-wette. Do fus euery day
tuye}, renewyng fe oyntment and muwdifying f e wounde} and
fyllyng fam of a lynne clofe y-wette, as it is seid 32 aboue. bis is fe
oyntment : l&ecipe coperose, sal nitri, afterwards mercurial a cinens
geneste,a cineris b nigri testudinis, atrament^', ointment, ana, parte
1 ; viridz's grec^ somwhat ; Of quikke-siluer double to fe quantite of
one of f e forseid ; Of bore} 3G grese resolued at fe fire and
mumlified, fat sufficef. ban medle first fe ashes \vith fe grese,
afterward fe tofer poudre}, and when fou hast wele y medled, put it
in a 3 — a " aschen of lirome " overlined. **— b "of blak snaile "
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Treatment of a Mormale or Ulcerated Leg. and finally La ri

Crank's ointment, though he had to use the knife. [• leaf 156, back]
It is best to cutaway the dead flesh in an ulcerated leg, and then to
apply a powder. Lan frank in his book gives advice about the cure of
a mormale. box and it schal be blak oyntment. With pis oyntment,
forsop, I cured fully pe gretter wondej of pe forseid legge, doyng in
pe maner seid afore ; J>e lesse wonde$, forsop, cured I with
vnguewto viridi, i.[e.] grene oyntment of lanfrank. Jjer was dede
flesch of bio colowr to pe brede of a peny ; pat dede flesch, forsoj),
was mich pikke, and, pat y-se, I kutte with a rasour a litel pe ouer
party of pat flesch ; Afterward I putte to larde, and so at pe last with
larde & with cuttyng I dissolued, i.[e.] lesyd it vtterly. }3at flesch pe?
-for remoue, eftsonej with pe oyntment of dyuylyn * aforeseid and a
elope wette in water I held pe wounde opne to pe brede of a peny,2
And pan eftsonej per brest out a wounde aboute pe side}, and it
bygan to large it vnto pat it was almost of pe same gretnej as it was
afore.3 )5at y-sene, I putted in four tymej poudre of litarge and
anoynted it about with vnguewto albo, and putte in pe wounde a
lynne elope wette in pe water of herb robert. Which cure semed to
me more p?-ofitable, and sowded better pe extremite} and glowep 4
pam vnto perfite halpe. *If pe mormale be euen aboue pe schyn-
bone, pat it be more sikerly and more sone cured it is profitable to
cutte pe dede flesch and putte it away if pe pacient consent. And if it
be cutte, alsone after pe cuttyng is to be putte iu a cloute wette in
whyte of ane ay al a ny^t. Afterward putte in poudre of white glasse
and of alum ^ucaryne, i.[e.] alww glasse, or alum plume or of bope.
And if pou se pe bone mortified, witte pou pat it is incurable or
vnnep for to merowe be cured. If pou trow it be curable, it is to be
helped with some cure of pe mormale in pe boke of lamfrank. Also,
as it is seid aboue, som tyrne a man is smytyn som party of pe legge
violently wit/
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A Treatise on Hcemoivhoids. 55 ward, if it be nede, of pe

sophene. At pe last be he scarified* in J)e leggej. I A I tretys of J>e
emoraide^y-drawen out after lam- [leaf 15T] 4 frank, a discrete
maistre of pe kyng$ of fraunce ; which made oVpaes86 tuo bokej of
cirurgie, be lesse and pe more. Also after maister bernard de gordon
in his lilie. Also after maister bartelmow in his passionarie. Also after
maister Richard Chiefly a . . compilation. 8 in his Micrologie. And
after maister Eoland, and mayster Guy; And after Roger Bawn And
maister lamarcii, And maister Gilbertyne ; And after oper experte
men whos doctryne I haue beholden & sene, and which I haue 12
founden moste experte in practicing, with helpe of our lord. I schal
schew pam in pis boke. Emoroys on Etymology greke is said flux of
blode, and it is seid of emak, fat is Emeroids. blode, and rois, flux.
Greke}, forsop, callep emeroys use the word 16 flux of blode in
what-ener parti of pe body it be; Bot general"5 1 Apud anence
latyne men l pis worde is appropriate to pe flux of the8Latinn blode
of J>e lure ; And pe veynes apperyng in )>e lure when M 2 quando
pai flwe,2 i.[e.] ren, and ar bolned and ake)>, pai ar called 20
emeroydej, bot neperle} vnproperly, sauand pe pece of be comon
puple. ffor when bai send out no blode, bot ar bolned, and akep,
and ychep or smertej) pai ar called by ober names anence lechej.
Lewed men and vnex- The unJ _ learned call 24 perte men callej) al
J>e infirniitej bredyng in fe lure everything emeroydej, or pilej, or
fics. ffrench men calle)? emeroydej Frenchmen fics, men of London
callej) J>am pilej. Neferlesse it is "figs," noct to strife agaynej fe vse
of spekyng, bot raper it 28 spede]) |>at lered men and experte
knawe Jje maner of spekyng and vse it. ffor John Damascen seij> '
It is hexiy for to chaunge noying custom, and most if it be olde.'
Neberlej of be name is no stryuyng whiles be but there -r-\. t
nothing 32 sekenej bene knowen. Dmerse auctowrej, forsof, hap
much in a , , name if all putte diuerse names to be sekenej of t>e
lure, and also are agreed •i -jj- j • AJU- as to the pai haue assigned
dme?-se causes and spicej, And pai condition. haue ymagined many
maners of curacions ; Of whiche 36 some more profitable and ofter
experte bene sewyngly to be noted vnder compendiowsnej to be
vtilite b of helyng. )3erfor for to trete schortly it is first to witte pat
be » ' ' garsed " overlined. b "i.[e.J profite" overlined.
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56 Piles, Condylomata, Warts — Causes and Symptoms.

Varieties of piles : " the deaf piles " of Avicenna. Condylomata, why
so called. The cause of condylomata ; their appearance. Verruca,
their causes; the symptoms of the inflamed variety : of the chronic
form. Piles due to congestion. emeroide3 if fai sende out blode fai ar
seid ry}tfully emeroyde}. If \er appere, forsof, in fe lure bolnyngs bio
or blak, or redne} to f e quawtite of a bene or gretter, fat is to f e
quawtite of a testicle of a cok or of a hounde, 4 as I haue oftyme
sene, som tyme occupiyng f e to half of J>e lure only, and som tyme
bofe, sich bene called of Avicen deef emeroyde}, for f er rynnef
noting out of Jam. And sich bolnyngs forsof, if fei be gretter, 8
puttyng out no blode, fai ar called condilomata, of condilo of greke,
fat is fe closed fist of a man. Condilomata, forsof, schewef pe
schappe of a fist y-closed, And condilomata bredef of gret malicious
or malencoli- 12 ous blode. After lamfrank, forsof, fe lesse bolnyng}
if f ei be blak or bio fai ar called attritos,1 or atreos, for fe blak
colowr of fam. If fai be rede fai ar called uve, i.[e.] grape^, and fai
haue fe schap of a rede vyne or 16 grape. And fai fat bene of blode
and of colre ar called morale},2 and fai ar like to mulberie} when fai
bigynne to wex rede. And som bene called verucale} 3 for fai ar like
to warte}, and sich haf fair bygynnyng of malen- 20 colye. And som
bene of blode, fof it be bot seldom, which ar called fics, 4 If fai be
made of ventosite 4 with grete strechyng of fe skynne. Al fe forseid
may be reduced vnto tuo fings : Oufe?1 of hotene} of humowrs, 24
or of mych aboundyng of blode. If fat hotene} be in cause, fat is
blode and colre, fise schal be fe signe — brennyng witA greuozts
prikkyng, and smertyng, and vnslepyng, and som tyme wt'tA ychyng
in fe lende} and 28 vfilh tenasmon and gret costyuene} of fe
wombe, and frist, and feblene} of goyng. Signe} of cold cause, fat is
of gret blode and malencoliows, bene fise — bolnyng vfiih hardne}
and derkne} and akyng — bot not scharp as 32 of hote cause — fe
colour of fe bolnyng bio or blak, smertyng in fe lure, wztA lousene}
of fe wombe and akyng, and greuowsne} or heuyne} of fe fie}.
Signe} if fe em[er]oide} be of multitude of blode bene fise, fat 36 is
to sey of fe veyne} apperyng in fe legge}. And it' 1 " blakej "
overlined. 2 "mulberiej" overlined. 3 ' ' warty " overlined. 4— 4 "after
gordon, and bai ar as war white bledders " overlined.
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Piles, their Pathology and Uses. 57 J et medicinas venarum

aperitivas suimmt. 3 Symptoinata. pai ren, pat pai ren mych & oft-
tyme, and pat pe pacient be of sanguyne habitude, ffor why ; in
pam is multitude of blocle * fat vsep not fleebotomye, and pat 4
drynkep copiously and oftymej wyne, and fat etep scharp pings, as
onyons, lekej, caule^, corny ne, and pat takep medicynes apertyue^
of veyne^,1 as bene scamonye i.[e.] aloe and euforbiu???, as
wittenessep all auctowrs 8 togidre and experte men. Emeroide3 ar
caused of malencoliows blode, which is pe fece of clene blode
aboundand in our body; which blode, forsop, for his yuel quality and
odiow« to nature, discretyue vertu enforcep for to cast 12 out to pe
helpyng of al pe body, helpyng pe vertu expulsyue of al pe membrej
togidre. And so purj pe strengpe of nature it is putte out fro pe
vayne kilyuz,2 pat liep to rig-bone of pe bak, which properly is
recep16 tacle of malencoliows blode. Which kylis, forsop, is diuided
into fiue brauchej pat bene ended about pe party of nature a; which
veyne$, forsop, when pai ar sorn tyme filled of melaucolioMS blode
pai distende, i.[e.] 20 strechep, so pe veyne^ pat ouper pe blode
brestep out or per ar gendred bolnyng^ of diuerse spice^ and
schapej. And also oper sinthomata,3 i.[e.] pe?'ile^, as scharp akyng
and prikkyng, brynnyng, ychyng, smertyng, thenasmon, i.[e.] 24
inordinate appetite of egestion, wit/i ful mich enforsyng and
nepe?'lesse he may do none egestion whan he coniep to pe pryue.
If, forsop, pe blode brist out it is called pe emoroyde} ; but if pat it
flowe temperatly it dop 28 many helpyngs and preseruep pe body
fro many sekene^ adnste and corrupte, as is Mania, malewcolia,
pleuresis, lepre, morfe, ydropisy, mormale, quartane, passions of pe
splene, and so of oper like. Bot as it p?-eseruep fro 32 pise when pat
it flewep temperitely, so when it is wont for to flewe and afte?-ward
cesep vtterly al pe forseid sekenes ar gendred. Also when pai flewe
ouer temperance pai bene cause of ptisyk or of ydropisy. 36 Wherfor
seip Galiene and ypocras after lamfrank ' Ich long lastyng and
ouermych puttyng out of blode is moste mi^ty cause for to make
ydropisy.' Jjerfor in pam in whome malencolio»s blode is multiplied
temperite fluying a " i.[e.] lure " overlined. [* leaf 157, back] Those
who are subject to congestive piles. The cause of piles. The
pathology of piles. Anatomy of the vena cava. Symptoms of piles.
Uses of piles when they only bleed moderately.
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58 The varieties of Bleeding Piles. What constitutes

moderate bleeding. H»\v to stop the bleeding from piles. The cause
of the bleeding in piles. [* leaf 158] The impatience of the present
generation. Bleeding piles are often concealed piles. of blode of fe
emeroyde^ helpef mych, ne it is no$t vtterly to be restreyned. It is
called temperite vse when f e pacientes felef f amself more li$t fat
jjai war wont l ; * cum patihauyng better appetite, and etyng and
slepyng more 4 sentiunt swetely or softely, and sich o]>er. Bot when
Jje pacientes soiito et felen famself more heuy, and fer schewe
malice of c'oiorati. appetite and foule colowr of body, fan is fe flwyng
ouer mych; wherfor it is alsone successyuely to be restreyned 8 and
turned away. Jperfor sife ]>er is nojt a litel hardnej in restreynyng of
fe emeroidej, ferfor many f ings ar be noted of J>e restreyning of
fain ; fat is to sey fat fe leclie wytte whefer fe flowyng be done 12 of
anathemasy or of diabrosi or of rixi; fat is whefer fe flowyng be
made of opnyng of veynej, fat is called anaf emasis ; or of fretyng of
f e veyne^, fat is called diabrosis; or of cleuyng or twynnyng, fat is
16 called rixis. f erf or if fe blode ybro^t to fe lure be aduste — for
when blode is aduste it is scharped2 — or if 2quiaciim sanguis false
flewme or colre be medled, fan oft-tymej f e veynej aduftu"r ar
freted and fai make fe fluxe. And for certayne sich 20 flux is of hard
restreynyng. ffor why ; fe substance of f e veyne yf reted may no^t
be * sonded 3 wttA-out disese and heuynes, sife fat it nedef a
medicyne corrosyue. And men now-of-daiej bene vupacient and yuel
tholyng, 24 And for-fi flowyng of diabrosi a is of hard curying. And f
e secundary is rixis b which also is cured with corrosyue^ in fe
bygynnyng. Anathemasis c is more Ii3tly cured fan fe ofer, bot
perauenture nojt without corrosyue}. 28 jjis I sey, if fe flowyng be
olde, Anathemasis is made for aboundance of blode or for ventosite
descendyng doune. Eixis, forsof, is made of ouer myche dryne$, of
cause witAin-forf e or of cause wi't/tout-forfe, or of hardnej 32 of
filfe^or forane hote aposteme, or any scharpe flowyng. Also flowyng
emoroydej somtyme ar hidde within, w?t/iout any bolnyngs
schewyng outward, fat of som fai ar denied to be dissenterikej and
yueh wrong.5 ffor why, in 36 5
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Signs of danger in Bleeding Piles. 59 passion is akyiig of be

wombe vtith puttyng out of blode 'J"h«y »«? sewyng. Also afte?"
gordon, scilicet de morbo, ' In euerv I , , • from dysenbing bat gob
out of be body bene 3 comon cause? ; teryand ' passio iliaca. i aut
ratione 4 Ouber by reson of be membre. or of vertue.1 or of membri
aut virtutis. humour. If it be bi reson of be membre. bat is for be
Bernard de . . , Gordon's membre is oue?' bmne. If it be for vertue,
ban it is Liiium for vertue retentyue is feble, and vertu expulsyue
strong, quoted. 8 If it be for humowr, bat is for ouber it is
malencoliows or for it is scharp, or subtile, or watrye. Also emoroidej
ar caused of scharpnes of blode and oue?' mych hete brennyng be
blode, as in colorik men bat bene of hote Piles are due 12 nature;
for blode when it is brent it geteb scharpnes, as of blood, it is seid
afore. Also oue?' mych flowyng of blode is made oube?- for
multitude of blode, as in bam bat drynkeb mvch wyne or ober metej
or drynkei bat to eating 1ft u- v u li j • and drinking AO multipliep
blode, or in bam bat bene sangyne com- too much, plexion. Also it is
made for yuel qualite of blode, as for it is oue?' scharp or subtile or
watry, as in bam bat vseb 0° yj^f^ rawe fruyte^, ffor raw fruyte}
gendrej) watry blode. }je 20 causes, forsob, y-knowen, propre cure
may be done to. Signe$ of dedly flowyng bene bise, bat is to sey : —
fflowyng of blode bryngyng to swouwyng is mortale a ; Also flowyng
of blode witA coldne^ of extremitej is mor- Danger 24 tale ; Also
flowyng bat comeb sodeynly and with bleeding hastinej is mortale ;
Also flowyng of blode bat bryngeb sudden to pale colow?', or grene,
or bio, or browne is werst and mortale ; Also quantite of blode
passing 4 pouwde is yuel, £allor28 and if it come to 24 it is deb.
fflowyng of blode w/t/t lijtnyng of be body is gode. In bam bat boleb
emoroidej be vryne schal be in cokmr remissed white with powdry
resolucions blak or bio residentej in be 32 bothme of be vessel. After
Egidi de vrinis, white and Quotes remisse ow for to be of
malencoliows blode oue?' aboundant corbeii de in be body, wherfor
naturel hete is febled. ffor why ; digestion waxeb raw, and of rawnes
of digestion is be 36 vryne discolored, * And it appereb \vikh poudry
resolucions r» ieaf iss, which bene resolued of malencoliows blode
blak and erbi abouredyng, and by contynuel waiej bai ar drawen to
be bladdar and putte out wit/t be vryne. And for bai ar 40 heuy and
erbi bai satle in be grounde. }3e same vryne a ' ' dedly " overlined.
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60 The Treatment of Bleeding Piles. How nolime-iangere

and lupus are produced. The blood is not confined. Choler and
melancholy are cribbed in the gall bladder and in the spleen. The
signs of melancholy. Treatment of bleeding piles by herb pills.
Phlebotomy is best in the overfed and idle whose piles bleed. also
may betokne in men vicea of pe splene, and in wymmen
witAholdyng of menstruej. And witte pou pat after gordon in '
clarificaci'one de vicio splenis ' }>at innatural humozirs may be
gendred in oper place fan in 4 pe lyuer, as in pe stoniak colre
peassyue,1 i.[e.] grene, and also colre eruginows, of which is
gendred ' noli-me-tangere,' and lupus. And also in pe veyue^ ar
gendred vnnaturale humowrs. Bot pe splene hap no vertu of
gendryng 8 any ping, sipe it is noping hot a receptakle of malencolie,
which is ane odious humowr to nature and to al membris of pe body
for his yuel qualitej. Also witte pou pat pe blode hape nouper
house,b ne receptakle, ne prison; 12 but colre and malencoly hap
prisons, pat is to sey colre in pe chiste of pe gall and malencoly in pe
splene. Also witte pou pat if pe pacient of emoroidej be of
malencoliows complexion, pise bene tokne$ : — smalnej c of 16
body, discolorwed, erpi, angry, waike of hert, heuy, and only ferpful
and couaitows. And witte pou pat if pe forseid pacient sende out
blode blak and pikke and stynkyng, pat pis flowyng is no3t to be
restreyned, hot if 20 it ouerflowe & pe pacient be febled. In euery-
ping, forsop, pe vertu of pe body is to be kept bifor al oper pings.
Agaynj pe flowyng of pe emoroidej distempre pou moste subtile
mele of whete, -with iuyse of millefoile, 24 and make perof piling,
and giffe hym euery day in pe mornyng 3 or 4 of pam distempered
vrith wyne of decoccion of millefoile, or plantayn, or burso pastoris,
or rede netle, or paruencw. Bot if pe pacient be of san- 28 guyne
complexion, and lifyng delicately and in ydelnej, and blode be
aboundand, pan pof per be sych flowyng it is no^t to be restreyned
hot if it ouer flow, pat is knowen by pe toknej aforeseid. J)erfor if
per faH ouer 32 mych flowyng, it is spedeful pat it be restreyned;
for, after galien, blode is norischyng of al me?nbrej, als Avele of
sadde as of softe, and al hap bigywnyng or spryngyng of blode ; and
fcr-als-mych it is seid pe frende of nature, 36 pe?'for if pe frende be
destroyed pe enemy waxep mi^ty. Jjerfor, after gordon, to pe curyng
of pe emoroidej is fleobotomy competent, if vertu and age sutf're it,
a " sekeiies " overlined. b " duellyng " overlined. c "or Iene3 "
overlined. prasina.
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The Treatment of Piles ~by Phlebotomy. 61 bobe for it

avoidej) matery goyng afore, and also it wit/idraweb be matery to be
contrary.* 'And fleobotomy uioodtobe ow to be done of be basilic
veyne of be arme for mater *«>»» the • ' basilic vein 4 goyng afore,
and afterward of be sophenis at be hele, be or the ~. * external
vtter sophe turneb be flowyng of be emoroide} and saphenous.
restreyneb be emoroyde} for euermore. Whiche fleobotomy, forsob,
continued *fro }ere to }ere, and namely I* fcanw] 8 about be fest of
seynt Micheli, bifore and after one} or twie}, or when-someuer be
pacient felej? tyklyng or The Weeding ychyng or prykkyng in be lure,
ban be he minusched as annually6 it is afore seid, and alsoue he
schal be cured. Also witte Michaelmas. 12 bou bat fleobotomye of be
inward sophenis of be leggej prouokeb be emoroide} and menstrue}
; And of be vtter Bleeding sophenis streyneb be emoroidej and
menstruej, and we- internal ' . saphenous serueb for certayne fro be
forseid passions. Sophenej lia<1 for Piles i c i and men°r16 bene bo
grete veyne^ bat ar strecned fro be kneej vnto rhagia. be ankle^ of
bobe partiej of be legge}. )3e maner of doyng of bis fleobotoraye is
bat it be done about be ^^my hour of euensong or latter, bat is in
be regnyng of saphenous 20 malencoliows blode, bat is fro be 9
hour of be daie vnto be 3 hour of be ni?t. Also witte bou bat fleo-
Evening is ' ' ' the proper botomye to be done vnder b be hele and in
saluatella of time for 1 phlebotomy. be handej, oweb no^t to be
done w»t» a blode iren bot 24 with a lancete, for hurtyng of be
synewe^, but if fat nede make it. Also witte bou bat he bat schal be
laten Treatment blode oweb for to putte his fete in hote watre, and
eft- phlebotomy, sone bam owe to be putte agayn, bat be blode go
out 28 bette?- ; And be pacient ow to abide still in be watre, vnto
bat be blode fat appered first blak dhaunge into fairer colow. Be bis
doct?-me boldly kept, for if it be no^t done competently it profiteth
nobing or litel ; ffor 32 certayne I haue cured for euermore, wiih on
latyng blode al-only of be vtter sophene, many men bat boled be
emoroidej many jeres, vnto be feblyng of be body. Bot witte bou
bat, after gordon, bat bof-al fleo36 botomye make blode fluxible,
neberlesse if it be done of be vtter sophene} it draweb be flowyng to
be opposite ; 1 et ita facit. and so it makyng x be flowyng for to be
turned fro be lure bat it flow nojt to be costomable place ; and 40
forbi it profiteb to al bat ar cured of be emoroide}, a " opposite "
overlined. b " at " overlined.
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62 Treatment of Piles ly Purgatives. and also to al pat ar

disposed to pe emoroide}, for The patients to be laten blode of pe
outward sophenis, one? or tuyei should be * v~* let blood m ver
and alse otte m autu??ipne fro }ere to }ere. fror twice every why ;
sich fleobotomye for certayne preseruep fro pe 4 autumn even
emoroide}, and avoidep paciente} fro al yuel humowrs piiesgbe '* *
and kepep pam in hele. And per be no blode-later redy, be per made
ventosyng with garsyng atuix pe buttoke} at pe rigebone ende or
aboue pe veynej, for it dope pe same 8 pat pe forseid fleobotomye,
bot nojt so expertly, bot nede The patient ^Q^Q no lawe. 1 And it is
to witte pat if pe leche may i Ad quod must be faciendum purged
gette pe ri}t reule by al pings in euery wirchmg after myrobaiani
bled. crafte, fleobotomye ow}t neuer to be done in pe emoroide} 12
tumconbyfore purgacion y-done, and pan sewyngly mynischyng.
quia myroAlso after gordon, in ouer mych flowyng of pe emoroide},
and also of menstruej, is competent a medicyne laxatyue to purge
corrupte humours pat inducep pe flowyng, pat 16 comprfmenpe
cause cesyng, pe etfecte cese. To which ping to be done
eos^mnores fotoebbest8 al pe mir[obalan)] a [con]ouenie/*t ffor
mira- "nt^Myr"fn^^nes^6 bolan) laxep afore pam and streynep
after pam . . . . a j^m' hoc and pai avoide al yuel humowrs. Jjerfor
be pai preparate 20 ™£^J>^_ .a decocte byfor pat pai be taken, for
bi Non debent • myrobaiani decoccion pair a decoqui antequam
back] *and so pe strength of laxatiueyng shuld be febled, bot
fumantur, bai ow to be resolued in rennyug liquore, as in mylk or 24
decoctionem J eorum gumraisins and whey, vfiih racyns & liquorice,
ellea walld pai schrenk b niitas in liquorice ' * fumum remay be used,
j,e stomake and lefe yuel tokenea byhyndc pam: and so soiveretur ,
- ,. , , . , et sic vis of be racyns and of pe liquore ar pai reuled pat
pai bryng purgandi der*n--\ t u i j /^ i • ii OC bilitaretur. in no
harme. Which, forsop, resolued and pe kirnelle^ -^o cast away, lat
pam lie al a ni^t in pe same liquor, and in pe mornyng pe colyng be
giffen to drynk. jjis medicyne, forsope, of mirobolan) is ful noble,
sipe it purgep humo^^rs of nijt placej, pat is citrine} colre,2 kebuli}
fleume,3 Indi 32 gj,"*^^.1 malencoly, i.fe.l blak colre, bellerici and
emblici colre 3 kebuii flavam. aduste. And so may al pe humowrs be
purged ; wherfor be mirabolan) hadde in reuerence, ffor pai availe
mich to pe emoroide} and menstrue} to be purged. And pis medi-
36 cyne for certayn curep euery flux of pe wombe, pat is to scy
diaria, dissenteria, liencaria. Also it is to wytte pat • A portion of the
leaf is missing. b "ronkle " overlined. c ' ' after " overlined.
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Treatment of Piles ly Diet and Medicines. 63 in pacientj of

be emoroide^ be ]>er neuer giffen medicyne} apertyues of veyuei
noube?' be ]>e moube, as scamone, or but never J \ scammony
aloe, or euforbium, comyn, lekej, onyans, garleke and sicb. or aloes.
4 ober scharp bing} ; ne be bar nojt putte opon be place, out take
aloen and comyn, which tuo, putte outward, streyneb, and witft-in-
forb taken opne]> veyne}. Bot it is to wytte bat in be forseid fluxes
of be wombe mirabolan) ow to be Jh,e myrobalans may 8 dissolued
in gote mylk,1 if it may be had, or in iuse or bejdvenin goat s milk.
water of fumitere or of playntane, or in rayne water or rose water, or
of veruene, or of anober stiptike herbe as millefoile or mynte. Also
witte bou, after al auctowrs, 12 bat pose pings bat restryneb
emoroide} restreyneb menstruej, and econverso : And be same
sekenej bat comeb of what is good . , , . -TO f°r Piles is be vice of
menstruej, comeb also of be emoroid, & econverso : equally good . ,
for menorand so by sewyng a bat ]>ai acorde in cure, perfor in cold
rhagia. 16 cause be J>ai giffentpat hetep and ingrossep J>e mater
of flowyng of blode, als wele of be party of mete} as of medicyne}.
Of J>e party, forsob, of mete^ be ber giffen milk, mele of whete
decocte, frese bene^, with canel, gret triticaf faba* ^ wyne> r^3e 2
an(i ^y^te. Of be party of medicyne be ber Remedies to fera cum
giffen note of cipresse 3 and be lefej, mirre, thure. mastike. the
treatcinnamomo, ' ment of vinum Iadanu7?i, storax calamita, anyse
rosted, and sich obe. piles. crassum, rube. And excercice and strong
frotyngs and swetyngs availejj. pressl CU 24 And if pe cause be
hote, be ber giffen lentesb vrith vinegre, 4 Portuiaca porcelane,4
sour milk, soben barly brede, substance of colej 6 cauiisdua- j?e tuo
watrej 5 bat it is decocte in y-cast away, perej, abjectis. coyncej 6 &
melde^,7 & sich obe?'. Medicyne} restrictyue} e Mespiia. 28 bene
bise ; Camphore, accacia, spodin, coriandre, sangwe's 7 Cotonea. , .
. ' » hEemat. draconts, sandali, lap[isj omoptoes,8 bole armoniac,
ypo9 Hypo- quistid,9 galley cupule, sumak, mirtett,10 quistidos. . , ,
., . . . . .c plantane, cincloile, qumquineruie, nbbewort, 32 bursa
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