Ukázka knihy z internetového knihkupectví www.kosmas.cz

AcAdemy of ScienceS of the czech Republic

DracunculoiD anD anGuillicoloiD nematoDes Parasitic in vertebrates

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the publisher thanks the for a financial contribution that madethe publication of this book possible.

U k á z k a k n i h y z i n t e r n e t o v é h o k n i h k u p e c t v í w w w . k o s m a s . c z , U I D : K O S 1 8 2 5 3 3

U k á z k a k n i h y z i n t e r n e t o v é h o k n i h k u p e c t v í w w w . k o s m a s . c z , U I D : K O S 1 8 2 5 3 3

U k á z k a k n i h y z i n t e r n e t o v é h o k n i h k u p e c t v í w w w . k o s m a s . c z , U I D : K O S 1 8 2 5 3 3

contents

preface . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13Historical review . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17methods of study of dracunculoid and anguillicoloid nematodes . . . . . . . . . . . . . . . . . . 37morphology and taxonomical features of dracunculoids and anguillicoloids . . . . . . . . 48biology of dracunculoid and anguillicoloid nematodes . . . . . . . . . . . . . . . . . . . . . . . . . . 63Pathogenicity and significance of dracunculoids and anguillicoloids . . . . . . . . . . . . . . 70list of dracunculoid and anguillicoloid nematodes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 77systematic review of dracunculoid and anguillicoloid nematodes . . . . . . . . . . . . . . . . . 84Superfamily dracunculoidea . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 84family daniconematidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 86

Genus Daniconema . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 87Genus Mexiconema . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 91Genus Syngnathinema . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 97

family dracunculidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 99Genus Avioserpens . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 100Genus Dracunculus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 113

family Guyanemidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 152Subfamily Guyaneminae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 152

Genus Guyanema . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 153Genus Histodytes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 165Genus Moravecia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 167Genus Pseudodelphis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 169

Subfamily travassosneminae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 174Genus Ichthyofilaria . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 175Genus Travassosnema . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 183

family lucionematidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 187Genus Lucionema . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 187

family micropleuridae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 190Subfamily micropleurinae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 191

Genus Granulinema . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 191Genus Kamegainema . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 196Genus Micropleura . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 198Genus Philonema . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 212Genus Protenema . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 226

Subfamily phlyctainophorinae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 229Genus Phlyctainophora . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 230

family philometridae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 234

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Subfamily Alineminae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 235Genus Alinema . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 235

Subfamily neophilometroidinae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 240Genus Neophilometroides . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 240

Subfamily philometrinae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 247Genus Buckleyella . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 248Genus Clavinema . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 250Genus Dentiphilometra . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 260Genus Nilonema . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 264Genus Paraphilometroides . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 267Genus Philometra . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 271Genus Philometroides . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 420Genus Rumai . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 474Genus Spirophilometra . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 476

family Skrjabillanidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 477Subfamily esocineminae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 478

Genus Esocinema . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 478Subfamily Skrjabillaninae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 481

Genus Molnaria . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 481Genus Sinoichthyonema . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 487Genus Skrjabillanus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 490

family tetanonematidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 502Genus Tetanonema . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 502

Genus incertae sedis: Lockenloia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 504Superfamily Anguillicoloidea . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 506family Anguillicolidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 506

Genus Anguillicola . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 506Genus Anguillicoloides . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 511

Host – parasite list . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 541pisces (fishes) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 541Amphibia (Amphibians) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 575Reptilia (Reptiles) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 576Aves (birds) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 579mammalia (mammals) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 581

references . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 585index . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 618

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Ukázka knihy z internetového knihkupectví www.kosmas.cz

Preface

the idea of writing the monograph presented here arose from the fact that,until now, no book dealing specifically with the nematodes of the spiruridsuperfamily dracunculoidea has been published. the only publication providinga survey of all dracunculoids, anguillicoloids and camallanoids known at thattime was the book “camallanata of animals and man and the diseases caused bythem” by ivashkin et al. (1971) published in Russian. now, it is rather out ofdate and the classification system used in it is not accepted by mosthelminthologists. moreover, many new interesting forms of dracunculoids andanguillicoloids were discovered since, mainly from fishes, which substantiallyextend the present knowledge about members of these two nematode groups.important new information has also accummulated about the biology,distribution and host–parasite relationships of some of them.

despite the fact that dracunculoids represent a large group of importantparasites with hosts belonging to all classes of vertebrates including man,their recognition is by far inadequate. Recent morphological, biological andmolecular studies of members of the family Anguillicolidae, which hasusually been assigned to the dracunculoidea, now show that this smallnematode group is phylogenetically distant from other dracunculoids andshould be placed in an independent superfamily Anguillicoloidea. Speciesdescriptions of dracunculoids and anguillicoloids, frequently incomplete andoften misleading, are scattered in many journals and, to date, there is nomonograph about these nematodes enabling their species identification.therefore, i considered it useful to compile the book dealing with theseparasites as a whole, in a similar way as my previous book treating thetrichinelloids parasitic in cold-blooded vertebrates (moravec 2001a).

my first experience with dracunculoids dates back to the beginning of thesixties of the last century, when as a university student working on theparasites of reptiles in former czechoslovakia, i discovered in grass-snakesin a South-Slovakian locality a strange, at that time almost unknownnematode Dracunculus oesophageus, previously reported only in italy. later,at the institute of parasitology, Academy of Sciences of the czech Republic(previously the czechoslovak Academy of Sciences), originally in prague and

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from 1985 in České budějovice, i had a possibility to study a great number ofdracunculoid species: mainly parasites of fishes, rarely those parasitizingreptiles and mammals and from different countries of the world. thesematerials were collected by me and my fellow-workers. Some were providedby foreign colleagues. Specimens of some species were borrowed for a re-examination from different foreign museums and other scientific institutions.Accordingly, the species descriptions in this monograph are often based onthe author’s studies while others were adapted from the respective literature.in addition to descriptions and illustrations, i included for each dracunculoidand anguillicoloid species the data on its hosts, site of localization,distribution and in particular their life cycles and bionomy, provided thatsuch data are available. my work in previous years on the life cycles ofseveral species of Philometra and Anguillicoloides that parasitize europeanfishes contributed to the knowledge of these parasites in this respect.

A large amount of literary data on dracunculoids and anguillicoloids hasaccummulated, particularly those dealing with the famous human pathogenDracunculus medinensis and, more recently, Anguillicoloides crassus,a highly pathogenic east-Asian parasite of eels (Anguilla spp.) introducedinto europe and some other continents about twenty five years ago. there arealso many reports on Philometra sp. in the extensive literature about fishparasites. therefore, i made no attempt to obtain every record of dracunculoids,which is nearly impossible. thus, there may be published host and localityrecords that are not included here. nevertheless, i believe that my lists arereasonably complete. however, when using this book for the identification ofdracunculoids, it is necessary to take into account that it only summarizes thepresent state of knowledge about this group of nematodes, which as a wholeremains insufficiently recognized. many species are still waiting for theirdiscovery.

i am grateful to prof. Vlastimil baruš from the institute of Vertebratebiology, AScR, brno for reading this book in manuscript and making criticalcomments and useful suggestions as to content and presentation. drs. davidmodrý from the faculty of Veterinary medicine in brno, Jiljí Sitko from theRegional museum in přerov and miloš Anděra from the national museum inprague helped with scientific names of host reptiles, birds and mammals,respectively. thanks are offered to irena husáková, institute of parasitology,AScR, České budějovice for technical assistance during the work. Specialthanks are due to dr. hassan hashmi, university of massachusetts, forrevising the english.

i would like to express my thanks to all who, during years of studies ondracunculoids, provided me with the type and other specimens, thus makingthe present work possible: david i. Gibson, the natural history museum,london; uK, Jean-lou Justine, museum national d’histoire naturelle, paris,

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france; Ahmed tayeh, World health organization, Geneva, Switzerland;marianne Køie, marine biological laboratory, copenhagen university,helsingør, denmark; leo margolis, pacific biological Station, fisheries andoceans, nanaimo, british columbia, canada; Anna Kohn, instituto oswaldocruz, Rio de Janeiro, brazil; Kálmán molnár and czaba Székely, Veterinarymedical Research institute, hungarian Academy of Sciences, budapest,hungary; lia paggi, istituto di parassitologia, università di Roma “laSapienza”, italy; david di cave, dipartimento Sanitá e biologia cellulare,università di Roma “tor Vergata”, Rome, italy; Kazuya nagasawa, instituteof far Seas fisheries, fisheries Agency of Japan, Shimizu, Japan; Kazuoogawa, department of Aquatic bioscience, Graduate School of Agriculturaland life Sciences, university of tokyo, tokyo, Japan; pin nie, Guitang Wang,Shangong Wu and huisheng Wu, laboratory of fish diseases, institute ofhydrobiology, chinese Academy of Sciences, Wuhan, china; h. hirose,nihon university, tokyo, Japan; Guillermo Salgado-maldonado, instituto debiología, universidad nacional Autónoma de méxico, mexico city, mexico;Victor Vidal-martínez and leopoldina Aguirre-macedo, centro de investigaciony de estudios Avanzados del ipn, unidad mérida, mérida, mexico; horsttaraschewski, institute of zoology, university of Karlsruhe, Karlsruhe,Germany; e. Karlsbakk, institute of fisheries and marine biology, universityof bergen, norway; maurice d. little and barry G. campbell, tulane medicalcenter, tulane university, new orleans, uSA; b. Jones, fisheries Researchcentre, ministry of Agriculture and fisheries, Wellington, new zealand;Shigehiko urawa, hokkaido Salmon hatchery, fisheries Agency of Japan,Sapporo, Japan; d. c. lee, South Australian museum, Adelaide, Australia;Shunya Kamegai, meguro parasitological museum, tokyo, Japan; ilan paperna,department of Animal Sciences, hebrew university of Jerusalem, Jerusalem,israel; puqin Wang, parasitology Research laboratory, fujian normaluniversity, china; Jo G. Van As, department of zoology and entomology,university of the free State, South Africa; Gill V. Spangenberg, underwaterWorld, San francisco, california, uSA; david G. huffman, freeman AquaticStation, Southwest texas State university, San marcos, texas, uSA; Russel p.hobbs, division of Veterinary and biomedical Sciences, murdoch university,murdoch, Australia; Winston R. Kay, department of zoology and entomology,School of life Sciences, the university of Queensland, brisbane, Australia;Juan m. timi, departamento de biología, universidad nacional de mar delplata, Argentina, ercument Genc, department of Aquaculture and fisheries,mustafa Kemal university, hatay, turkey; Atheer h. Ali, department offisheries and marine Resources, college of Agriculture, basrah university,basrah, iraq; clive R. Kennedy, department of zoology, university ofexeter, exeter, uK; G. Arbocco, museo civico di Storia naturale, Genoa,italy; G. V. Vasilkov, ViGiS, moscow, Russia; dalisey l. Ribu, department of

Preface 9

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microbiology and parasitology, the university of Queensland, brisbane,Australia; Ahmed hafiz, marine Research Section, ministry of fisheries,Republic of maldives; Robert Konečný, umweltbundesamt / environmentAgency, Vienna, Austria; céline levron, laboratoire parasites et ecosystèmesméditerranées, université de corse, corte, france; thorsten Walter, marinepathology, Group, institut für meereskunde Kiel, Kiel, Germany; isaure duburon, college of biology, university of charlestown, South carolina, uSA;patricia pilitt and eric p. hoberg, uS national parasite collection, beltsville,maryland, uSA; predrag cakić, institute of biological Research, belgrade,yugoslavia; Vladimíra hanzelová and marta Špakulová, parasitologicalinstitute, Slovak Academy of Sciences, Košice, Slovak Republic; iva dyková,Radim ergens, ivan fiala, Roman Kuchta, Jiří lom and tomáš Scholz,institute of parasitology, Academy of Sciences of the czech Republic, Českébudějovice, czech Republic; milan Gelnar, Andrea Šimková and ludmilaVojtková, department of zoology and ecology, faculty of Sciences, masarykuniversity, brno, czech Republic; Vlastimil baruš, institute of Vertebratebiology, Academy of Sciences of the czech Republic, brno, czech Republic;and Antonín prouza and Stanislav Červinka, State Veterinary institute, Českébudějovice, czech Republic.

for help in securing specimens and for providing me with necessaryliterature, i am also indebted to: Roy c. Anderson, department of zoology,university of Guelph, Guelph, ontario, canada; p. Ghittino, istituto zoopro -filactico Sperimentale del piemonte e della liguria, torino, italy; A. m.parukhin, institute of biology of Southern Seas, Academy of Sciences of theukr. SSR, Sevastopol, ukraine (then ukr.SSSR); Robin m. overstreet, Gulfcoast Research laboratory, the university of Southern mississippi, oceanSprings, mississippi, uSA; thomas l. crosby, Virginia State university,petersburg, Virginia, uSA; marcelo e. oliva, universidad de Antofegasta,Antofegasta, chile; Valentina A. tikhomirova, tver, Russia; Sergey G. Sokolov,institute of ecology and evolution, Russian Academy of Sciences, moscow,Russia; Annie J. petter, museum national d’histoire naturelle, paris, france;and Sandy cairncross and Ralph muller, department of infectious and tropicaldiseases, london School of hygiene and tropical medicine, london, uK.

many of the illustrations in the book have been redrawn from publishedworks. for this, i am grateful to the authors whose names, with date ofpublication, appear in the captions to the relevant figures. the full title andpublisher of each source is cited in the reference list. i thank the followingpublishers for their kind permission to use illustrations in their publications:Academia (figs. 17, 122, 124, 143, 155A–e, 175A–e, G–J, 192, 199, 203, 205),Acta Hydrobiologica Sinica (fig. 86 c–e), Acta Parasitologica (figs. 110, 117,124, 126, 130, 138, 139, 159, 163, 168, 170, 174 A–h), Acta Societatis ZoologicaeBohemicae (former Acta Societatis Zoologicae Bohemoslovacae) (figs. 50, 121,

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127), Acta Veterinaria Hungarica (figs. 118, 155 f), Acta Zoologica Mexicana(fig. 165 A–d), Acta Zoologica Sinica (fig. 209), Acta Zootaxonomica Sinica(fig. 132 A–c), American Journal of Hygiene (fig. 32 b–l), American Journal ofTropical Medicine (fig. 28 f–h), Angewandte Parasitologie (fig. 197), Annalsand Magazine of Natural History (fig. 62 c), Parasite (former Annals deParasitologie Humaine et Comparée) (figs. 16, 23 e, h–z, 33 l–n), Arquivos doInstituto Biologigo, S. Paulo (figs. 95, 182), Atas da Sociedade de Biologia doRio de Janeiro (fig. 90 b–d, f, 196), Bulletin du Muséum National d’HistoireNaturelle [© publications Scientifiques du muséum national d’histoirenaturelle, paris] (figs. 39, 71, 72, 119, 128 c–m), Bulletin of the Nansei RegionalFishery Research Laboratory (fig. 85), Bulletin de la Société Zoologique deFrance (fig. 22 d–f, i, J), Canadian Journal of Fisheries and Aquatic Sciences(fig. 131), Canadian Journal of Zoology (figs. 28 e, i, 29, 48, 65, 67, 74 A–c, f, G,i–l, 180 A, bd–f, h–m,), Chinese Journal of Zoology (fig. 82 d–f), J. & A.churchill, london (fig. 113 c), Ciencias Marinas (fig. 46 A, b), ComparativeParasitology (former Proceedings of the Helminthological Society of Washington)(figs. 28 A–d, 74 d, e, h, 105 c, d, G, h), Comptes Rendus des Séances (etMémoires) de la Société de Biologie et de Ses Filiales, Paris (fig. 60 A–c),Contributions to Canadian Biology and Fisheries (fig. 65 A, b, d, e, f, h),Diseases of Aquatic Organisms [© inter-Research] (fig. 223), ExperimentalPathology and Parasitology (fig. 174 i, J), Folia Parasitologica (figs. 3–6, 10–13,21, 25, 26, 35, 37, 38, 41, 44, 46 c–J, 52–55, 63, 77, 78, 84, 91, 92, 96 b, 106–109, 115,125, 133–137, 140, 142, 143–147, 152, 156, 164, 165 e–G, 169, 171, 183, 190, 198,208, 210–222), Helminthologia (fig. 68), international Academic publishers(fig. 20), Japanese Journal of Zoology (figs. 49, 64 b, d, 86 A, b, 123, 141, 194 c,d), Japanese Journal of Parasitology (fig. 193 d–f), Journal of Helminthology(figs. 81, 89, 93, 94, 104 A, b, 105 A, b, e, f, 120, 153, 166, 167 d, e, 172 d–f, 177,189), Journal of the Medical College in Keijo (fig. 82 A–c), Journal ofParasitology (figs. 14, 22 K, 30 A–f, l, 34 A, b, e, f, 45, 56–58, 75, 76, 79, 87, 88,102, 148, 186 c, d, f, G, h, 207), Mémoires de l’Institut Scientifique duMadagascar (fig. 23 A–d, f, G), Memórias do Instituto Oswaldo Cruz (figs. 24,51), nauka, moscow (figs. 18, 19, 111 e, 167 A–c, 175 f, 176, 206), naukovadumka, Kiev (figs. 116, 128 A, b, 129 e, f, 191 e, f), ocean press, beijing(fig. 179), Ophelia (fig. 47 c–f), Parasitology (figs. 64 A, e, f, 90 A, e),Parasitology Research (former Zeitschrift für Parasitologie) [© Springer](figs. 27 A–d, 34 c, d, G, 42, 59, 157), Parazitologiya (figs. 70, 178, 204),Proceedings of the Academy of Natural Sciences of Philadelphia (fig. 167 f),Proceedings of the Latvian Academy of Sciences (figs. 185, 195), PublicacionesBiologicas Instituto de Investigaciones Cientificas, Monterrey (fig. 99), ResearchBulletin of the Panjab University (fig. 132 d–f), Revista de Medicina Veterinaria,Buenos Aires (fig. 60 d), Revue Suisse de Zoologie (figs. 36, 40, 80), Revue deZoologie Africaine (fig. 96 A, c–e), Rivista di Parassitologia (fig. 129 A–d),

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Science press, beijing (figs. 103, 112), Systematic Parasitology [© Springer](figs. 43, 66, 83, 114, 160, 161, 194 A, b, e, f, 202), taylor and francis, london(fig. 62 A, d), Transactions of the American Microscopical Society (fig. 15),Transactions of Researches on Fish Diseases (China) (fig. 191 A–d),Transactions of the Royal Society of South Australia (fig. 151), TrudyGelmintologicheskoy Laboratorii (GELAN) (figs. 185 c, h, 201), TrudyLeningradskogo Obshchestva Estestvoispytateley (fig. 69 b, c, e), TrudyZoologicheskogo Instituta AN USSR (fig. 97 c, d), Veterinary Medical Journal(Egypt) (fig. 182), Zentralblatt für Bakteriologie (fig. 193 A–c), Zentralblatt fürParasitenkunde, Infektionskrankheiten und Hygiene (fig. 73), ZoologicheskiyZhurnal (figs. 69 A, d, 101, 173, 187), Zeitschrift für wissenschaftliche Zoologie(fig. 111 A, b, d, G, h), Zoologia Polonica (fig. 162), Zoologische Jahrbücher(fig. 150), Zoologischer Anzeiger (figs. 47 A, b, 99, 158).

this work was accomplished within the author’s grant project no. 524/03/0061from the Grant Agency of the czech Republic. i wish to thank the editorialboard of the Academy of Sciences of the czech Republic for the financialsupport that made the publication of this book possible. i am most gratefulfor the excellent editorial and production work of mr. pavel zátka from thepublishing house Academia who helped in many ways, especially in workingout the format of the book.

12 Preface

Ukázka knihy z internetového knihkupectví www.kosmas.cz

introduction

nematodes of the spirurid superfamilies dracunculoidea Stiles, 1907 andAnguillicoloidea yamaguti, 1935, which are characterized by certainmorphological features and some biological peculiarities, represent a largeand diverse group of parasites with worldwide distribution. members arenoted for the presence of a simple oval or triangular mouth surrounded bya variable number of cephalic papillae usually arranged in two circles. thebuccal capsule, if present, is usually reduced. the oesophagus is largelydivided into muscular and glandular parts but may be entirely muscular. twospicules or a sclerotized copulatory plate may be present or absent. Alldracunculoids are viviparous, whereas anguillicoloids are oviparous. Afterfertilisation, dracunculoid females grow markedly as first-stage larvae filltheir uteri. in some groups, the vulva and anus atrophy in fully gravid femalesand larvae are dispersed into the environment by bursting as they come incontact with water, whereas in others their larvae are released into tissues.many dracunculoids exhibit marked sexual dimorphism, in which females arehighly modified and considerably larger than males. the body size of gravidfemales in different species ranges between 1 mm in Lucionema balatonensefrom european pikeperch to more than 1 m in Philometra sp. from wreckfish(moravec 2004b) or Avioserpens mosgovoyi from aquatic birds.

According to moravec (2004b), the superfamily dracunculoidea includesthe following nine families: dracunculidae Stiles, 1907, micropleuridae bayliset daubney, 1926, philometridae baylis et daubney, 1926, Anguillicolidaeyamaguti, 1935, tetanonematidae Skryabin et Shikhobalova, 1948, Skrja -billanidae Shigin et Shigina, 1958, Guyanemidae petter, 1974, daniconema -tidae moravec et Køie, 1987, and lucionematidae moravec, molnár etSzékely, 1998. the monotypic dracunculoid genus Lockenloia Adamson etcaira, 1991 with L. sanguinis Adamson et caira, 1991 from sharks has notbeen assigned to any family and is considered incertae sedis. however, in thisbook, the family Anguillicolidae is removed from dracunculoidea and isplaced in a separate superfamily, Anguillicoloidea (see preface).

the hosts of the numerous species of these two nematode superfamilies are members of all main classes of vertebrates (fishes, amphibians, reptiles,

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birds, mammals), including man. however, their highest species and morpho -logical diversity occurs in fishes. Some species parasitize a wide range of hostswhile others exhibit a relatively narrow host specificity. the dracunculoidsare mostly parasites of various tissues and organ cavities and different speciesattack, for example, the skin and subcutaneous tissue, body musculature, eyes,orbits, gills and the swimbladder (in fish), kidneys, gonads and the circulatorysystem, or are found in the body cavity of the host (ivashkin et al. 1971;chabaud 1975; moravec 2004b).

the dracunculoid and anguillicoloid life cycles involve aquatic crustaceans(copepods, ostracods or branchiurids) as intermediate hosts and, consequently,the transmission and the occurrence of these parasites in vertebrate definitive hosts is always associated with the aquatic environment. therefore,it is not surprising that the absolute majority of dracunculoids and allanguillicoloids are found in fishes, both freshwater and marine. of the ninedracunculoid and anguillicoloid families, eight (89%) include the forms fromfishes, whereas the family dracunculidae contains a few species parasitic tobirds (species of Avioserpens) and reptiles and mammals (members ofDracunculus). the micropleuridae contains four species of Micropleura fromreptiles, two species of the monotypic genera Kamegainema and Protenemafrom amphibians, two species of each Granulinema and Phlyctainophorafrom elasmobranchs (sharks) and four species of Philonema from teleosteanfishes. of a total number of 37 valid genera of dracunculoid and anguillicoloidnematodes with 165 recognized species, 32 genera (86%) with 145 species(88%) are found in fishes.

dracunculoid nematodes are widely distributed among vertebrates,particularly in freshwater, brackish-water and marine fishes, whereas anguilli -colloids adapted only to freshwater eels (Anguilla spp.). About 450 fishspecies, belonging to 131 fish families and 31 orders, have been reported asthe hosts of these parasites. besides the well known human parasiteDracunculus medinensis, many other dracunculoids are known as pathogensin animals such as caudate amphibians, turtles, snakes, crocodiles, birds andsome mammals. Some fish dracunculoids and anguillicoloids are highlypathogenic, which are known to be agents of serious diseases of fish witheconomic importance, causing sometimes mass mortalities in stocks of bothfree-living and cultured fish (e.g., anguillicolosis of eels, philometroidosis ofcarps, philonemosis of salmonids). the frequent destruction of gonads causedby Philometra spp. in cultured marine perciform fishes may considerablydecrease the reproduction of these fishes and may even cause full parasiticcastration (Ramachandran 1975; Sakaguchi et al. 1987; moravec, ogawa et al.2002; moravec, Glamuzina et al. 2003). fish philometrids (Philometra sp.and Philometroides sp.) were also recorded as facultative human parasites(deardorff et al. 1986; Kuroda et al. 1991).

14 introduction