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Heinz Mehlhorn

Animal
Parasites
Diagnosis, Treatment,
Prevention
Animal Parasites
Heinz Mehlhorn

Animal Parasites
Diagnosis, Treatment, Prevention
Heinz Mehlhorn
Department of Parasitology
Heinrich Heine University
D€usseldorf, Germany

This is an updated translation of the 7th edition of the German book “Die Parasiten der Tiere”
(2012) published by Springer Spektrum.

ISBN 978-3-319-46402-2 ISBN 978-3-319-46403-9 (eBook)


DOI 10.1007/978-3-319-46403-9

Library of Congress Control Number: 2016956719

# Springer International Publishing Switzerland 2016


This work is subject to copyright. All rights are reserved by the Publisher, whether the whole or part of
the material is concerned, specifically the rights of translation, reprinting, reuse of illustrations,
recitation, broadcasting, reproduction on microfilms or in any other physical way, and transmission
or information storage and retrieval, electronic adaptation, computer software, or by similar or
dissimilar methodology now known or hereafter developed.
The use of general descriptive names, registered names, trademarks, service marks, etc. in this
publication does not imply, even in the absence of a specific statement, that such names are exempt
from the relevant protective laws and regulations and therefore free for general use.
The publisher, the authors and the editors are safe to assume that the advice and information in this
book are believed to be true and accurate at the date of publication. Neither the publisher nor the
authors or the editors give a warranty, express or implied, with respect to the material contained
herein or for any errors or omissions that may have been made.

Cover illustration: Left: Scanning electron micrograph of the anterior end of the hookworm Ancylostoma
caninum showing typical teeth. Right: Scanning electron micrograph of the anterior end of a tapeworm of
the genus Taenia showing a species specific crown of hooks besides the suckers. Photos Heinz Mehlhorn

Printed on acid-free paper

This Springer imprint is published by Springer Nature


The registered company is Springer International Publishing AG
The registered company address is: Gewerbestrasse 11, 6330 Cham, Switzerland
The Structure of the Book and How to Use the
Diagnostic Keys

Contents

Chaps. 1–3 explain what parasitism means, where such animals live and how they
can be diagnosed most easily.
Chaps. 4–6 present the most important parasites of animals. Using the tables in
Chap. 2, where the different parasites are listed according to their site of occur-
rence, it becomes easy to find the relevant chapter in the book where a detailed
description is given in addition to micrographs and summarizing tables.

How to Use the Diagnostic Keys

In several chapters, a key is provided, which helps to diagnose some important


parasites. The keys should be used as follows:

– The user of the book gets in general 2 (rarely 3) choices of answers on questions
related to morphologic aspects of a given parasitic stage (explained in the text
and/or shown on micrographs).
– If the user has decided, he/she finds at the right side of the question a number in
bold appearance.
– Then he/she goes to the next question written behind this number in the next line
and so on. It is, however, important to read always all possible answers before a
decision is made!
– If at the right side the name of a parasite or a group of parasites occurs, the
diagnosis is completed and the user follows to the given page number, where the
searched parasite is described in detail.

v
vi The Structure of the Book and How to Use the Diagnostic Keys

Text

The parasites of all animals in the close surroundings of humans are described in
detail in Chaps. 4–6, which are presented in a sequence with respect to their
organization as protozoans, worms (helminths) and ectoparasites. Each parasite
and its closely related groups are described in the following subheadings:

1. Name: The origin of the species names is explained in order to understand the
often complicated names.
2. Geographic distribution/epidemiology: This section gives insights into
where the parasite occurs (worldwide, locally, etc.), and it is shown whether
they are of high or low importance.
3. Biology/morphology: This section covers aspects of the appearance and repro-
duction of the parasites.
4. Symptoms of disease: The most important symptoms of disease introduced by
this parasite are listed.
5. Diagnosis: This topic contains information how to find most easily a supposed
parasite in the body of a host or in its feces, etc.
6. Pathway of infection: This section describes how the host can be infected with
a supposed parasite.
7. Prophylaxis: This section gives short information about how infections with
the diagnosed parasite can be avoided.
8. Incubation period: This is the period ranging from the day of infection until
first symptoms of disease occur.
9. Prepatent period: This is the period from the day of infection with the
diagnosed parasite until first parasitic stages can be diagnosed, which can be
transmitted. The prepatent period is often shorter than the incubation period so
that other hosts can be infected without knowing it.
10. Patency: Period during which a parasite can be transmitted to another host.
11. Therapy: Presentation of methods/compounds that are in common use to treat
infections. Note: Medication may differ in different countries due to local laws.
12. Further reading: This section gives some rather new and/or older but impor-
tant papers, which help to approach relevant literature concerning a diagnosed
parasite.

Addenda

Addendum A: Antiparasitic Drugs Used for Animals

This addendum contains tables where different medicaments are listed that can be
used for the treatment of infections occurring in the different host groups. It gives
more details than in the special Chaps. 4–6.
The Structure of the Book and How to Use the Diagnostic Keys vii

Addendum B: Diagnostic Stages

This addendum encloses comparative tables with additional micrographs of stages


of several parasites of the different hosts.

Addendum C: Questions to Test Obtained Knowledge

This addendum lists 100 questions concerning important animal parasites offering
always five potential answers, but only one of them is correct. This allows us to
control our own knowledge and is recommended especially for students before
examinations. Correct answers are listed at the end of this addendum.

Addendum D: Origin of Figures

The origins of the macro- or microscopical figures of the parasites used in this book
are listed.
Preface

Parasites endanger not only the health of humans but also that of animals which live
together with humans in homes and on farms. Many groups of parasites are in
addition transferable from animals to humans and back. The pathways of
transmissions of endoparasites and ectoparasites have obtained increasing impor-
tance in our days of globalization and global warming. While globalization offers
easy transportation of agents of diseases from one end of the world to the other,
global warming increases the chances of many parasites (especially ectoparasites)
to enter new biotopes in formerly cold regions. Before this background, it seems
necessary to stabilize and to increase the knowledge on parasites with respect to
occurrence, transmission and control. Thus, the contents of this volume are
addressed to veterinarians and students of veterinary medicine but also to all people
who keep farm animals and especially to those who are owners of pet animals.
In order to make it easy to find the wanted information, the book presents the
relevant knowledge on each parasite under the following 12 subheadings:

1. Name
2. Geographic distribution/epidemiology
3. Biology/morphology
4. Symptoms of disease
5. Diagnosis
6. Pathway of infection
7. Prophylaxis
8. Incubation period
9. Prepatent period
10. Patency
11. Therapy
12. Further reading

The correct diagnosis of the different parasitic stages of animals is not only
possible by the description of the relevant methods but also supported by more than
600 micrographs, by many diagrammatic representations as well as by comparative
tables. The listed control measurements represent the actual knowledge in the year
of the appearance of this book and should always be checked.

ix
x Preface

Each chapter on a given parasite ends with the presentation of some recent
papers, which allow a retrospective of important features of the parasite.
Hundred questions—each with five choices—allow personal control of our own
knowledge. They had been tested in many student examinations to confirm a solid
knowledge.
This book represents the first English edition after seven previous ones in
German.

D€usseldorf, Germany Heinz Mehlhorn


June 2016
Acknowledgements

The representation of a book such as this one, considering a broad spectrum of


many features of the same topic, is not possible without the help of experienced
colleagues and friends. Thus, this book is based on common books and articles
published with renowned colleagues such as Dieter D€uwel, Dieter Eichenlaub,
Alfred Otto Heydorn, Thomas L€oscher, Werner Peters ({), Gerhard Piekarski ({),
Wolfgang Raether and Eberhard Schein ({). Furthermore, I am deeply indebted to
my colleagues Johannes Eckert (Z€urich), Axel Haberkorn (Bayer Leverkusen),
Heinz Hänel (Hoechst AG, Frankfurt), Alfred Otto Heydorn (Berlin), Sven Klimpel
(Frankfurt), Gerd Lehmann (Aalbaum), Brigitte Loos-Frank (Stuttgart), Gerhard
Piekarski ({) (Bonn), Rainer Pospichel (Bayer AG), Regine Ribbeck (Leipzig) and
Eberhard Schein ({) (Berlin) for their contribution of beautiful pictures, their
advices and/or nice glass slides to prepare rare aspects of parasites. Dr. Volker
Walldorf (D€ usseldorf) and Mr. Fried Theissen ({) (Essen) contributed nice
drawings especially prepared from drafts.
My son Tim, Mrs. Diehl and my wife Birgit helped in translating the text into
English. The preparation of the text was done by Mrs. Inge Schaefers and Mrs.
Susanne Walter. Presentation of the figures was accomplished by Mrs. Isabelle
Mehlhorn, Mrs. Susanne Walter and Mr. Bernd Pr€umm.
The staff at Springer Heidelberg (Drs. Andrea Schlitzberger and Lars K€orner)
finished this book in its hopefully very useful and agreeable final version.

D€usseldorf, Germany Heinz Mehlhorn


June 2016

xi
Contents

1 Phenomenon Parasitism . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1
1.1 Who Are Parasites? . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1
1.2 Parasites and Their Hosts . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1
2 Where Are Which Parasites? . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11
2.1 Parasites Within Feces or Intestine . . . . . . . . . . . . . . . . . . . . . 12
2.2 Parasites in Blood . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14
2.3 Parasites in Saliva . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18
2.4 Parasites Inside Lymph Fluid . . . . . . . . . . . . . . . . . . . . . . . . . 18
2.5 Parasites in Urine . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18
2.6 Parasites in/on Mucous Layers . . . . . . . . . . . . . . . . . . . . . . . . 19
2.7 Parasites in/on Tissues . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19
2.7.1 Parasites in Macrophages . . . . . . . . . . . . . . . . . . . . . . 19
2.7.2 Parasites in Liver and Spleen . . . . . . . . . . . . . . . . . . . 19
2.7.3 Parasites Inside Muscles . . . . . . . . . . . . . . . . . . . . . . . 19
2.7.4 Parasites in Lung, Trachea and Gills . . . . . . . . . . . . . . 19
2.7.5 Parasites in Sexual Organs . . . . . . . . . . . . . . . . . . . . . 19
2.7.6 Parasites in Bones . . . . . . . . . . . . . . . . . . . . . . . . . . . 20
2.7.7 Parasites in Kidneys . . . . . . . . . . . . . . . . . . . . . . . . . . 20
2.7.8 Parasites in the Swim Bladder of Fish . . . . . . . . . . . . . 20
2.7.9 Parasites Inside the Brain . . . . . . . . . . . . . . . . . . . . . . 20
2.7.10 Parasites in Eyes . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20
2.7.11 Parasites in the Skin . . . . . . . . . . . . . . . . . . . . . . . . . . 20
2.8 Parasites on the Skin, in the Fur or in Plumage of Birds . . . . . . 21
3 Methods to Diagnose Parasites . . . . . . . . . . . . . . . . . . . . . . . . . . . . 23
3.1 Investigation of the Feces . . . . . . . . . . . . . . . . . . . . . . . . . . . . 23
3.1.1 Macroscopical Procedures . . . . . . . . . . . . . . . . . . . . . 24
3.1.2 Microscopical Procedures . . . . . . . . . . . . . . . . . . . . . . 25
3.2 Investigation of Parasites Inside Blood . . . . . . . . . . . . . . . . . . . 30
3.3 Further Cytological and Serological Methods . . . . . . . . . . . . . . 31

xiii
xiv Contents

4 Protozoan Parasites . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 33
4.1 Trichomonads . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 33
4.1.1 Tritrichomonas foetus . . . . . . . . . . . . . . . . . . . . . . . . . 33
4.1.2 Trichomonads of Donkeys and Horses . . . . . . . . . . . . 38
4.1.3 Trichomonas Species and Tetratrichomonas Species
of Birds . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 39
4.1.4 Histomonas meleagridis (Histomoniasis) . . . . . . . . . . . 43
4.2 Giardia Species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 46
4.2.1 Giardia Species of Dogs and Cats . . . . . . . . . . . . . . . . 51
4.2.2 Giardia Species of Ruminants . . . . . . . . . . . . . . . . . . 52
4.2.3 Giardia Species of Birds . . . . . . . . . . . . . . . . . . . . . . 53
4.2.4 Spironucleus Species (syn. Hexamita) . . . . . . . . . . . . . 54
4.3 Trypanosomes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 55
4.3.1 Trypanosomes (Salivaria): e.g. Agents of Nagana . . . . 65
4.3.2 Trypanosoma brucei evansi (Agent of Surra) . . . . . . . . 67
4.3.3 Stercorarian Trypanosomes . . . . . . . . . . . . . . . . . . . . . 68
4.3.4 Trypanosoma brucei equiperdum . . . . . . . . . . . . . . . . 70
4.3.5 Trypanosomes of Fish . . . . . . . . . . . . . . . . . . . . . . . . 73
4.4 Leishmania Species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 76
4.4.1 Leishmania Species of Dogs . . . . . . . . . . . . . . . . . . . . 79
4.4.2 Leishmaniasis of Cats, Equids and Cattle . . . . . . . . . . 82
4.5 Amoebae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 83
4.5.1 Entamoeba histolytica . . . . . . . . . . . . . . . . . . . . . . . . 83
4.5.2 Amoebas of Reptiles and Frogs . . . . . . . . . . . . . . . . . 87
4.5.3 Malpighamoeba mellificae . . . . . . . . . . . . . . . . . . . . . 88
4.5.4 Acanthamoeba Species and Related Groups . . . . . . . . . 90
4.6 Apicomplexa (Sporozoa) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 91
4.6.1 Eimeria Species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 92
4.6.2 Isospora and Cystoisospora Species . . . . . . . . . . . . . . 114
4.6.3 Cryptosporidium species . . . . . . . . . . . . . . . . . . . . . . 120
4.6.4 Sarcocystis Species . . . . . . . . . . . . . . . . . . . . . . . . . . 128
4.6.5 Toxoplasma gondii . . . . . . . . . . . . . . . . . . . . . . . . . . . 153
4.6.6 Hammondia Species . . . . . . . . . . . . . . . . . . . . . . . . . . 160
4.6.7 Neospora Species . . . . . . . . . . . . . . . . . . . . . . . . . . . . 162
4.6.8 Besnoitia Species . . . . . . . . . . . . . . . . . . . . . . . . . . . . 164
4.6.9 Further Species with Tissue Cysts . . . . . . . . . . . . . . . . 168
4.6.10 Babesia Species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 171
4.6.11 Theileria Species . . . . . . . . . . . . . . . . . . . . . . . . . . . . 187
4.6.12 Agents of Malaria or Malaria-Like Diseases . . . . . . . . 195
4.7 Ciliates and Flagellates . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 207
4.7.1 Balantidium coli . . . . . . . . . . . . . . . . . . . . . . . . . . . . 208
4.7.2 Buxtonella sulcata . . . . . . . . . . . . . . . . . . . . . . . . . . . 211
4.7.3 Ichthyophthirius multifiliis and Related Ciliates . . . . . . 211
Contents xv

4.8 Microsporidia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 216


4.8.1 Encephalitozoon cuniculi . . . . . . . . . . . . . . . . . . . . . . 216
4.8.2 Microsporidia of Fishes . . . . . . . . . . . . . . . . . . . . . . . 220
4.8.3 Nosema Species of Bees . . . . . . . . . . . . . . . . . . . . . . . 222
4.9 Myxozoa . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 224
4.9.1 Myxobolus (syn. Myxosoma) cerebralis in Fishes . . . . . 227
4.9.2 Tetracapsuloides bryosalmonae of Salmonids . . . . . . . 229
4.9.3 Sphaerospora renicola . . . . . . . . . . . . . . . . . . . . . . . . 229
4.10 Blastocystis Species (Blastocystosis) . . . . . . . . . . . . . . . . . . . . 230
4.11 Pneumocystis Species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 232
Further Reading (Joint List for This Chapter) . . . . . . . . . . . . . . . . . . . 234
5 Worms (Helminths) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 251
5.1 Platyhelminthes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 251
5.1.1 Trematodes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 252
5.2 Tapeworms (Cestodes) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 296
5.2.1 Tapeworms of Carnivores . . . . . . . . . . . . . . . . . . . . . . 310
5.2.2 Tapeworms of Ruminants and Pigs . . . . . . . . . . . . . . . 334
5.2.3 Tapeworms of Horses (Equids) . . . . . . . . . . . . . . . . . . 341
5.2.4 Tapeworms of Birds . . . . . . . . . . . . . . . . . . . . . . . . . . 346
5.2.5 Tapeworms of Rodents and Rabbits . . . . . . . . . . . . . . 349
5.2.6 Tapeworms of Reptiles and Amphibia . . . . . . . . . . . . . 351
5.2.7 Tapeworms of Fishes . . . . . . . . . . . . . . . . . . . . . . . . . 352
5.3 Nematodes (Roundworms) . . . . . . . . . . . . . . . . . . . . . . . . . . . 356
5.3.1 Morphology of the Nematodes . . . . . . . . . . . . . . . . . . 363
5.3.2 Development of the Nematodes . . . . . . . . . . . . . . . . . 373
5.3.3 Nematodes as Agents of Disease in Animals . . . . . . . . 377
5.4 Tongue Worms (Pentastomida) . . . . . . . . . . . . . . . . . . . . . . . . 464
5.4.1 Linguatula serrata (Tongue Worms, Pentastomida) . . . 467
5.4.2 Other Pentastomids of Reptiles . . . . . . . . . . . . . . . . . . 468
5.5 Acanthocephala (Thorny-Headed Worms) . . . . . . . . . . . . . . . . 470
5.5.1 Macracanthorhynchus hirudinaceus
(Giant Thorny-Headed Worm of Swine) . . . . . . . . . . . 474
5.5.2 Thorny-Headed Worms of Birds . . . . . . . . . . . . . . . . . 475
5.5.3 Acanthocephala of Fishes . . . . . . . . . . . . . . . . . . . . . . 476
5.6 Leeches (Hirudinea, Annelida) . . . . . . . . . . . . . . . . . . . . . . . . 479
Further Reading (Joint List for This Chapter) . . . . . . . . . . . . . . . . . . . 484
6 Ectoparasites (Arthropods: Acari, Insecta, Crustacea) . . . . . . . . . . 499
6.1 Arachnids, Chelicerata . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 499
6.1.1 Ticks . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 503
6.1.2 Mites . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 525
xvi Contents

6.2 Insects . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 571


6.2.1 Lice (Phthiraptera) . . . . . . . . . . . . . . . . . . . . . . . . . . . 574
6.2.2 Fleas (Order Aphaniptera, Siphonaptera) . . . . . . . . . . . 588
6.2.3 Bloodsucking Bugs (Order: Rhynchota, Heteroptera) . . . 603
6.2.4 Order Diptera . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 607
6.3 Crustaceans (Crustacea) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 646
Further Reading (Joint List for This Chapter) . . . . . . . . . . . . . . . . . . . 653

Addendum A: Antiparasitic Drugs Used for Animals . . . . . . . . . . . . . . 663

Addendum B: Diagnostic Stages . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 671

Addendum C: Questions to Test Obtained Knowledge . . . . . . . . . . . . . . 681

Addendum D: Origin of Figures . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 697

Author Index . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 699

Subject Index . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 701


About the Author

Prof. Dr. Heinz Mehlhorn has investigated


parasites, their transmission pathways and sig-
nificant control measures for over 40 years. He
has published more than 20 books and 250 origi-
nal publications and received 25 patents on
antiparasitic drugs, some of which he uses at
his university spin-off company Alpha-Biocare
(founded in 2000). As a university instructor, he
had the pleasure to introduce many students to
the topics in parasitology. Many of them are now
professors or in leading industrial positions. In
television and radio broadcasts, he regularly
informs the public about relevant parasitological
problems in our days of globalization and global
warming.

xvii
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