Aquatic Oligochaete Biology Developments in Hydrobiology 51

Series editor H. J. Dumont Aquatic Oligochaete Biology

Proceedings of the 4th International Symposium on Aquatic Oligochaete Biology

Edited by J. L. Kaster

Reprinted from Hydrobiologia, vol. 180 (1989)

Kluwer Academic Publishers

Dordrecht / Boston / London Library of Congress Cataloging-in-Publication Data

~quatic 01 igochaete bIology! edited by Jerry L. Kaster. p. cm. -- (Developments In hydroblology ; 51) "Proceedings of the 4th InternaTional Sycposium on Aquatic 01 igochaete Biology ... March 13-18 [1988] at the State University. Baton Rouge"--Pref.

1. Ollgochaeta--Congresses. 2. Aquat ic invertebrates--Congresses. I. Kaster. Jerry L. II. International SymposIum on AquatIc 01 igochaete Biology (4th 1988 LouIsiana State UniversIty. Baton Rouge. La.) III. SerIes. QL391.A6A68 1989 595.1 '46--dc20 89-11141

ISBN-13: 978-94-010-7577-0 e-ISBN-13:978-94-009-2393-5 DOl: 10.1007/978-94-009-2393-5 Kluwer Academic Publishers incorporates the publishing programmes of Dr W. Junk Publishers, MTP Press, Martinus Nijhoff Publishers, and D. Reidel Publishing Company.

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Copyright

© 1989 by Kluwer Academic Publishers, Dordrecht Softcover reprint of the hardcover 1st edition 1989 All rights reserved. No part of this publication may be reproduced, stored in a retrieval system, or transmitted in any form or by any means, mechanical, photocopying, recording, or otherwise, without the prior written permission from the copyright owners. v

Contents

Preface by J. L. Kaster VII In memoriam: Dr. Oarence James Goodnight IX

Part One: Taxonomy and Evolution 1. Comments on the classification of the Clitellata by P. C. Holt 1 2. Did the lumbriculids provide the ancestors of the branchiobdellidans, acanthobdellidans and leeches? by R. O. Brinkhurst & S. R. Gelder 7 3. Phylogeny and origins of Enchytraeidae by K. A. Coates 17 4. Preliminary investigations of hybridization/reticulate evolution in Guaranidrilus (Enchytraeidae: Oligochaeta) by K. A. Coates 35 5. Preliminary report on the Enchytraeidae (Oligochaeta) of West by B. Healy 47 6. New species of Phallodrilus (Oligochaeta, Tubificidae) from caves of northern Spain and southwestern France by P. Rodriguez & N. Giani 57

Part Two: Anatomy, Ultrastructure and Physiology 7. Neurobehavioral specializations for respiratory movements and rapid escape from predators in posterior segments of the tubificid Branchiura sowerbyi by C. D. Drewes & M. J. Zoran 65 8. Different models of tubificid spermatozeugmata by M. Ferraguti, G. Grassi & C. Erseus 73 9. Body wall organization in enchytraeids by R. Valvassori, M. de Eguileor, G. Lanzavecchia & G. Searl 83 10. Morphogenesis of body wall muscle fibers in Enchytraeus minutus by G. Lanzavecchia, M. de Eguileor, R. Valvassori, L. Di Lernia & C. Cambiaso 91 11. Histochemical analysis of oligochaete body wall by M. de Eguileor, S. Daniel, F. Cotelli, R. Valvassori & G. Lanzavecchia 99 12. T -system in muscles of microdriles by M. de Eguileor, R. Valvassori, G. Lanzavecchia & L. Di Lernia 109 13. Histophysiology of digestion and observations on the structure of the alimentary canal in the ectosymbiont Chaetogaster limnaei limnaei Baer, 1827 (Annelida: Oligochaeta) by S. R. Gelder 115 VI

Part Three: Life Histories and Population Dynamics 14. Population analysis in mass cultures of Tubifex tubifex by C. Bonacina, G. Bonomi & C. Monti 127 15. Density-dependent processes in cohorts of Tubifex tubifex, with special emphasis on the control of fecundity by C. Bonacina, G. Bonomi & c. Monti 135 16. Cohort cultures of Tubifex tubifex forms by A. Paoletti 143 17. Biological studies upon Enchytraeus variatus Bouguenec & Giani 1987 in breeding cultures by V. Bouguenec & N. Giani 151 18. Population dynamics of Tubificoides amplivasatus (Oligo chaeta, Tubificidae) in the Oresund, Denmark by C. Erseus & R. J. Diaz 167 19. Intrinsic and extrinsic variables controlling the productivity of asexual populations of Nais spp, (Naididae, Oligo chaeta) by J. Juget, V. Goubier & D. Barthelemy 177

Part Four: Commmunity Studies and Pollution Biology 20. Meiobenthic naidid and aelosomatid oligochaetes from the pro fundal zone, and relations of species to eutrophication by J. Sarkka 185 21. Observations of predator-prey interaction on dispersal of an oligochaete prey, Limnodrilus hoffmeisteri by J. L. Kaster 191 22. Pollution and tidal benthic communities of the James River Estuary, by R. J. Diaz 195 23. The role of oligochaetes in the management of waters by P. F. M. Verdonschot 213 24. Colonization of a manmade gravel bar by Oligochaeta by C. R. Bingham & A. C. Miller 229 25. Aquatic toxicity test for enchytraeids by J. Roembke & Th. Knacker 235 26. Acute and chronic effects of an anionic surfactant on some freshwater tubificid species by S. Casellato & P. Negrisolo 243 Hydrobiologia 180: VII, 1989. J. L. Kaster (ed.) Aquatic Oligochaete Biology. VII

Preface

This volume reflects the proceedings of the 4th International Symposium on Aquatic Oligochaete Biology, unique in that it represents the first such meeting to be held in the United States. This meeting was hosted by Dr. Walter J. Harman during March 13-18 at the Louisiana State University, Baton Rouge, Louisiana, U.S.A. For the many that are acquainted with Dr. Harman, you might have guessed that the meeting would be a distinguished success and it was, with all of the grandeur that only a southern setting could provide. This success was measured not only by the enthusiastic scientific exchange that centered on the theme of our favorite subject, aquatic oligochaetes, but also by the encouraging atmosphere provided by Dr. Harman and his staff that established early in the meeting a sense of cooperation among participants. These proceedings are an outgrowth of this symposium and include 26 papers presented by a truly international cast of contributors. The 'global' nature of the participation in the symposium and these proceedings is indicative of a growing interest in aquatic oligochaetes. This volume will serve to enhance information exchange among oligochaete researchers. Evidence of the international composition of these proceedings is given by a broad geographical basis: France, Italy, Sweden, Federal Republic of Germany, The Netherlands, Ireland, Finland, Argentine, Canada and of course the United States. The Italian constituency provided the largest portion of papers being presented, a total of seven. The content of the presented papers was equally as broad, including studies of taxonomy, pollution ecology, productivity, species interactions, reproductive behavior, morphology, neural biology, genetics, evolutionary relation• ships and much more. The scope of these proceedings as well as the symposium presentations represents a balanced mixture of biological topics. To ensure a high level of scientific quality, all manuscripts were reviewed by outside peer reviewers. The reviewing process is difficult and time consuming, so I wish to thank those individuals who donated both time and effort in making this volume a success. I especially thank Mary Forlenza from the Center for Great Lakes Studies for painstakingly typing and proofing the manuscripts. We have unfortunately lost from our ranks one of our valued colleagues, Dr. Clarence J. Goodnight, who died in August, 1987. We will surely miss his continued contributions to oligochaete biology. I wish to thank Dr. Brinkhurst for authoring the 'In Memoriam' honoring Dr. Goodnight, to whom this volume is dedicated. As a final note, I would like to thank Drs. Harman and Brinkhurst for giving me the opportunity to edit this volume.

JERRY L. KASTER January, 1989 Hydrobiologia 180: IX-X, 1989. J. L. Kaster (ed.) Aquatic Oligochaete Biology IX

Dr. Clarence James Goodnight A.B., A.M., Ph.D. In Memoriam

Dr. C. J. Goodnight, born May 30, 1914 in Gillespie, , died on August 9, 1987. Dr. Goodnight taught at Brooklyn College, City, Purdue University and Western University, Kalamazoo. He was widely travelled, and was associated with many Universities and laboratories. His list of positions with major sciencific societics included the posts of Secretary to the North American (then Midwest) Benthological Society, President of the American Microscopical Society, and Secretary, American Society of Zoologists. Dr. Goodnight one of the first American scientists to contribute to the study of aquatic oligochaetes in relation to pollution, beginning with his key to the group in the 1959 revision of Ward and Whipple. His list of grants and publications indicates a rapid shift from the study of pollution indicators in the early 1960s to an interest in the accumulation of radionuclides by tubificids. This was added to his much earlier interest in the Branchiobdellida, which resulted in publications dating from 1939-1943. His pollution studies, often in association with graduate students, broke new ground and it is quite usual to find his name among the first to publish in each phase of aquatic oligochaete biological work in North America. This is evidenced still in his later papers on spermatogenesis in Limnodrilus, identifying two sperm types and recognizing the effects of phagocytes in the sperm sac. x

Dr. Goodnight made a very significant contribution to the study of aquatic oligochaetes and provided one of the very few centers for the training of graduate students in our field at a significant period in its development. He would have surely contributed to these Proceedings of the first of our meetings to be held in the United States, and this volume is dedicated to his memory. The following citations provide a synopsis of Dr. Goodnight's contributions to oligochaete biology.

R. O. BRINKHURST Sidney, B.C. Canada

1940. New records ofbranchiobdellids (Oligochaeta) and their Crayfish hosts. Jour. Tenn. Acad. Sci. 15: 170-171. 1941. A new branchiobdellid from . J. Parasitol 27 (suppl): 34. 1942. A new branchiobdellid from Kentucky. Trans. am. Micro. Soc. 61: 272-273. 1943. Reports on a collection ofbranchiobdeliids. J. Parasitol29: 100-103. 1954. The effects of various concentrations of ions on the asexual reproduction of Aeolosoma. Proc. Ind. Acad. Sci. 63: 269-270 (with F. I. Kamemoto). The effects of various concentrations of ions on the asexual reproduction of the oligochaete Aeolosoma hemprichi. Trans. am. Micro. Soc. 75: 219-228 (with F. I. Kamemoto). 1957. The ecology of freshwater oligochaetes in West Central . Bull. Ecol. Soc. of America 38: 60. 1959. Oligo chaeta in Freshwater Ecology Edited by W. T. Edmondson. John Wiley and Sons, New York. 1960. Oligochaetes as indicators ofpoliution. Water and Sewage Works 107: 311 (with L. S. Whitley). 1961. Oligochaetes as indicators of pollution. Proceedings of the 15th Annual Industrial Waste Conference, 1960: 139-142 (with L. S. Whitley). 1965. Studies on the chemical composition of several aquatic oligochaetes. Amer. Zool. 5: 220 (with B. K. Whitten). The effect of environmental factors on the respiration of TUbifex. Am. MidI. Nat. 74: 418-428 (with D. J. Fowler). The accumulation of Sr-89 in tubificid worms. Bull. Ecol. Soc. Amer. 46: 100 (with B. K. Whitten). 1966. Stronium-89 and Calcium-45 accumulation in an aquatic oligochaete. Am. Zool. 6: 508 (with B. K. Whitten). Toxicity of some common insecticides to tubificids. J. Water Poll. Cont. Fed. 38: 227-235 (with B. K. Whitten). 1967. The accumulation of Sr-89 and Ca-45 by an aquatic oligochaete. Physiol. Zool. 40: 371-385 (with B. K. Whitten). 1968. The role of tubificid worms in the transfer of radioactive phosphorus in an aquatic ecosystem. Proc. Second National Symposium on Radioecology pp. 270-277. (with B. K. Whitten). 1972. A new species oftubificid oligochaete from Central America, Limnodrilus bulbiphallus n. sp. Trans. am. Microsc. Soc. 91: 579-585 (with E. M. Block). 1973. Relationships between Phosphorus-32 accumulation in algae, bacteria, and tubificids. Proc. Third National Symposium on Radioecology 410-416 (with W. L. Stromberg). 1976. The effect on X-radiation on the coelomic cells of the tubificid Limnodrilus hoffmeisteri. Trans. am. Microsc. Soc. 95: 23-24 (with E. M. Block). 1980. Spermatogenesis in Limnodrilus hoffmeisteri (Annelida, Tubificidae), a morphological study ofthe development of the two sperm types. Trans. Amer. Microsc. Soc. 99: 368-384 (with E. M. Block). 1981. Spermatogenesis in Limnodrilus hoffmeisteri (Oligochaeta, Tubificidae): the role ofphagocytes in the sperm sac. Micron 12: 71-72 (with E. M. Block and J. Mathews). 1982. Observations on the life history of Limnodrilus hoffmeisteri (Annelida, Tubificidae) from the Little Calumet River in temperate North America. Int. J. Invert. Reprod. 4: 239-247.