The genus Gregarina was created by Dufour in 1828 for "un nouveau genre de ver qui vit en troupeau dans les intestine de divers insectes". These unicellular parasites were discovered before the cell theory was developed by Schwann & Schleiden in 1839 and completed by Virchow in 1855, and therefore, Dufour supposed that they were helminths. He named Gregarina ovata and G. cuneata, the two first known species found in earwigs and beetles respectively. Many other species were further described over the whole world, aided by the easy detection of their large extracellular stages, even with simple binocular magnifiers, and by their abundance in a wide range of invertebrates, especially insects. The first major contributions to our knowledge of gregarines are due to Moniker (1850-60), Lankaster (1860-85), Schneider (1875), Min-gazzini (1891-93), and Siedlecki (1899) including the first observation of the process of fertilization in the cyst of Lankesteria ascidae. The occurrence of Microsporidia inside marine gregarines was established by Caullery & Mesnil (1897-1919). Leger (1904-1910) accurately described the gametogenesis of Stylocephalus longicollis and other species. In the first part of the zoth century, the occurrence of an asexual schizogonic process was demonstrated in marine gregarines by Brasil (1904-1910). Various classifications were proposed by Pixell-Goodrich (1915-1950), Ray (1930), Ganapati and coworkers (1937-46), Kamm (1922), Reichenow (1929-32) and other authors.
The chapter Gregarines written in 1953 by Pierre-Paul Grasse (Traite de Zoologie, Tome I, Masson Ed.) was the masterwork of what was known about this parasitic group at the middle of the zoth century, both in its biology, cytology at the light micro-scopic level and systematics. Since then, the development of methodological tools has considerably improved our knowledge of Gregarines and their place in the evolution of the Apicomplexa. The present revised and updated edition of the chapter Gre-garines covers more than 6o years of publications, recording scientific investigations conducted in both cell biology and systematics of marine and terrestrial gregarines. In the early 6o's, electron microscopy became the determinant for defining the cellu-lar organization of these parasites. During that period, two main schools emerged in France, one focused on the ultrastructure of marine gregarines under the leadership of Emile Vivier at Lille University, the other on the ultrastructure of terrestrial gregarines under the direction of Pierre-Paul Grasse in the Laboratory 'Evolution des titres organ-ises' (Faculty of Sciences, Paris). Since 1953, gregarine systematics were emphasized by Geus (1969) and Levine (1988) in book chapters and more recently by Clopton & Perkins (z000) in the Illustrated Guide of Protozoa (Lee, Leedale & Bradbury Ed.).
The Apicomplexa (Levine, 1970) are able to parasitise the majority of animal taxa due to their ability to develop outside and also inside host cells. The extracellular de-velopment of most gregarines explains their morphological variability and the pre-dominance of their sexual reproduction with many gametes in large size gamonts. In contrast the Coccidia and Hematozoa are able to enter and develop intracellularly in their vertebrate hosts, while their pathogenicity appears as a consequence of their asexual proliferation, a process additional to the formation of small intracellular gamonts. The systematics of gregarines is based on morphological, biological and molecular criteria that appear to be correlated with their localization in their hosts and to reflect their adaptation to the large diversity of environments, as well as the feeding behaviours and development of their invertebrate hosts. The emphasizing on the distribution of the different gregarines taxa in marine, freshwater and terrestrial hosts is therefore an important theme of this publication. The data presented in chap-ters in and iv is intended to facilitate the identification of species encountered by those who do not specialise in these parasites.
An important part of the information herein reported was collected from 1970 to 1980 by Jean Theodorides for a supplement of the first edition of the Traite de Zoologie. The supplement remained unpublished after the death of P. P. Grasse, in 1985. This volume is dedicated to the memory of Jean Theodorides (1927-1999) and Emile Vivier (1923-2on), for their contribution to the knowledge of Gregarines.

VOLUME 1

PREFACE 1

ACKNOWLEDGMENTS 2

CHAPTER I INTRODUCTION: GREGARINES AMONG APICOMPLEXA 7

      The high morphological diversity of the vegetative stages of gregarine

      trophozoites 12

      Schizogony and taxonomy of gregarines 13

      Life cycles of gregarines 14

      A. Life cycle of a Eugregarine: Lecudina tuzetae 14

      B. Life cycle of an Archigregarine: Selenidium pendula, the type species 18

      C. Life cycle of a Neogregarine: Mattesia oryzaephili 18

      Bibliography 21

CHAPTER II BIOLOGY OF GREGARINES AND THEIR HOST-PARASITE INTERACTIONS 25

      II.1 Biology of gregarines 26

      II.1.1 Cell and molecular organization of vegetative gregarines: the trophozoites 26

      II.1.1.1 Cell and molecular organization of the cortex of trophozoites: the epicyte 26

      II.1.1.2 Cytoskeleton and cell motility 54

      II.1.1.3 Diversity in nutrition: cell surface, mucron, epimerite 71

      II.1.1.4 Cytoplasmic organelles of trophozoites 87

      II.1.2 Syzygy and cell recognition between the gamonts 102

      II.1.2.1 Types of syzygy 102

      II.1.2.2 Syzygy with multiple associations 105

      II.1.2.3 Cell junctions between gamonts in syzygy 706

      II.1.2.4 Cell recognition and sexual differences between gamonts 108

      II.1.3 Biology and cell organization of gregarines during gamogony 110

      II.1.3.1 Rotational movements and secretions during cyst formation 110

      II.1.3.2 Syncytial nuclear divisions 172

      II.1.3.3 Cellularization 176

      II.1.3.4 Gametes and fertilization 178

      II.1.3.5 Centriole/basal body in Eukaryotes and the mToc/basal body structures

      in gregarines 133

      II.1.3.6 Abnormal encystment: parthenogenesis? 135

      II.1.4 Biology and cell organization of gregarines during sporogony 136

      II.1.4.1 Zygote, meiosis and metagamic mitosis 136

      II.1.4.2 Secretion of the sporoblast wall and cytoplasmic events within the

      sporoblast 139

      II.1.4.3 Biogenesis of sporozoites 141

      II.1.4.4 Importance of spore structures in taxonomy of Gregarines 144

      II.1.4.5 Dissemination of the gregarine spores 144

      II.1.5 Transition from sporozoite to trophozoite 150

      II.2 Diversity of host-parasite interactions 158

      II.2.1 Behaviour of gregarines 158

      II.2.1.1 Habitat 158

      II.2.1.2 Host specificity 160

      II.2.1.3 Effects of gregarines on their hosts 167

      II.2.2 Electrophysiological properties of the cortical membranes in gregarine trophozoites 162

      II.2.3 Endosymbionts and hyperparasitemia with Microsporidia 162

      II.2.3.1 Bacteria 162

      II.2.3.2 Microsporidia 162

      II.2.4 Experimental control of gregarine life cycles 172

      II.2.4.1 Marine gregarines 172

      II.2.4.2 Terrestrial gregarines 173

      II.2.5 Relationships between host and parasite life cycles and chronology of

      gregarine life cycles 173

      II.2.5.1 Marine gregarines 173

      II.2.5.2 Terrestrial gregarines 175

      Bibliography 178

CHAPTER III MARINE GREGARINES 197

      III.1 General presentation 197

      III.1.1 Definition of Aseptatorina and Septatorina 797

      III.1.2 Intermediates between the Aseptatorina and Septatorina 798

      III.1.3 Gregarines of Polychaeta and other marine Lophotrochozoa 200

      III.1.4 Gregarines of marine Acoela, Nemertodermatida flatworms, and

      Chaetognatha 203

      III.1.5 Gregarines in the marine Ecdysozoa: crustaceans 203

      III.1.6 Gregarines of Echinodermata and Hemichordata: the marine

      Deuterostomia Xenambulacria 205

      III.1.7 Gregarines of Tunicata: the marine Deuterostomia Urochordata 205

      III.2 Archigregarinorida Grasse, 7953: the Archigregarines 206

      III.3 Eugregarinorida Leger, 7900 240

      III.3.1 Eugregarinida Aseptatorina Chalcravarty, 1960 240

      III.3.1.1 Coelomic Eugregarinorida Aseptatorina 240

      III.3.1.2 Intestinal Eugregarinorida Aseptatorina 266

      III.3.2 Eugregarinida Septatorina Lankester, 1885 325

      Bibliography 355

VOLUME II

CHAPTER IV SYSTEMATICS OF TERRESTRIAL AND FRESH WATER

GREGARINES 377

      Introduction 377

      Classification - diagnosis of families 377

      Order Eugregarinorida Leger, rgoo 377

      Suborder ASEPTATORINA Chakravarty, 1960 378

      Suborder SEPTATORINA Lankester, 1885 380

      Order Neogregarinorida Grasse,1953 (Schizogregarinida Léger, rgoo) 389

      IV.1 Gregarines of Platyhelminthes: Turbellaria 394

      IV.2 Gregarines ofAnnelids: Clitellata 398

      IV.2.1 Gregarines of Hirudinea 398

      IV.2.2 Gregarines of Oligochaeta 399

      IV.3 Gregarines of Chelicerata: Arachnida 427

      IV.3.1 Gregarines of Acari (Oribatida) 428

      IV.3.2 Gregarines of Opiliones 431

      IV.4 Gregarines of Myriapoda 436

      IV.15 Gregarines of Hexapoda: Entognatha, Insecta 467

      IV.5.1 Gregarines of Entognatha 467

      IV5.2 Gregarines of primitive insects 469

      IV.5.2.1 Gregarines of Archaeognatha 469

      IV.5.2.2 Gregarines of Zygentoma 472

      IV.5.3 Gregarines of Odonata 474

      IV.5.4 Gregarines of Ephemeroptera 488

      IV.5.5 Gregarines of Blattaria 492

      IV.5.6 Gregarines of Isoptera 498

      IV.5.7 Gregarines of Plecoptera 504

      IV.5.8 Gregarines of Orthoptera 506

      IV.5.9 Gregarines of Dermaptera 530

      IV.5.10 Gregarines of Embioptera 530

      IV.5.11 Gregarines of Psocoptera 532

      IV.5.12 Gregarines of Hemiptera 533

      IV.5.13 Gregarines of Coleoptera 536

      IV.5.14 Gregarines of Neuroptera (= Nevroptera) 630

      IV.5.15 Gregarines of Raphidioptera 630

      IV.5.16 Gregarines of Hymenoptera 637

      IV.5.17 Gregarines of Mecoptera 634

      IV.5.18 Gregarines of Siphonaptera 635

      IV.5.19 Gregarines of Diptera 638

      IV.5.20 Gregarines of Trichoptera 654

      IV.5.21 Gregarines of Lepidoptera 664

      Bibliography 672

CREDITS 711

INDEX ON TAXONOMICAL NAMES 715

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