Cambridge University Press

978-0-521-85008-7 - Ecology of the Acanthocephala

C. R. Kennedy
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Ecology of the Acanthocephala

Commonly known as spiny-headed worms, acanthocephalans have

a characteristic eversible proboscis, which bears rows of recurved
spines used for attachment to the intestine of their host.
Acanthocephalans are endoparasitic and an extremely successful
group found in almost all marine, freshwater and terrestrial systems,
infecting a huge range of definitive (usually vertebrate) and
intermediate (usually arthropod) hosts during their life cycles. This
volume is concerned with the ecology (distribution and abundance)
of the Acanthocephala, and through this approach aims to reveal the
huge success of this group of parasites. The acanthocephalans have
evolved differently from all other groups of parasites, and as such
represent a distinct and alternative pathway of parasite evolution
and host
parasite interactions. Written for parasitologists, ecologists
and zoologists who are interested in learning about a different
pathway of parasite evolution, this book is aimed at postgraduate
and research scientists.

is Emeritus Professor of Parasitology at the

C. R. KENNEDY

University of Exeter.

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Cambridge University Press
978-0-521-85008-7 - Ecology of the Acanthocephala
C. R. Kennedy
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Ecology of the
Acanthocephala
C. R. KENNEDY
School of Biological Sciences,
University of Exeter

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Cambridge University Press
978-0-521-85008-7 - Ecology of the Acanthocephala
C. R. Kennedy
Frontmatter
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CAMBRIDGE UNIVERSITY PRESS

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Published in the United States of America by

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ß Cambridge University Press 2006

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 2006

Printed in the United Kingdom at the University Press, Cambridge

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ISBN-10 0-521-85008-8 hardback

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978-0-521-85008-7 - Ecology of the Acanthocephala
C. R. Kennedy
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Contents

Preface page vii

1 Introduction 1
1.1 Biodiversity and the Acanthocephala 1
1.2 An ecological approach to the Acanthocephala 8

2 Life cycles and transmission 12

2.1 General strategy 12
2.2 The adult stage 14
2.3 Eggs 16
2.4 Intermediate hosts and cystacanths 18
2.5 Paratenic hosts 20
2.6 Post-cyclic transmission 24
2.7 Variation in life cycles 26

3 Biogeography and distribution 28

3.1 Phylogenetic influences 28
3.2 Biogeographical influences 32
3.3 Host features 43

4 Specificity 52
4.1 General strategy 52
4.2 Criteria for specificity 56
4.3 Spatial variations in specificity 63
4.4 Strain specificity 69

5 Host
parasite interactions 75
5.1 Distribution of acanthocephalans within hosts 75
5.2 Relations in vertebrate hosts 81
5.3 Relations in invertebrate hosts 84

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978-0-521-85008-7 - Ecology of the Acanthocephala
C. R. Kennedy
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vi Contents

5.4 Alterations in intermediate host behaviour 87

5.5 Significance of behavioural changes in hosts 99

6 Population dynamics 101

6.1 General considerations 101
6.2 Seasonality and short-term studies 104
6.3 Regulatory mechanisms 110
6.4 Long-term studies 116
6.5 Mathematical models 121

7 Community dynamics 125

7.1 General considerations 125
7.2 Richness and dominance of communities 127
7.3 Spatial and temporal variability 137
7.4 Niche dynamics 141
7.5 Interspecific interactions 146

8 Introductions and extinctions 158

8.1 Dispersal 158
8.2 Introductions and colonisation 165
8.3 Rarity 172
8.4 Extinctions 174

9 Relations to ecosystem changes 179

9.1 Biological tags 179
9.2 Acanthocephalans as indicators of pollution 183
9.3 Impact on ecosystems 192

10 Conclusions and overview 200

References 211
Index 241

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C. R. Kennedy
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Preface

To do science is to search for repeated patterns, not simply to accumulate facts.

The only rules of the scientific method are honest observations and accurate logic.
To be great science it must also be guided by a judgement, almost an instinct,
for what is worth studying.
MacArthur (1972)

We conclude that the complicated interplay between virulence and

transmissibility of parasites leaves room for many coevolutionary paths to be
followed, with many endpoints.
Anderson & May (1982)

The Acanthocephala are generally considered to be a small and

rather insignificant group of parasites. They are a minor phylum
of interest to specialists; as they rarely, and then accidentally, infect
humans or cause significant disease or disorder to their farmed and
domestic animals, they receive a cursory treatment even in most
parasitology textbooks. Nevertheless, there have been some books
devoted exclusively to them, principally Acanthocephala of Domestic
and Wild Animals (Petrochenko, 1956, 1958), Vol. V of Systema Helminthum
(Yamaguti, 1963), An Ecological Approach to Acanthocephalan Physiology
(Crompton, 1970) and Biology of the Acanthocephala (Crompton & Nickol,
1985). The first two of these books are fundamentally systematic
in their approach, whereas the latter two are far more biological.
Crompton & Nickol’s (1985) book does contain some chapters on
acanthocephalan ecology, but it was published 20 years ago, and more
recent information on their ecology can only be found in the primary
scientific literature. Much recent information is being presented at the
International Acanthocephalan Workshops, but proceedings of these
are not published.

vii

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C. R. Kennedy
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viii Preface

Parasitology can be viewed as a special branch of ecology since it

is concerned with the relationships between organisms and their
environment. This present volume attempts to present a comprehen-
sive account of the ecology of the Acanthocephala. By so doing, it aims
to show how an ecological approach can open up a new perspective on
the phylum such that acanthocephalans can now be seen as a common,
widespread and very successful group of parasites. The pattern of their
evolution has differed in many respects from those of all other groups
of helminth parasites and so they present a distinctive and character-
istic path of host
parasite co-evolution.
Their impact on their hosts extends beyond host individuals onto
populations and communities of free-living organisms and indeed into
whole ecosystems. They are also proving to be very sensitive indicators
of some forms of pollution. It is intended, therefore, that this book
will be of interest to a wider readership, especially to freshwater
and marine biologists and also to ecologists in general, as well as to
parasitologists who may be excited by the prospect of recognising and
studying a different pattern of parasite evolution. It is aimed primarily
at postgraduates and researchers and it assumes at least an under-
graduate knowledge of parasitology and ecology. It challenges some
current ideas and paradigms on biodiversity, by showing what can be
achieved in evolutionary terms by a group of organisms that comprises
a small number of species and exhibits a high degree of morphological
uniformity. It aims to provoke an interest in the group and to stimulate
the reader. It also attempts to pluck the phylum from the obscurity of
current textbooks and place it on a par with, for example, cestodes and
nematodes as being a group worthy of study for its own sake.
No book can be comprehensive in its coverage, and this one is no
exception. I am solely responsible for the selection of examples, which
have been chosen from a wide range of hosts and habitats around the
world. Nearly all examples in figures and tables have appeared in
published form previously and I must thank all the authors and publi-
shers for permitting me to use them: the source of each is acknowl-
edged fully in the legends to figures and captions to tables. While the
author alone must be held responsible for presentation and any errors,
no book can ever be completed without the help of many people. I
would like particularly to thank Ward Cooper, the then Commissioning
Editor at Cambridge University Press, for his advice and efforts on my
behalf and also his successor Dominic Lewis for being available to
answer my many queries and deal with problems. Many of the ideas
presented here have been developed in discussion with friends and

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Preface ix

colleagues over a long period of time; and although it is invidious

to single out particular persons I would like to mention Al Bush,
David Crompton, Bahram Dezfuli, Celia Holland, John Holmes,
Brent Nickol and Phil Whitfield in particular who have all contributed,
often unknowingly and unwittingly, to the development of my ideas.
I owe them a great deal as colleagues and friends. Other ideas
have emerged from discussions with my postgraduate students and
postdoctoral fellows and to them, and especially to Rachel Bates and
Alastair Lyndon, I express my thanks.
Many other people have helped me in a variety of ways,
by providing me with photocopies and advice, and I would particularly
like to express my thanks to Omar Amin, Chris Arme, David Gibson,
Eileen Harris, Robert Poulin, Jamie Stevens and Bernd Sures. Two other
people deserve special thanks. The first of these is Ian Tribe, who,
although a botanist, read through the manuscript for me and made
many very valuable suggestions. Finally, and above all, I must thank my
wife Pat. My interest in acanthocephalans was set alight by some of the
questions raised in her thesis and now 30 years later I am still trying to
explain and understand the significance of some of her findings. She
was in large measure responsible for planting the idea of this book in
my mind and in getting me to start on it despite my insistence that
I would never write another book. She has supported me and
encouraged me continually throughout the writing, through the
creative times and through the inevitable frustrating times. Quite
simply, this book could not have existed without her and both I and
anyone who reads it and enjoys it owes her an enormous debt.

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