GUIDE TO MONOGENOIDEA OF FRESHWATER FISH OF PALAEARTIC AND AMUR REGIONS O.N. PUGACHEV, P.I. GERASEV, A.V. GUSSEV, R. ERGENS, I. KHOTENOWSKY Scientific Editors P. GALLI O.N. PUGACHEV D. C. KRITSKY LEDIZIONI-LEDIPUBLISHING © Copyright 2009 Edizioni Ledizioni LediPublishing Via Alamanni 11 Milano http://www.ledipublishing.com e-mail: [email protected] First printed: January 2010 Cover by Ledizioni-Ledipublishing ISBN 978-88-95994-06-2 All rights reserved. No part of this publication may be reproduced, stored in a retrieval system, transmitted or utilized in any form or by any means, electonical, mechanical, photocopying or oth- erwise, without permission in writing from the publisher. Front cover: /Dactylogyrus extensus,/ three dimensional image by G. Strona and P. Galli. 3 Introduction; 6 Class Monogenoidea A.V. Gussev; 8 Subclass Polyonchoinea; 15 Order Dactylogyridea A.V. Gussev, P.I. Gerasev, O.N. Pugachev; 15 Suborder Dactylogyrinea: 13 Family Dactylogyridae; 17 Subfamily Dactylogyrinae; 13 Genus Dactylogyrus; 20 Genus Pellucidhaptor; 265 Genus Dogielius; 269 Genus Bivaginogyrus; 274 Genus Markewitschiana; 275 Genus Acolpenteron; 277 Genus Pseudacolpenteron; 280 Family Ancyrocephalidae; 280 Subfamily Ancyrocephalinae; 282 Genus Ancyrocephalus; 282 Subfamily Ancylodiscoidinae; 306 Genus Ancylodiscoides; 307 Genus Thaparocleidus; 308 Genus Pseudancylodiscoides; 331 Genus Bychowskyella; 332 Order Capsalidea A.V. Gussev; 338 Family Capsalidae; 338 Genus Nitzschia; 338 Order Tetraonchidea O.N. Pugachev; 340 Family Tetraonchidae; 341 Genus Tetraonchus; 341 Genus Salmonchus; 345 Family Bothitrematidae; 359 Genus Bothitrema; 359 Order Gyrodactylidea R. Ergens, O.N. Pugachev, P.I. Gerasev; 359 Family Gyrodactylidae; 361 Subfamily Gyrodactylinae; 361 Genus Gyrodactylus; 362 Genus Paragyrodactylus; 456 Genus Gyrodactyloides; 456 Genus Laminiscus; 457 Subclass Oligonchoinea A.V. Gussev; 457 Order Diclybothriidea; 460 Family Diclybothriidae; 460 Genus Diclybothrium; 460 Genus Paradiclybothrium; 461 Order Mazocraeidea; 465 Suborder Mazocraeinea; 466 Family Mazocraeidae; 466 Genus Mazocraes; 466 Suborder Discocotylinea I.A. Khotenovsky, O.N. Pugachev; 466 Family Diplozoidae: 466 Subfamily Diplozoinae; 470 Genus Paradiplozoon; 470 4 Genus Inustiatus; 506 Genus Eudiplozoon; 508 Genus Sindiplozoon; 508 Genus Diplozoon; 512 Family Discocotylidae; 514 Genus Discocotyle; 514 Family Octomacridae; 517 Genus Octomacrum; 517 Host–parasite list; 519 Literature; 537 Index List of parasites; 548 List of fishes; 559 5 GUIDE TO MONOGENOIDEA OF FRESHWATER FISH OF PALAEARTIC AND AMUR REGIONS INTRODUCTION The first key to parasites of freshwater fish of the USSR was published in Russian in 1962 and was translated into English in 1964. It was a one volume book that included keys for all parasite groups to the species level. Its translation is only one available internationally to present day. The second edition, also in Russian, was published between 1984 and 1987 and consists of three volumes (editor-in-chief O.N. Bauer). The second volume deals mostly with monogenenans (editor A.V. Gussev) and includes 24 keys and 16 supplements. In this volume, the Dactylogyri- dae (sensu Bychowsky et Nagibina, 1978) includes seven genera, and two separate keys can be used to identify over 200 nominal Dactylogyrus species. Three keys identify about 50 species in two subfamilies of the Ancyrocephalidae (sensu Bychowsky, 1937), and the Tetraonchidae in- cludes 20 species. Keys for four genera in the Gyrodactylidae identify about 150 nominal species. For the polyopisthcotyleans, several keys can be used to identify more than 30 species from 10 genera. Overall, about 500 species of monogeneans can be identified to species level using the keys and supplements in this book. Thus, this volume became popular among ichthyoparasitolo- gists, despite it being written in Russian. Currently, it is a rare book. More than 20 years have passed since the second edition was published, and many new monogenan species have been described. New data from different countries exists, and some re- visions to different monogenean taxa have been published. Therefore, preparation of a second edition English language “Guide to the Monogenoidea of Freshwater Fish of the Palaearctic and Amur Regions” seems to be of current importance. Because fish parasites, especially monogeneans, follow their host’s distribution, in gen- eral fish zoogeography reflects monogenean distribution. The terms “Palaearctic” and “Nearctic” are not zoogeographical units but can be used for notational convenience (Banarescu, 1992).Palaearctic defines the Eurasian part of the Holarctic that includes the Euro-Mediterranean, Circumpolar (the Asian part) (Berg, 1949), Baikal, and western Mongolian subregions. The Sa- hara marks a sharp southern border between northwestern Africa which belongs to Palaearctic and contains mainly European aquatic fauna, and tropical African and Ethiopian regions. In con- trast, vague borders exist in western Asia: Southern Anatolia, the entire Near East, central and southern Iran, and parts of Afghanistan must be considered as transitional areas between the Euro-Mediterranean subregion and the South Asian subregion of the Sino-Indian region (Ba- narescu, 1992). The same is true for the Amur River, which is transitional area between the Cir- cumpolar subregion and the Sino-Indian region. Considering that a number of fish species from the Amur River were introduced into European aquaculture, a separate key for Dactylogyrus spe- cies of Amur River fishes is included in this edition, in contrast to species from other transitional areas which are inserted to a common keys. Some precise data on monogenean distribution can be found in Pugachev’s (2002) “Checklist of the Freshwater Fish Parasites of the Northern Asia” and Moravec’s (2001) “Checklist of the Metazoan Parasites of Fishes of the Czech Republic and Slo- vak Republic (1873–2000).” When adding newly described species and species from new geographic regions to the keys from the 1985 publication, it seemed prudent to retain the structure of the keys. This pro- vides continuity between the previous and current editions of the book that would have been lacking if the keys were completely revised. Moreover there is no modern accepted monoge- nean classification. Although the more conservative monogenean classification was used in this edition, it does not greatly influence the identification process. Therefore, some monogenean groups maintain their systematic status (e.g., order Tetraonchidea, family Ancyrocephalidae, etc.). However, some groups obviously need revision (e.g., genera Dactylogyrus and Gyrodac- tylus). Many changes were made in this edition of the book. For simplicity, the small theoretical chapters (e.g., infection pathways of monogeneans and marine monogenean species) from the second edition were excluded from this one. A new family—Bothitrematidae, with the genus Bothitrema—was added to the common key of families. The key for Dactylogyrus now includes 62 additional species, and the key for Gyrodactylus now includes 25 new species. The key for the 6 GUIDE TO MONOGENOIDEA OF FRESHWATER FISH OF PALAEARTIC AND AMUR REGIONS Tetraonchidae was completely revised, and the key for the Diplozoidae was reconstructed from Khotenovsky (1985b). Altogether 97 species were added to the different keys, yielding a total of 622 valid species with more than 200 synonyms and 714 figures for the Palaearctic monogeneans and species which are not found yet in the Palaearctic but theirs discovery in the future is ex- tremely probable. The principle behind dichotomic diagnostic keys consists of comparing characters pre- sented as numbered items in the form of “thesis – antithesis.” Before beginning the process of identifying a given monogenean, detailed data must be collected, including the form and size of the body, site, or organ (e.g., haptoral armament, copulatory organ, vaginal armament, etc.). The subclass is the first taxon to be determined, followed by the order and family to which a given specimen belongs. The identification process proceeds as follows. Identification begins with the first key item. Both the thesis and antithesis must be read carefully. The antithesis number is in parenthe- ses. If the parasite possesses all of the features included in item 1 and there is no Latin name at the end of that item, then proceed to item 2 and again compare the thesis and antithesis. If the fea- tures of a given thesis do not match the specimen characteristics, proceed to the antithesis and then to the next item. This process continues until the Latin name is reached at the end of a suit- able item that corresponds to the characters of the specimen in question. The keys to species con- tain a brief description with a reference to the appropriate figures, hosts, and geographical distri- bution. Geographical distribution is characterized mainly using either host range or river basins or sea basins. In some cases, state names were used, taking into account the latest changes in Europe and the former Soviet Union. Latin names of fish species are in accordance with the “Catalogue of Fish Names” (Eschmeyer, 1998; www.calacademy.org) or Fishbase (www.fishbase.org) All keys that are part of the Russian edition were translated by O.N. Bauer and O.N. Pugachev. All indexes, host- parasite lists, and literature were prepared by Mrs. Irine Pugacheva. The authors would like to thank Dr. Nina Bogutskaya for consultations about fish classification (especially the