Coccidiosis in Livestock, Poultry, Companion Animals, and
Humans - 1st Edition
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Praise for the Book
Coccidiosis in Livestock, Poultry, Companion Animals, and Humans is a long needed update and review for this group
of clinically important protozoan parasites that infect a wide variety of hosts. It’s been 25 years since the coccidia were
thoroughly reviewed with updated information so it’s timely and needed. The group of internationally recognized authors
and subject matter experts have updated important aspects about Coccidiosis including detailed chapters on their biology,
phylogeny, immunity, vaccination, modern molecular biology techniques, anticoccidial drugs, and new information on the
coccidia of cattle. Coccidiosis in Livestock, Poultry, Companion Animals, and Humans will be a must have comprehensive
resource for a wide range of global scientific disciplines for upper level undergraduate and graduate students, and research-
ers that include veterinary parasitologists, veterinarians, physicians, clinicians, and animal and poultry scientists.
Dr. Daniel E. Snyder
Veterinary Parasitology Consultant
Daniel E. Snyder, DVM, PhD Consulting, LLC
This book focuses on the genera Eimeria, Cyclospora and Cystisospora, which are apicomplexan parasites, known as
conventional coccidia, and cause coccidiosis. Coccidiosis is one of the most important diseases of livestock, particularly
poultry, but occurs also in many other farm and companion animals, as well as in humans. Coccidiosis has a huge economic
impact, and represents a constant threat to the food supply. The extensive use of coccidiostats, especially against Eimeria in
poultry, has led to widespread drug resistance, and this problem needs to be solved in the years to come, as there are many
challenges ahead in order to ensure food security, but also food safety, for an ever-growing population on this planet. A
plethora of studies on different aspects of coccidian biology, phylogeny, pathogenesis and treatment and prevention strate-
gies have been carried out and published during the last 20–30 years, and it has become difficult to keep track and maintain
an overview of the research developments in the field. Thus, this book summarizes in a concise and very structured manner
the current knowledge on coccidiosis caused by conventional coccidia, and as such fills an important gap.
The driving force behind this book has been Dr. Jitender P. Dubey, which is clearly one of the most eminent parasitolo-
gists on this planet, and also known for his excellent books on Toxoplasma, Cryptosporidium, Sarcocystis and Neospora.
Here, Dr. Dubey recruited scientists of high international reputation from all over the world to contribute chapters on the
basic biology of coccidia, phylogeny, host immunity, vaccination, genomics, transcriptomics and proteomics, and on the
current state of anti-coccidial drugs. These seven chapters provide an excellent and up-to-date overview on the covered
subjects. The following chapters are dedicated to coccidiosis in the different major livestock species, including cattle, water
buffaloes, pigs, sheep, goats, and most important, poultry. However, also other host species are covered, including camelids,
rabbits, turkeys, ducks, horse and other equids, dogs, cats, and also humans. In these chapters the reader will encounter
concise information on the different species of coccidia found in the different host species, the life cycle and morphological
features, data on prevalence, on the pathogenesis, diagnosis, management and treatment options.
The book also contains stunning illustrations, showing the morphological, histological and structural features of the
different coccidia, and how infections affect their hosts and cause disease. In addition, many tables are provided that sum-
marize all this information in a concise and clear manner. Overall, I believe that this book represents an invaluable, highly
useful resource and “must-have” for all interested in coccidian parasites, including students, biologists, veterinarians, para-
sitologists, government, academia and industry.
Dr. Andrew Hemphill
Institute of Parasitology
Department of Infectious Diseases and Pathobiology
Universität Bern
Switzerland
Coccidiosis in Livestock,
Poultry, Companion
Animals, and Humans
Edited by
J. P. Dubey
CRC Press
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This book is dedicated to the memory of Drs. László P. Pellérdy (1907–1974),
Norman D. Levine (1912–1999), Datus M. Hammond (1911–1974),
Erich O. Scholtyseck (1918–1985), and Peter L. Long (1928–2005)
for their contributions to coccidiosis.
Contents
Preface..................................................................................................................................................................................... ix
Editor....................................................................................................................................................................................... xi
Contributors...........................................................................................................................................................................xiii
Chapter 1
Biology of Intestinal Coccidia.................................................................................................................................................. 1
J. P. Dubey, D. S. Lindsay, M. C. Jenkins, and C. Bauer
Chapter 2
Phylogeny of Coccidian Parasites........................................................................................................................................... 37
P. C. Thompson and B. M. Rosenthal
Chapter 3
Host Immunity in Coccidiosis................................................................................................................................................ 43
H. S. Lillehoj, W. H. Kim, and A. Chaudhury
Chapter 4
Vaccination..............................................................................................................................................................................51
M. C. Jenkins
Chapter 5
Genomics, Transcriptomics, and Proteomics of the Eimeria Species.................................................................................... 59
D. Blake and F. Tomley
Chapter 6
Anticoccidial Drugs of Livestock and Poultry Industries...................................................................................................... 65
S. Noack, H. D. Chapman, and P. M. Selzer
Chapter 7
Coccidiosis in Cattle............................................................................................................................................................... 79
B. Bangoura and A. Daugschies
Chapter 8
Coccidiosis in Water Buffaloes (Bubalus bubalis)..................................................................................................................91
J. P. Dubey
Chapter 9
Coccidiosis in Sheep............................................................................................................................................................... 99
J. M. Molina and A. Ruiz
Chapter 10
Coccidiosis in Goat (Capra hircus)...................................................................................................................................... 109
A. Ruiz and J. M. Molina
Chapter 11
Coccidiosis of Pigs................................................................................................................................................................ 125
A. Joachim and A. Shrestha
vii
viii Contents
Chapter 12
Coccidiosis in Old World Camels..........................................................................................................................................147
J. P. Dubey and R. K. Schuster
Chapter 13
Coccidiosis in South American Camelids.............................................................................................................................153
J. P. Dubey
Chapter 14
Coccidiosis in Rabbits (Oryctolagus cuniculus)....................................................................................................................159
M. Pakandl and X. Liu
Chapter 15
Coccidiosis in Chickens (Gallus gallus)................................................................................................................................169
B. Jordan, G. Albanese, and L. Tensa
Chapter 16
Coccidiosis in Poultry in China.............................................................................................................................................175
X. Suo
Chapter 17
Coccidiosis in Turkeys (Meleagris gallopavo)..................................................................................................................... 209
T. Rathinam and U. Gadde
Chapter 18
Coccidiosis in Ducks (Anas spp.)...........................................................................................................................................217
S. Wang and X. Suo
Chapter 19
Coccidiosis in Horses and Other Equids...............................................................................................................................231
C. Bauer and J. P. Dubey
Chapter 20
Coccidiosis in Dogs (Canis familiaris)................................................................................................................................. 245
J. P. Dubey and D. S. Lindsay
Chapter 21
Coccidiosis in Cats (Felis catus)........................................................................................................................................... 255
J. P. Dubey
Chapter 22
Coccidiosis in Humans......................................................................................................................................................... 267
S. Almeria, H. N. Cinar, and J. P. Dubey
References..............................................................................................................................................................................313
C. K. Cerqueira-Cézar, F. H. A. Murata, O. C. H. Kwok, and R. Calero-Bernal
Index......................................................................................................................................................................................375
Preface
Coccidiosis is one of the most important diseases of live- Domestic Animals, Long, CRC Press, 1990), and many of
stock, particularly poultry. Billions of dollars are spent their authors are now deceased. Furthermore, in the last three
on prevention worldwide to minimize coccidiosis in poul- books by Dr. Long, a lot of text was related to Sarcocystis and
try. Traditionally, coccidiosis was considered an intestinal Toxoplasma. Thus, there is a need for a book dealing exclu-
disease, caused by several distinct Eimeria species. The sively with conventional coccidia (Eimeria, Cyclospora,
disease is so important and pervasive that until recently and Cystoisospora). Here, I enlisted scientists with interna-
all poultry feed was medicated with anticoccidial drugs, tional repute and specialized knowledge to write chapters
mainly antibiotics. With the rapid development of drug on all major livestock species including cattle, pigs, sheep,
resistance, and the increasing concern of feeding antibiot- and goats. Special emphasis is given to poultry coccidiosis
ics to food animals and potential impact on public health, because of economic impact. There are chapters on phylog-
research has been directed to finding alternative methods eny, molecular biology, host-pathogen immunobiology and
of control of coccidiosis in poultry. Until 1970, coccidian immunoprophylaxis, vaccines, genetics and genomics, biol-
parasites were considered host specific and confined to ogy, and chemotherapy. There is a special chapter on coc-
mainly the intestine. With the discovery of Toxoplasma as a cidiosis in chickens in China because most of the literature is
coccidian of cats and infections in all warm-blooded hosts in Chinese and often in local journals. There is also a chapter
and subsequent discovery of Sarcocystis, and Neospora as on intestinal coccidiosis in humans, including Cyclospora.
coccidian parasites with generalized infections, research We provide concise, authoritative, up-to-date information
interest and activities in “conventional” coccidia, Eimeria, on coccidiosis, especially research in the last 28 years. Each
declined because of lack of funding and scientific research- host species chapter includes information on different spe-
ers. There has been an explosion of research on non-Eime- cies of coccidia, life cycle and biology, prevalence, epidemi-
ria coccidians in the last 40 years. I have been involved in ology, pathogenesis, diagnosis, and treatment. All references
writing books on three subjects published by CRC Press: are alphabetized by the first author and appear at the end of
Toxoplasma (Toxoplasmosis of Animals and Humans, first the book for the benefit of readers. Originals of each research
edition, Dubey and Beattie, 1988; second edition, Dubey, papers were obtained to avoid mistakes in citations.
2010), Sarcocystis (Sarcocystosis of Animals and Humans, It is hoped that this book will be useful to biologists,
first edition, Dubey, Speer, and Fayer, 1989; second edition, veterinarians, parasitologists, and researchers from govern-
Dubey, Calero-Bernal, Rosenthal, Speer, and Fayer, 2016), ment, academia, and industry.
and Neospora (Neosporosis in Animals, Dubey, Hemphill, I would like to acknowledge those who made this pos-
Calero-Bernal, and Schares, 2017). Additionally, I edited a sible; I cannot possibly list all of them. Christian Bauer from
multiauthored book on Cryptosporidium (Cryptosporidiosis Germany and David Lindsay from the United States, and
in Man and Animals, Dubey, Speer, and Fayer, 1990). Camila Cezar, Fernando Murata, Meghan Sadler, and Oliver
All books on conventional coccidiosis were writ- Kwok from my laboratory worked tirelessly to make this
ten more than 25 years ago (The Coccidia, Hammond and book possible.
Long, University Park Press, 1973; The Biology of Coccidia,
Long, Edward Arnold Press, 1982; Coccidiosis of Man and J. P. Dubey
ix
Editor
J. P. Dubey, MVSc, PhD, DSc, Sarcocystis, and related cyst-forming coccidian parasites of
was born in India. He earned his humans and animals. He has published over 1500 research
veterinary degree in 1960 from papers in international journals. In 1985, he was chosen
Veterinary College, Mhow, India to be the first recipient of the Distinguished Veterinary
and masters in veterinary parasi- Parasitologist Award by the American Association of
tology in 1963, from Veterinary Veterinary Parasitologists. Dr. Dubey is the recipient of the
College, Mathura, India. He 1995 WAAVP Pfizer Award for outstanding contributions
earned a PhD in medical microbi- to research in veterinary parasitology. He also received the
ology in 1966 from the University 2005 Eminent Parasitologists Award by the American Society
of Sheffield, England. Dr. Dubey of Parasitologists. The Thomson Institute for Scientific
received postdoctoral training Information identified him as one of the world’s most cited
from 1968 to 1973 with Dr. J.K. authors in plant and animal sciences for the last decade. In
Frenkel, Department of Pathology and Oncology, University 2003, he was selected for the newly created Senior Science and
of Kansas Medical Center, Kansas City, Kansas. From 1973 to Technology Service (SSTS) and is one of the few such scien-
1978, he was an associate professor of veterinary parasitology, tists and executives within the USDA’s Agricultural Research
Department of Pathobiology, Ohio State University, Columbus, Service (USDA-ARS); selection for this position is by invi-
Ohio, and professor of veterinary parasitology, Department tation only, on approval by the Secretary of Agriculture. In
of Veterinary Science, Montana State University, Bozeman, 2010, Dr. Dubey was elected to the U.S. National Academy
Montana, from 1978 to 1982. He is presently a senior scientist, of Sciences, Washington, DC, and inducted in the USDA-
Animal Parasitic Diseases Laboratory, Beltsville Agricultural ARS Hall of Fame. In 2018, he was the first recipient of the
Research Center, Agricultural Research Service, United States William C. Campbell One-Health Award by the American
Department of Agriculture (USDA), Beltsville, Maryland. Association of Veterinary Parasitologists. In the same year, he
Dr. Dubey has spent more than 55 years researching coc- received the honorary Doctor of Science degree from McGill
cidian parasites, including Eimeria, Toxoplasma, Neospora, University, Montréal, Québec, Canada.
xi
Contributors
G. Albanese H. N. Cinar
Departments of Population Health and Poultry Science U.S. Food and Drug Administration
Poultry Diagnostic and Research Center Laurel, Maryland
University of Georgia
Athens, Georgia A. Daugschies
Institute for Parasitology
S. Almeria Leipzig University
Food and Environmental Microbiology Leipzig, Germany
Office of Applied Research and Safety Assessment
Food and Drug Administration
J. P. Dubey
Laurel, Maryland
U.S. Department of Agriculture
B. Bangoura Agricultural Research Service
Wyoming State Veterinary Laboratory Animal Parasitic Diseases Laboratory
Department of Veterinary Sciences Beltsville, Maryland
University of Wyoming
Laramie, Wyoming U. Gadde
Huvepharma Inc.
C. Bauer Peachtree City, Georgia
Institute of Parasitology
Justus Liebig University Giessen M. C. Jenkins
Giessen, Germany U.S. Department of Agriculture
Agricultural Research Service
D. Blake Animal Parasitic Diseases Laboratory
Parasite Genetics, Pathobiology and Population Sciences Beltsville, Maryland
Royal Veterinary College
North Mymms A. Joachim
Hertfordshire, United Kingdom Institute of Parasitology, Department of Pathobiology
Veterinärmedizinische Universität Wien/Vetmeduni
R. Calero-Bernal Vienna
Department of Animal Health Wien, Austria
Complutense University of Madrid
Madrid, Spain
B. Jordan
C. K. Cerqueira-Cézar Departments of Population Health and
Oak Ridge Institute for Science and Education (ORISE) Poultry Science
Research Fellow Poultry Diagnostic and Research Center
U.S. Department of Agriculture University of Georgia
Agricultural Research Service Athens, Georgia
Animal Parasitic Diseases Laboratory
Beltsville, Maryland W. H. Kim
Oak Ridge Institute for Science and Education (ORISE)
H. D. Chapman Research Fellow at the Animal Biosciences and
Department of Poultry Science Biotechnology Laboratory
University of Arkansas U.S. Department of Agriculture
Fayetteville, Arkansas Agricultural Research Service
Beltsville, Maryland
A. Chaudhury
Oak Ridge Institute for Science and Education (ORISE) O. C. H. Kwok
Research Fellow at the Animal Biosciences and U.S. Department of Agriculture
Biotechnology Laboratory Agricultural Research Service
U.S. Department of Agriculture Animal Parasitic Diseases Laboratory
Beltsville, Maryland Beltsville, Maryland
xiii
xiv Contributors
H. S. Lillehoj A. Ruiz
Poultry Immunology, Genomics and Disease Resistance Departmento de Patologia Animal
Animal Biosciences and Biotechnology Laboratory Produccion Animal, Bromatologia y Technologia de los
U.S. Department of Agriculture Alimentos
Agricultural Research Service University de Las Palmas de Gran Canaria
Beltsville, Maryland Las Palmas, Spain
D. S. Lindsay R. K. Schuster
Department of Biomedical Science and Pathology FTA Parasitology, FTA Tropical Veterinary Medicine
Virginia Maryland College of Veterinary Medicine Central Veterinary Research Laboratory
Center for One Health Research Dubai, United Arab Emirates
Virginia Tech
Blacksburg, Virginia
P. M. Selzer
Molecular Discovery—Pharma R&D
X. Liu
Boehringer Ingelheim Vetmedica GmbH
College of Veterinary Medicine
Ingelheim am Rhein, Germany
China Agricultural University
Beijing, China
A. Shrestha
J. M. Molina Institute of Parasitology
University of Las Palmas de G. C. Department of Pathobiology
Faculty of Veterinary Medicine. Parasitology Unit Veterinärmedizinische Universität Wien/Vetmeduni
Las Palmas de Gran Canaria, Spain Vienna
Wien, Austria
F. H. A. Murata
Oak Ridge Institute for Science and Education (ORISE) X. Suo
Research Fellow College of Veterinary Medicine
U.S. Department of Agriculture China Agricultural University
Agricultural Research Service Beijing, China
Animal Parasitic Diseases Laboratory
Beltsville, Maryland L. Tensa
Departments of Population Health and Poultry Science
S. Noack Poultry Diagnostic and Research Center
Molecular Discovery—Pharma R&D University of Georgia
Boehringer Ingelheim Vetmedica GmbH Athens, Georgia
Ingelheim am Rhein, Germany
P. C. Thompson
M. Pakandl
U.S. Department of Agriculture
Department of Research and Development
Agricultural Research Service
BIOPHARM, Research Institute of Biopharmacy and
Animal Parasitic Diseases Laboratory
Veterinary Drugs
Beltsville, Maryland
Prague, Czech Republic
T. Rathinam F. Tomley
Huvepharma Inc. The Royal Veterinary College
Peachtree City, Georgia North Mymms, Hatfield
Hertfordshire, United Kingdom
B. M. Rosenthal
U.S. Department of Agriculture S. Wang
Agricultural Research Service College of Veterinary Medicine
Animal Parasitic Diseases Laboratory China Agricultural University
Beltsville, Maryland Beijing, China
Chapter 1
Biology of Intestinal Coccidia
J. P. Dubey, D. S. Lindsay, M. C. Jenkins, and C. Bauer
CONTENTS
1.1 Introduction and Brief History........................................................................................................................................ 1
1.2 Genus Eimeria (syn. Coccidium)..................................................................................................................................... 2
1.2.1 Life Cycle of Eimeria.......................................................................................................................................... 2
1.2.1.1 Oocyst................................................................................................................................................... 2
1.2.1.2 Infection of the Host............................................................................................................................. 5
1.2.1.3 Asexual Development........................................................................................................................... 5
1.2.1.4 Sexual Development............................................................................................................................. 6
1.2.1.5 Extraintestinal Coccidia....................................................................................................................... 6
1.2.1.6 Description of Eimeria Species............................................................................................................ 6
1.3 Genus Cystoisospora....................................................................................................................................................... 8
1.4 Ultrastructure..................................................................................................................................................................14
1.4.1 Eimeria Species................................................................................................................................................. 22
1.4.1.1 Oocysts, Sporocysts, Sporozoites....................................................................................................... 22
1.4.1.2 Schizonts and Merozoites................................................................................................................... 22
1.4.1.3 Microgamonts and Macrogamonts..................................................................................................... 24
1.4.2 Cystoisospora.................................................................................................................................................... 25
1.5 In Vitro Cultivation........................................................................................................................................................ 25
1.6 Pathogenesis and Lesions of Intestinal Coccidiosis...................................................................................................... 27
1.7 Diagnosis....................................................................................................................................................................... 29
1.8 Control........................................................................................................................................................................... 36
References............................................................................................................................................................................... 36
1.1 INTRODUCTION AND BRIEF HISTORY husbandry practices, such as better management, hygiene,
and sanitation, it is still almost impossible to raise live-
Coccidia have long been recognized as one of the most stock coccidia free.
important groups of parasites that infect animals, espe- Until the discovery of the two-host (polyxenous) life
cially domestic livestock. Most of the stages of the life cycle cycle of Toxoplasma gondii in 1970, conventional coc-
require a microscope for visualization, but some accumu- cidia of the genus Eimeria were believed to have a simple
lations of stages cause gross lesions that can be observed oro-fecal one-host transmission life cycle with infections
in the intestinal tract. Coccidia were not discovered until confined mostly to the intestines of the host. T. gondii is a
the invention of the microscope by Leeuwenhoek, who tissue-dwelling coccidian that uses all warm-blooded ani-
probably saw coccidian life-cycle stages (now recognized mals, including humans, as intermediate hosts, but only
as Eimeria stiedai) in the liver of a rabbit.973–975 However, felids can serve as the definitive host.420 Since the discovery
their economic importance was not realized until the early of the life cycle of T. gondii, a heteroxenous development
1900s when outbreaks of bloody enteritis, caused by the has been demonstrated in several other coccidia as well. As
coccidian E. tenella, were observed in the ceca of chick- stated in the preface, this book is concerned with those coc-
ens.228,232,1717 Despite improvements in modern animal cidia that have a direct life cycle completed in a single host,
