Parasitic Protozoa of Farm Animals and Pets Monica Florin-Christensen Leonhard Schnittger Editors 123 Parasitic Protozoa of Farm Animals and Pets Monica Florin-Christensen Leonhard Schnittger Editors Parasitic Protozoa of Farm Animals and Pets Editors Monica Florin-Christensen Leonhard Schnittger Center for Research on Veterinary Center for Research on Veterinary and Agronomic Sciences and Agronomic Sciences National Institute of Agricultural National Institute of Agricultural Technology Technology Hurlingham, Argentina Hurlingham, Argentina National Council of Scientific and National Council of Scientific and Technological Research (CONICET) Technological Research (CONICET) Buenos Aires, Argentina Buenos Aires, Argentina School of Exact, Chemical and Natural School of Exact, Chemical and Natural Sciences (FCEQN), University of Morón Sciences (FCEQN), University of Morón Morón, Province of Buenos Aires Morón, Province of Buenos Aires Argentina Argentina ISBN 978-3-319-70131-8 ISBN 978-3-319-70132-5 (eBook) https://doi.org/10.1007/978-3-319-70132-5 Library of Congress Control Number: 2017963966 © Springer International Publishing AG, part of Springer Nature 2018 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. The publisher remains neutral with regard to jurisdictional claims in published maps and institutional affiliations. Printed on acid-free paper This Springer imprint is published by the registered company Springer International Publishing AG part of Springer Nature The registered company address is: Gewerbestrasse 11, 6330 Cham, Switzerland Dedicatory To our children, Nicolas, Ana, Julieta, and Uriel Contents 1 Introduction into Parasitic Protozoa . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1 Leonhard Schnittger and Monica Florin-Christensen 2 Cryptosporidium . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11 Mariela L. Tomazic, Carlos Garro, and Leonhard Schnittger 3 Eimeria . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 55 Berit Bangoura and Arwid Daugschies 4 Sarcocystis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 103 Cecilia Decker Franco, Leonhard Schnittger, and Monica Florin-Christensen 5 Neospora . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 125 Dadin Prando Moore and Maria Cecilia Venturini 6 Toxoplasma . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 149 Gastón Moré, Maria Cecilia Venturini, Lais Pardini, and Juan Manuel Unzaga 7 Besnoitia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 169 Heinz Mehlhorn 8 Theileria in Ruminants . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 187 Henry Kiara, Lucilla Steinaa, Vishvanath Nene, and Nicholas Svitek 9 Babesia in Domestic Ruminants . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 215 Sabrina Ganzinelli, Anabel Rodriguez, Leonhard Schnittger, and Mónica Florin-Christensen 10 Babesia of Domestic Dogs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 241 Gad Baneth 11 Equine Piroplasmids . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 259 Massaro W. Ueti and Donald P. Knowles 12 Trypanosoma . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 271 Silvina E. Wilkowsky vii viii Contents 13 Leishmania . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 289 Anabel Elisa Rodriguez, José Octavio Estévez, María Cecilia Nevot, Alejandra Barrios, and Monica Florin-Christensen 14 Trichomonas . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 313 Esther Collántes-Fernández, Marcelo C. Fort, Luis M. Ortega-Mora, and Gereon Schares 15 Amoebae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 389 Julia Walochnik 16 Genomics and Genetic Manipulation of Protozoan Parasites Affecting Farm Animals . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 413 Carlos E. Suarez, Heba F. Alzan, and Brian M. Cooke About the Editors Monica  Florin-Christensen and Leonhard  Schnittger are biologists and researchers of the National Council of Scientific and Technological Research of Argentina (CONICET). They lead a research group that investigates on molecular parasitology, focusing on pathogenic protists of veterinary importance, at the Institute of Pathobiology, National Institute of Agricultural Technology (INTA, Argentina). Both are Professors of molecular biology and population genetics at the School of Exact, Chemical and Natural Sciencies, University of Moron, Argentina. ix Introduction into Parasitic Protozoa 1 Leonhard Schnittger and Monica Florin-Christensen Abstract Parasitic protozoans are protists—i.e., unicellular eukaryotes. Besides, they share the common features of being heterotrophic, motile in at least one of their stages, and dependent on a host for survival. Protists are a highly diverse evolu- tionary unrelated grouping that is at present taxonomically divided into at least five supergroups or eukaryotic kingdoms: SAR, Excavata, Amoebozoa, Archaeplastida, and Ophistokonta. The protozoan parasites that infect farm ani- mals and pets, and are the subject of this book, belong to the first three super- groups. These organisms can cause significant morbidity and mortality in domestic animals, thus provoking substantial economic losses associated with livestock production or major concern for pet owners. Additionally, some are zoonotic and of public health relevance. Human domestication has led to a huge world population of a few selected farm and pet animals, usually raised in close proximity. This has provided an ideal and spacious ecological niche for parasitic protozoans that invade and thrive in these hosts. In addition, international travel has promoted global distribution of most parasitic protozoa. A long history of coadaptation has existed between parasites and their wild ancestor hosts, whereas adaptation to domestic animals has been much shorter from an evolutionary per- spective. While the first scenario resulted in unapparent infections that ensured L. Schnittger (*) • M. Florin-Christensen Center for Research on Veterinary and Agronomic Sciences National Institute of Agricultural Technology Hurlingham, Argentina National Council of Scientific and Technological Research (CONICET), Buenos Aires, Argentina School of Exact, Chemical and Natural Sciences (FCEQN), University of Morón, Morón, Province of Buenos Aires, Argentina e-mail: [email protected] © Springer International Publishing AG, part of Springer Nature 2018 1 M. Florin-Christensen, L. Schnittger (eds.), Parasitic Protozoa of Farm Animals and Pets, https://doi.org/10.1007/978-3-319-70132-5_1 2 L. Schnittger and M. Florin-Christensen parasite perpetuation, the second represents an unbalanced situation resulting in clinical signs and mortality. This calls for continuous efforts in the development and application of control measures. 1.1 Protozoa as a Subgroup of Protists Protozoa represent a subgroup of the protists. Then, what are protists? Originally, the name referred to all eukaryotic unicellular organisms that were joined into the taxo- nomic kingdom Protista in the year 1866 by Ernst Haeckel. The group of protists was further subdivided into (1) animal-like protozoans, i.e., heterotrophic/parasitic motile eukaryotes; (2) plantlike protophyta or algae, i.e., autotrophic unicellular eukaryotes; and (3) fungus-like saprophytes, i.e., unicellular eukaryotic decomposers with extracel- lular digestion and spore production. However, with the advancement of science, it became clear that these subdivisions are artificial, as they do not reflect valid taxonomic groups based on evolutionary relatedness. Nevertheless, all protists that are non-photo- synthetic and non-fungus-like are, for convenience, often referred to as protozoans. The simplest definition of a protist is exclusive: a eukaryote that is neither an ani- mal nor a plant or a fungus. However, based on genetic similarities or differences, protists represent a paraphyletic assembly, i.e., an artificial collection of diverse organ- isms—possibly only connected to the last common eukaryotic ancestor—that spans the entire eukaryotic tree and consists of at least five monophyletic supergroups. Each of these supergroups represents a taxonomic kingdom that is more diverse than and different from the traditional taxonomic kingdoms of animals, plants, or fungi. Possibly, protists are the least understood group of organisms as compared to prokary- otes, animals, fungi, and plants. This is likely because they have undergone by far the most extensive, continuous, and ongoing changes in their classification and naming over the last decades, making it difficult to approach this group by anybody than the expert. Consequently, protists have been gradually omitted from most biology text- books, as noticed by Adl et al. (2007). Protists display an immense morphological variety at a cellular level, reflecting the extensive capacity of the eukaryotic cell to evolve and adapt to an enormous diversity of ecological niches. This morphological variety is, in contrast to higher animals and plants, only accessible—i.e., can only be visualized and studied—by the use of light or electron microscopy. Importantly, mor- phological characteristics of protists that can be studied by light microscopy—e.g., an amoeboid cell form—are often counterintuitive with regard to their phylogenetic ori- gin as they may represent convergent evolutionary developments. They are therefore of limited value to classify and scientifically name these organisms. In contrast, ultra- structural differences that can only be appreciated by electron microscope studies— e.g., the apical complex of apicomplexan protozoans—reveal character traits that often allow recognition and delineation of phylogenetically related groups.