THE UNIVERSITY OF KANSAS NATURAL HISTORY MUSEUM SPECIAL PUBLICATION NO. 25 20 JULY 2017 Planetary Biodiversity Inventory (2008–2017): Tapeworms from Vertebrate Bowels of the Earth Edited by JANINE N. CAIRA Department of Ecology and Evolutionary Biology, University of Connecticut, Storrs, Connecticut, 06269-3043, USA AND KIRSTEN JENSEN Department of Ecology and Evolutionary Biology, Biodiversity Institute, University of Kansas, Lawrence, Kansas, 66045, USA NATURAL HISTORY MUSEUM THE UNIVERSITY OF KANSAS LAWRENCE, KANSAS SPECIAL PUBLICATION NATURAL HISTORY MUSEUM, THE UNIVERSITY OF KANSAS SERIES EDITOR: Kirsten Jensen TYPE SETTING AND LAYOUT: Kirsten Jensen ORIGINAL COVER ART: Kendel Craig COVER DESIGN: Kirsten Jensen & Janine N. Caira SPECIAL PUBLICATION NO. 25 pp. i–ix, 1–463 Published 20 July 2017 ISBN-10: 0-89338-002-4 ISBN-13: 978-0-89338-002-1 © THE UNIVERSITY OF KANSAS, NATURAL HISTORY MUSEUM Chapters in this volume were peer-reviewed. This Special Publication is electroniclly available at http://hdl.handle.net/1808/24421 and http://tapeworms.uconn.edu/finalpub.html PRINTED BY YURCHAK PRINTING, INC., LANDISVILLE, PENNSYLVANIA CONTENTS ACKNOWLEDGMENTS ............................................................................................................................viii PREFACE .................................................................................................................................................ix 1 AN OVERVIEW OF TAPEWORMS FROM VERTEBRATE BOWELS OF THE EARTH Janine N. Caira, K. Jensen, Boyko B. Georgiev, Roman Kuchta, D. Timothy J. Littlewood, Jean Mariaux, Tomáš Scholz, Vasyl V. Tkach, and Andrea Waeschenbach .............................1 2 AMPHILINIDEA Tomáš Scholz and Roman Kuchta ..........................................................................................21 3 BOTHRIOCEPHALIDEA Roman Kuchta and Tomáš Scholz ..........................................................................................29 4 CARYOPHYLLIDEA Tomáš Scholz and Mikuláš Oros ............................................................................................47 5 CATHETOCEPHALIDEA Janine N. Caira, Veronica M. Bueno, and Kirsten Jensen .....................................................65 6 CYCLOPHYLLIDEA Jean Mariaux, Vasyl V. Tkach, Gergana P. Vasileva, Andrea Waeschenbach, Ian Beveridge, Yana Dimitrova, Voitto Haukisalmi, Stephen E. Greiman, D. Timothy J. Littlewood, Arseny A. Makarikov, Anna J. Phillips, Tantely Razafiarisolo, Vincent Widmer, and Boyko B. Georgiev ...................................................................................................................77 7 DIPHYLLIDEA Janine N. Caira, Verónica A. Ivanov, Kirsten Jensen, and Fernando P. L. Marques ...........149 8 DIPHYLLOBOTHRIIDEA Roman Kuchta and Tomáš Scholz ........................................................................................167 9 GYROCOTYLIDEA Roman Kuchta, Tomáš Scholz, and Haakon Hansen ...........................................................191 10 HAPLOBOTHRIIDEA Roman Kuchta and Tomáš Scholz ........................................................................................201 11 LECANICEPHALIDEA Kirsten Jensen, Joanna J. Cielocha, Kaylee S. Herzog, and Janine N. Caira ........................207 12 LITOBOTHRIIDEA Janine N. Caira, Kaitlin Gallagher, and Kirsten Jensen .......................................................231 13 NIPPOTAENIIDEA Tomáš Scholz, Jan Brabec, and Roman Kuchta ....................................................................243 14 ONCHOPROTEOCEPHALIDEA I Alain de Chambrier, Tomáš Scholz, Jean Mariaux, and Roman Kuchta ..............................251 15 ONCHOPROTEOCEPHALIDEA II Janine N. Caira, Kirsten Jensen, and Verónica A. Ivanov ...................................................279 16 PHYLLOBOTHRIIDEA Timothy R. Ruhnke, Janine N. Caira, and Maria Pickering ................................................305 17 RHINEBOTHRIIDEA Timothy R. Ruhnke, Florian B. Reyda, and Fernando P. L. Marques..................................327 iii 18 SPATHEBOTHRIIDEA Roman Kuchta and Tomáš Scholz ........................................................................................349 19 TETRABOTHRIIDEA Jean Mariaux, Roman Kuchta, and Eric P. Hoberg .............................................................357 20 “TETRAPHYLLIDEA” VAN BENEDEN, 1850 RELICS Janine N. Caira, Kirsten Jensen, and Timothy R. Ruhnke ...................................................371 21 TRYPANORHYNCHA Ian Beveridge, Mohammed Haseli, Verónica Ivanov, Adriana Menoret, and Bjoern C. Schaeffner ...............................................................................................................401 22 A MOLECULAR FRAMEWORK FOR THE CESTODA Andrea Waeschenbach and D. Timothy J. Littlewood ..........................................................431 APPENDIX. PUBLICATIONS CITING THE NATIONAL SCIENCE FOUNDATION PLANETARY BIODIVERSITY INVENTORY PROGRAM GRANTS DEB 0818696 AND 0818823 .......................................................453 iv Contributors BEVERIDGE, IAN Department of Veterinary Science, University of Melbourne, 250 Princes Highway, Werribee, Victoria 3030, Australia. E-mail: [email protected] BRABEC, JAN Institute of Parasitology, Biology Centre of the Czech Academy of Sciences, Branišovská 31, 370 05 České Budějovice, Czech Republic. E-mail: [email protected] BUENO, VERONICA M. Department of Ecology and Evolutionary Biology, University of Connecticut, 75 N. Eagleville Rd., Storrs, CT 06269-3043, USA. E-mail: [email protected] CAIRA, JANINE N. Department of Ecology and Evolutionary Biology, University of Connecticut, 75 N. Eagleville Rd., Storrs, CT 06269-3043, USA. E-mail: [email protected] DE CHAMBRIER, ALAIN Muséum d’Histoire Naturelle de Genève, P.O. Box 6434, CH-1211 Geneva 6, Switzerland. E-mail: [email protected] CIELOCHA, JOANNA J. Biology Department, Science Center 220J, Rockhurst University, 1100 Rockhurst Road, Kansas City, MO 64110, USA. E-mail: [email protected] DIMITROVA, YANA D. Institute of Biodiversity and Ecosystem Research, Bulgarian Academy of Sciences, 2 Gagarin Street, 1113 Sofia, Bulgaria. E-mail: [email protected] GALLAGHER, KAITLIN Department of Ecology and Evolutionary Biology, University of Connecticut, 75 N. Eagleville Rd., Storrs, CT 06269-3043, USA. E-mail: [email protected] GEORGIEV, BOYKO B. Institute of Biodiversity and Ecosystem Research, Bulgarian Academy of Sciences, 2 Gagarin Street, 1113 Sofia, Bulgaria. E-mail: [email protected] GREIMAN, STEPHEN E. Department of Biology, Georgia Southern University, P.O. Box 8042-1, Statesboro, GA 30460, USA. E-mail: [email protected] HANSEN, HAAKON Norwegian Veterinary Institute, Section for Parasitology, Ullevålsveien 68, 0454 Oslo, Norway. E-mail: [email protected] HASELI, MOHAMMAD Department of Biology, Faculty of Sciences, University of Guilan, Rasht, Iran. E-mail: [email protected] HAUKISALMI, VOITTO Piettasenkatu 38 A 23, 33580 Tampere, Finland. E-mail: [email protected] HERZOG, KAYLEE S. Department of Ecology and Evolutionary Biology and the Biodiversity Institute, University of Kansas, 1200 Sunnyside Ave., Lawrence, KS 66045, USA. E-mail: [email protected] v HOBERG, ERIC P. Animal Parasitic Diseases Laboratory, Agricultural Research Service, US Department of Agriculture, Beltsville, MD, 20705-2350, USA and Department of Invertebrate Zoology, National Museum of Natural History, Smithsonian Institution, Washington, D.C., 20560-0163, USA. E-mail: [email protected] IVANOV, VERÓNICA A. Instituto de Biodiversidad y Biología Experimental y Aplicada - IBBEA (CONICET-UBA), and Departamento de Biodiversidad y Biología Experimental, Facultad de Ciencias Exactas y Naturales, Universidad de Buenos Aires, Ciudad Universitaria, Pabellón II, piso 4, Lab. 52, C1428EHA, Buenos Aires, Argentina. E-mail: [email protected] JENSEN, KIRSTEN Department of Ecology and Evolutionary Biology and the Biodiversity Institute, University of Kansas, 1200 Sunnyside Ave., Lawrence, KS 66045, USA. E-mail: [email protected] KUCHTA, ROMAN Institute of Parasitology, Biology Centre of the Czech Academy of Sciences, Branišovská 31, 370 05 České Budějovice, Czech Republic. E-mail: [email protected] LITTLEWOOD, D. TIMOTHY J. Department of Life Sciences, Natural History Museum, Cromwell Road, London, SW7 5BD, UK. E-mail: [email protected] MAKARIKOV, ARSENY A. Institute of Systematics and Ecology of Animals, Siberian Branch of Russian Academy of Sciences, 11 Frunze Str., 630091, Novosibirsk, Russia. E-mail: [email protected] MARIAUX, JEAN Muséum d’Histoire Naturelle de Genève, P.O. Box 6434, CH-1211 Geneva 6, Switzerland and Department of Genetics and Evolution, University of Geneva, CH-1205 Geneva, Switzerland. E-mail: [email protected] MARQUES, FERNANDO P. L. Departamento de Zoologia, Instituto de Biociências, Universidade de São Paulo, Rua do Matão, tv. 14, no. 101, 05508-090 Cidade Universitária, São Paulo, Brasil. E-mail: [email protected] MENORET, ADRIANA Instituto de Biodiversidad y Biología Experimental y Aplicada - IBBEA (CONICET-UBA), Facultad de Ciencias Exactas y Naturales, Universidad de Buenos Aires, Ciudad Universitaria, Pabellón II, piso 4, Lab. 52, C1428EHA, Buenos Aires, Argentina. E-mail: [email protected] OROS, MIKUláš Institute of Parasitology of the Slovak Academy of Sciences, Hlinkova 3, 040 01 Košice, Slovakia. E-mail: [email protected] PHILLIPS, ANNA J. Department of Invertebrate Zoology, National