Handbook of Zoology: Volume 3 Gastrotricha and Gnathifera

This section of the Handbook of Zoology is intended as a comprehensive and exhaustive account of the biology of the taxa Gastrotricha, Nematoda, Nematomorpha, Priapulida, Kinorhyncha, Loricifera, Gnathostomulida, Micrognathozoa, Rotifera, Seisonida and Acanthocephala, covering all relevant topics such as morphology, ecology, phylogeny and diversity. The series is intended to be a detailed and up-to-date account of these taxa. As was the case with the first edition, the Handbook is intended to serve as a reliable resource for decades. Many of the taxa of this volume are comparatively unknown to many biologists, despite their diversity and importance for example in meiofaunal communities (Gastrotricha, Rotifera, Gnathostomulida), their fascinating recent discoveries (Loricifera and Micrognathozoa), their importance as parasites (many nematodes, Nematomorpha, Acanthocephala) and their importance for evolutionary questions (e.g. Priapulida, Gastrotricha). The groups covered range from those poor in species (such as Micrognathozoa with 2 known species) to the species-rich and diverse Nematoda and their ca. 20.000 described species. While each taxon is covered by one chapter, nematodes are treated in several chapters dedicated to their structural, taxonomic and ecological diversity. * Comprehensive account of the morphology, ecology, phylogeny and diversity of theGastrotricha and Gnathifera * Including newly described Micrognathozoa * Taxa with important roles in marine meiofaunal communities * Recent discoveries and evolutionary questions treated List of contributing authors 1 Gastrotricha 1.1 Introduction 1.2 Morphology 1.2.1 General and external morphology 1.2.2 Integument 1.2.3 Musculature 1.2.4 Nervous system 1.2.5 Sensory structures 1.2.6 Intestinal system 1.2.7 Body cavities and connective tissue 1.2.8 Excretory system 1.2.9 Reproductive organs 1.2.9.1 Female gonad 1.2.9.2 Male gonad 1.2.9.3 Frontal and caudal organ 1.2.9.4 Reproductive system of the Paucitubulatina 1.2.9.5 Additional accessory structures 1.2.10 Gametes 1.2.10.1 Spermatozoa 1.2.10.2 Spermatogenesis and spermiogenesis 1.2.10.3 Eggs 1.2.10.4 Oogenesis 1.3 Reproduction and development 1.3.1 Reproductive biology 1.3.2 Cleavage and development 1.4 Physiology 1.5 Phylogeny 1.6 Systematics 1.6.1 Order Macrodasyida Brunson, 1950 1.6.1.1 Family Cephalodasyidae Hummon & Todaro, 2010 1.6.1.2 Family Dactylopodolidae Strand, 1929 1.6.1.3 [Family Lepidodasyidae Remane, 1927] 1.6.1.4 Family Macrodasyidae Remane, 1926 1.6.1.5 Family Planodasyidae Chandrasekhara Rao & Clausen, 1970 1.6.1.6 Family Thaumastodermatidae Remane, 1927 1.6.1.7 Family Turbanellidae Remane, 1927 1.6.1.8 Family Xenodasyidae Todaro, Guidi, Leasi & Tongiorgi, 2006 1.6.1.9 Family Redudasyidae Todaro, Dal Zotto, Jondelius, Hochberg, Hummon, Kånneby & Rocha, 2012 1.6.2 Order Chaetonotida Brunson, 1950 1.6.2.1 [Suborder Multitubulatina d’Hondt, 1971] 1.6.2.1.1 [Family Neodasyidae Remane, 1929] 1.6.2.2 Suborder Paucitubulatina d’Hondt, 1971 1.6.2.2.1 Family Muselliferidae Leasi & Todaro, 2008 1.6.2.2.2 Family Xenotrichulidae Remane, 1936 1.6.2.2.3 Family Chaetonotidae Gosse, 1864 1.6.2.2.4 Family Dasydytidae Daday, 1905 1.6.2.2.5 Family Neogosseidae Remane 1927 1.6.2.2.6 [Family Dichaeturidae Remane, 1927] 1.6.2.2.7 Family Proichthydidae Remane 1927 1.6.2.3 Gastrotricha incertae sedis 1.7 Biogeography 1.8 Ecology Acknowledgments Literature 2 Phylum Gnathostomulida 2.1 Introduction and history of research (Fig. 2.1) 2.2 Morphology 2.2.1 General and external morphology (Figs. 2.2–2.6) 2.2.2 Integument (Figs. 2.7 and 2.8) 2.2.3 Musculature (Figs. 2.9 and 2.10) 2.2.4 Nervous system (Fig. 2.11) 2.2.5 Sensory structures (Fig. 2.12) 2.2.6 Intestinal system (Figs. 2.13–2.36) 2.2.6.1 Pharynx 2.2.6.2 Jaws 2.2.6.3 Basal plate and jugum 2.2.7 Body cavities and connective tissue 2.2.8 Excretory system (Fig. 2.37) 2.2.9 Reproductive organs (Figs. 2.38–2.41) 2.2.9.1 Female organs 2.2.9.2 Male organs 2.2.10 Gametes 2.2.10.1 Eggs and oogenesis 2.2.10.2 Sperm and spermiogenesis (Figs. 2.39 E and 2.42–2.46) 2.2.10.2.1 Spermiogenesis 2.3 Reproduction and development 2.3.1 Reproductive biology 2.3.2 Oviposition and cleavage 2.3.3 Development (Fig. 2.47) 2.4 Biology and physiology (Figs. 2.48 and 2.49 2.5 Ecology 2.6 Phylogeny 2.7 Systematics 2.7.1 Phylum Gnathostomulida (Ax, 1956) Riedl, 1969 2.7.1.1 Order Filospermoidea Sterrer, 1972 2.7.1.1.1 Family Haplognathiidae, Sterrer 1972 2.7.1.1.2 Family Pterognathiidae Sterrer, 1972 2.7.1.2 Suborder Conophoralia Sterrer, 1972 2.7.1.2.1 Family Austrognathiidae Sterrer, 1971 2.7.1.3 Order Bursovaginoidea Sterrer, 1972 2.7.1.4 Suborder Scleroperalia Sterrer, 1972 2.7.1.4.1 Family Gnathostomariidae Sterrer, 1972 2.7.1.4.2 Family Rastrognathiidae Kristensen & Nørrevang, 1977 2.7.1.4.3 Family Agnathiellidae Sterrer, 1972 2.7.1.4.4 Family Mesognathariidae Sterrer, 1972 2.7.1.4.5 Family Paucidentulidae terrer, 1998 2.7.1.4.6 Family Clausognathiidae Sterrer, 1992 2.7.1.4.7 Family Problognathiidae Sterrer & Farris, 1975 2.7.1.4.8 Family Onychognathiidae Sterrer, 1972 2.7.1.4.9 Family Gnathostomulidae Sterrer, 1972 2.8 Biogeography 2.9 Paleontology Acknowledgments Literature 3 Micrognathozoa 3.1 Introduction 3.2 Morphology 3.2.1 General and external morphology 3.2.2 Integument 3.2.3 Musculature 3.2.4 Nervous system 3.2.5 Sensory structures 3.2.6 Intestinal system 3.2.7 Body cavities and connective tissue 3.2.8 Excretory system 3.2.9 Reproductive organs 3.2.10 Gametes 3.3 Reproduction and development 3.4 Physiology 3.5 Phylogeny 3.6 Systematics 3.7 Biogeography 3.8 Paleontology 3.9 Ecology Literature 4 Rotifera 4.1 Introduction 4.2 Morphology 4.2.1 General and external morphology 4.2.1.1 Morphology of the female 4.2.1.2 Coloniality 4.2.1.3 Corona 4.2.1.4 Morphology of the male 4.2.2 Integument 4.2.2.1 Pedal glands 4.2.2.2 Retrocerebral organ 4.2.2.3 Sheats and tubes 4.2.3 Musculature 4.2.3.1 Musculature of the female 4.2.3.2 Musculature of the male 4.2.4 Nervous system 4.2.5 Sensory structures 4.2.6 Intestinal system 4.2.6.1 Intestinal system of the female 4.2.6.2 Trophi 4.2.6.3 Intestinal system of the male 4.2.7 Body cavity 4.2.8 Excretory system 4.2.8.1 Excretory system of the female 4.2.8.2 Excretory system of the male 4.2.9 Reproductive organs 4.2.9.1 Reproductive organs of the female 4.2.9.2 Reproductive organs of the male 4.2.10 Gametes 4.3 Reproduction and development 4.3.1 Reproductive biology 4.3.2 Cleavage 4.3.3 Development 4.4 Physiology 4.5 Phylogeny 4.6 Systematics 4.6.1 Classification 4.6.2 Keys 4.6.2.1 Key to higher taxa 4.6.2.2 Key to families of Bdelloidea 4.6.2.3 Key to families of Ploima 4.6.2.4 Key to families of Flosculariacea 4.6.2.5 Key to families of Collothecacea 4.6.3 Characterization of families 4.7 Biogeography 4.8 Paleontology 4.9 Ecology 4.9.1 Feeding ecology 4.9.2 Habitat 4.9.2.1 Freshwater and limnoterrestrial habitats 4.9.2.2 Saline environments 4.9.2.3 Symbiotic associations Literature 5 Acanthocephala: functional morphology 5.1 Introduction 5.2 General aspects 5.3 Body wall: host-parasite interface 5.4 Acanthocephalan muscles 5.5 Reproductive system and reproduction 5.6 Development: host-parasite interface in the intermediate host 5.7 Excretion and osmoregulation 5.8 Nervous and sensory receptor system 5.9 Phylogenetic considerations Literature 6 Acanthocephala 6.1 Introduction 6.2 Phylogeny 6.3 Systematics 6.3.1 Background to classification 6.3.2 Selection of diagnostic characters 6.3.3 Characterization of the classes of the Acanthocephala 6.3.4 Characterization of orders and families of the Acanthocephala 6.3.4.1 Archiacanthocephala 6.3.4.1.1 Apororhynchida: Apororhynchidae Shipley, 1899 (Fig.6.2) 6.3.4.1.2 Gigantorhynchida: Gigantorhynchidae Hamann, 1892 (Fig.6.3) 6.3.4.1.3 Moniliformida: Moniliformidae Van Cleave, 1924 (Figs.6.1 and 6.3) 6.3.4.1.4 Oligacanthorhynchida: Oligacanthorhynchidae Southwell & MacFie, 1925 (Fig.6.3) 6.3.4.2 Eoacanthocephala 6.3.4.2.1 Gyracanthocephalida: Quadrigyridae Van Cleave, 1920 (Fig.6.4) 6.3.4.2.2 Neoechinorhynchida: Dendronucleatidae Sokolovskaia, 1962 6.3.4.2.3 Neoechinorhynchida: Neoechinorhynchidae Ward, 1917 (Fig.6.4) 6.3.4.2.4 Neoechinorhynchida: Tenuisentidae Van Cleave, 1936 (Fig.6.5) 6.3.4.3 Palaeacanthocepha 6.3.4.3.1 Echinorhynchida 6.3.4.3.2 Echinorhynchida: Arhythmacanthidae Yamaguti, 1935 (Fig. 6.5) 6.3.4.3.3 Echinorhynchida: Cavisomidae Meyer,1932 (Fig.6.5) 6.3.4.3.4 Echinorhynchida: Diplosentidae Tubangi & Masilungan, 1937 (Fig. 6.6) 6.3.4.3.5 Echinorhynchida: Echinorhynchidae Cobbold, 1876 (Fig.6.5) 6.3.4.3.6 Echinorhynchida: Fessisentidae Van Cleave, 1931 (Fig.6.5) 6.3.4.3.7 Echinorhynchida: Heteracanthocephalidae Petrochenko, 1956 (Fig. 6.6) 6.3.4.3.8 Echinorhynchida: Illiosentidae Golvan, 1960 (Fig.6.6) 6.3.4.3.9 Echinorhynchida: Isthmosacanthidae Smales, 2012 (Fig.6.6) 6.3.4.3.10 Echinorhynchida: Pomphorhynchidae Yamaguti, 1939 (Fig.6.7) 6.3.4.3.11 Echinorhynchida: Rhadinorhynchidae Travassos, 1923 (Fig.6.7) 6.3.4.3.12 Echinorhynchida: Sauracanthorhynchidae Bursey, Goldberg & Kraus,2007 (Fig. 6.9) 6.3.4.3.13 Echinorhynchida: Transvenidae Pichelin & Cribb, 2001 (Fig.6.7) 6.3.4.3.14 Heteramorphida: Pyrirhynchidae Amin & Van Ha, 2008 (Fig.6.8) 6.3.4.3.15 Polymorphida: Centrorhynchidae Van Cleave, 1916 (Fig.6.8) 6.3.4.3.16 Polymorphida: Plagiorhynchidae Golvan, 1960 (Fig.6.8) 6.3.4.3.17 Polymorphida: Polymorphidae Meyer,1931 (Fig.6.8) 6.3.4.4 Polyacanthocephala 6.3.4.5 Uncertain assignment 6.5 Taxonomic keys 6.5.1 Key to the classes of the Acanthocephala 6.5.2 Key to the orders and families of the Archiacanthocephala 6.5.3 Key to the orders and families of the Eoacanthocephala 6.5.4 Key to the orders and families of the Palaeacanthocephala Literature 7 Ecology of the Acanthocephala 7.1 Introduction 7.2 Life cycle 7.3 Behavior 7.4 Population ecology 7.5 Community ecology 7.6 Environmental parasitology Literature Index

This section of the Handbook of Zoology is intended as a comprehensive and exhaustive account of the biology of the taxa Gastrotricha, Nematoda, Nematomorpha, Priapulida, Kinorhyncha, Loricifera, Gnathostomulida, Micrognathozoa, Rotifera, Seisonida and Acanthocephala, covering all relevant topics such as morphology, ecology, phylogeny and diversity. The series is intended to be a detailed and up-to-date account of these taxa. As was the case with the first edition, the Handbook is intended to serve as a reliable resource for decades.

Many of the taxa of this volume are comparatively unknown to many biologists, despite their diversity and importance for example in meiofaunal communities (Gastrotricha, Rotifera, Gnathostomulida), their fascinating recent discoveries (Loricifera and Micrognathozoa), their importance as parasites (many nematodes, Nematomorpha, Acanthocephala) and their importance for evolutionary questions (e.g. Priapulida, Gastrotricha). The groups covered range from those poor in species (such as Micrognathozoa with 2 known species) to the species-rich and diverse Nematoda and their ca. 20.000 described species. While each taxon is covered by one chapter, nematodes are treated in several chapters dedicated to their structural, taxonomic and ecological diversity.

Annotation This section of the Handbook of Zoology is intended as a comprehensive and exhaustive account of the biology of the taxa Gastrotricha, Nematoda, Nematomorpha, Priapulida, Kinorhyncha, Loricifera, Gnathostomulida, Micrognathozoa, Rotifera, Seisonida and Acanthocephala, covering all relevant topics such as morphology, ecology, phylogeny and diversity. The groups covered include the diverse Nematoda with ca. 20.000 described species. As with the first edition, the Handbook is intended to serve as a reliable resource for decades The Handbook of Zoology provides an in-depth treatment of the entire animal kingdom covering both invertebrates and vertebrates. It publishes comprehensive overviews on animal systematics and morphology and covers extensively further aspects like physiology, behavior, ecology and applied zoological research.