Cladocera Developments in Hydrobiology 35

Series editor H.J. Dumont Cladocera Proceedings of the Cladocera Symposium, Budapest 1985

Edited by L. Forro and D.G. Frey

Reprinted from Hydrobi%gia 145

1987 DR W. JUNK PUBLISHERS a member of the KLUWER ACADEMIC PUBLISHERS GROUP DORDRECHT/BOSTON/LANCASTER Distributors for the United States'and Canada: Kluwer Academic Publishers, P.O. Box 358, Accord Station, Hingham, MA 02018-0358, USA for the UK and Ireland: Kluwer Academic Publishers, MTP Press Limited, Falcon House, Queen Square, Lancaster LA! lRN, UK for all other countries: Kluwer Academic Publishers Group, Distribution Center, P.O. Box 322, 3300 AH Dordrecht, The Netherlands

Library of Congress Cataloging in Publication Data

Cladocera Symposium (1985 : Budapest, Hungary) C1adocera : proceedings of the C1adocera Symposium, Budapest, 1985.

(Developments in hydrobio1ogy ; 35) 1. C1adocera--Congresses. I. Forro, L. II. Frey, David G. (David Grover), 1915- III. Title. IV. Series. QL444.B83C58 1985 595.3'2 86-18558 ISBN- 13: 978-94-010-8292-1

ISBN-13: 978-94-010-8292-1 e-ISBN-13: 978-94-009-4039-0 DOl: 10.1007/978-94-009-4039-0

Copyright

© 1987 by Dr W. Junk Publishers, Dordrecht. Sof tcover reprint of the hardcover 1st edition 1987

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 of the publishers, Dr W. Junk Publishers, P.O. Box 163, 3300 AD Dordrecht, The Netherlands. Preface

There is a strong tendency now for specialists in various groups of organisms, or in various processes and their consequences, to get together at more or less regular intervals to discuss their subjects. This has hap• pened e.g. with the Chironomidae, Ephemeroptera, and Plecoptera among the aquatic insects, and the Rotifera, Copepoda, and Ostracoda among the zooplankton. Finally the scientists working with the Cladocera, who have been isolated from one another for a long time except on a personal basis, have decided to do the same, largely through the initiative and drive of L. Forro. Thus, the First International Symposium on Cladocera was held in Budapest on 12-16 August 1985. Most of the roughly 60 attendees were from Euro• pean countries, although a fair number came from Australia, Canada, India, Nigeria, and the United States. The accompanying table presents a list of attendees by country of origin. Regrettably, a number of persons who had expressed interest in coming were unable to do so. Their names and, in two instances countries, enclosed in brackets, are given to indicate the overall extent of topics that would have been discussed. Persons who presented papers or posters are indicated by an asterisk (*). The names of persons who chaired sessions are printed in italics. A total of 34 papers and 10 posters was presented, most of which are included in this volume. Besides three days for presentation of papers, there was a half day for discussion (,Mediterranean Workshop') and another half day for an excursion to the bend of the Danube. Most of the work in organizing this symposium was done by L. Forro and his associates in aquatic studies in Hungary, aided by an International Organizing Committee consisting of H. J. Dumont (Belgium), L. For• ro (Hungary), D. G. Frey (USA), G. Fryer (Great Britain), and N. N. Smirnov (Soviet Union). Logistic ar• rangements for the meeting were handled largely by IBUSZ, the Hungarian tourist agency. Besides getting specialists from around the World together to talk about their own particular interests in Cladocera for the first time, the group has decided to have further symposia at 3-year intervals, the next of which will be held in Czechoslovakia in 1988. It has also initiated efforts to establish a newsletter to keep investigators informed of activities and progress in the various laboratories. Other positive results are hoped for. For example, at present there is no central source of information to which one can turn to learn about where type material of various taxa is stored and what voucher material is also available. Because of the ravages of wars and other episodes of uncertain curation, many of the older specimens have been lost completely. An inventory of such materials by country or prominent scientist would be most valuable. Anyone reading this volume will see immediately that many topics relating to the Cladocera are represent• ed. The answer to the question, 'What are Cladocera?' , is that this is not a monophyletic group but probably includes four separate orders. We are all interested in the evolution of Cladocera, which ultimately involves knowledge of genetic processes. Persons working with the Daphniidae, principally various species in Daphnia and Simocepha/us, have been able to demonstrate by electrophoresis that hybridization occurs naturally in nature, although there seems to be little introgression between the parents. On the one hand, it is presumed that the great diversity of Bosmina in the Baltic Region resulted from hybridization between a few invading species in the early Holocene, but, on the other hand, paleolimnological studies of lakes in northern Germa• ny do not reveat the sequence of taxa predicted. Natural hybridization in the Chydoridae seems not to occur VI at all, or at least, if it does, to result in any gene exchange between parent populations. Distributions of spe• cies are being disrupted by man, such as the recent introductions of two species of Moina from North Ameri• ca and East Asia into the ricefields of Italy. Except for such man-related movements, however, the distribu• tion of the Chydoridae is stabile. Many of what are called species today are in reality groups of species, with, seemingly, a different group member(s) on each continent. There seems to be no true cosmopolitanism in this family. Very careful studies on feeding by Daphnia in Lake Balaton have demonstrated that these animals depend not so much on algae for their energy and substance as on the organic films that cover the mineral silt and clay particles in the water. Other studies reported are concerned in nice detail with processes in the Cladocera - competition among species and how a number of closely related species eating much the same food can co-exist at the same time, or birth, growth, and mortality of the Bosmina population in the River Danube in Hungary, and the detailed response of cladoceran communities to changes in lakes resulting from changes in watershed conditions, such as those induced by acid precipitation. This is a beginning. The symposium was highly stimulating to the participants, and indications are that the enthusiasm generated will lead to various types of cooperation, and eventually will lead to a comprehen• sive understanding of the Cladocera, their evolution and their involvement in various ecological processes.

D. G. Frey & L. Forr6

[ARGENTINA] FINLAND [Paggi, 1. C. (Santo Tome S. E)] *Ranta, E. (Helsinki) AUSTRALIA [Sarvala, 1. (Turku)] *Benzie, 1. A. H. (Canberra) *Uimonen-Simola, P. (loensuu) *Shiel, R. 1. (Adelaide) FRANCE [Timms, B. V. (Cooranbong)] *Amoros, C. (Lyon) AUSTRIA [Rey, 1. (Toulouse)] *Moritz, C. (Innsbruck) GERMAN DEMOCRATIC REPUBLIC (DDR) *Unterweger, A. T. (Salzburg) *Flossner, Mr. & Mrs. D. (lena) *Wiener, W. (Salzburg) [Horn, W. (Wiinschendorf)] BELGIUM *Lieder, U. (Berlin) *Dumont, H J. (Gent) GERMAN FEDERAL REPUBLIC (BRD) CANADA *Hojmann, W. (PIon) *Boucherle, M. M. (Kingston, Ont.) Hollwedel, Mr. & Mrs. W. (Varel) *Chengalath, R. (Ottawa, Ont.) *Rieder, Mr. & Mrs. N. (Karlsruhe) *Hann, B. 1. (Winnipeg, Man.) *Wolf, H. G. (PIOn) *Hebert, P. D. N. (Windsor, Ont.) HUNGARY *Paterson, M. 1. (Halifax, N. S.) *Bothar, A. (God) *Schwartz, S. S. (Windsor, Ont.) *Entz, B. (Tihany) CZECHOSLOVAKIA Forr6, L. (Budapest) *Hrbacek, 1. (Praha) *G.-T6th, L. (Tihany) *Hudec, 1. (Kosice) INDIA *Korinek, V. (Praha) [Battish, S. K. (Ludhiana)] Stifter, P. (Benatky nad lizerou) *Michael, R. G. (Shillong) DENMARK [Sharma, B. K. (Shillong)] *R0en, Mr. & Mrs. U. (K0benhavn) IRELAND ENGLAND *Duigan, C. A. (Dublin) Dayton, 1. (Birmingham) ITALY *Fryer, G. (Ambleside) *de Bernardi, R. (Pallanza) *Crosetti, D. (Roma) VII

*Manca, M. (Pallanza) USA *Margaritora, Mrs. & Mr. F. G. (Roma) *Baas, D. (Bloomington, IN) [Morocco] *Berner, D. B. (Philadelphia, PA) [Lamoot, E. (Azrou)] *Cotten, C. A. (Bloomington, IN) [Ramdani, M. (Rabat)] * Frey, Mr. & Mrs. D. G. (Bloomington, IN) NIGERIA [USSR] *Egborge, A. B. M. (Benin) [Aladin, N. V. (Leningrad)] NORWAY [Glagolev, S. M. (Moskva)] [Klemetsen, A. (Troms0)] [Korovchinsky, N. M. (Moskva)] Nilssen, J. P. (Oslo) [Makrushin, A. V. (Leningrad)] ROMANIA [Radzimovsky, V. D. (Kiev)] *Negrea, Mr. & Mrs. S. (Bucuresti) [Rivier, I. K. (Borok)] SPAIN [Smirnov, N. N. (Moskva)] *Alonso, M. (Barcelona) [Villar, R. V. (Moskva))" SWEDEN VENEZUELA *Bengtsson, J. (Uppsala) *Zoppi de Roa, E. (Caracas) Milbrink, G. (Uppsala)

1. Goran Milbrink 17. Catherine Duigan 33. Jan Bengtsson 2. ? 18. John A. H. Benzie 34. Donatella Crosetti 3. Riccardo de Bernardi 19. Jaroslav Hrb.kek 35. Bela-Entz 4. Ulrik R0en 20. Rama Chengalath 36. Hans Georg Wolf 5. Ulrich Lieder 21. Clara Cotten 37. Fiorenza G. Margaritora 6. Werner Hollwedel 22. R. George Michael 38. Anna Bothar 7. Karl Kraus 23. Evelyn Zoppi de Roa 39. Igor Hudec 8. Claude Amoros 24. Wolfgang Wiener 40. Norbert Rieder 9. Dorothy B. Berner 25. Andreas Unterweger 41. Laszlo G-Toth 10. Mary M. Botl,cherle 26. Paul D. N. Hebert 42. Pavel Stifter 11. Michael Paterson 27. Dietrich FlOssner 43. Marina Manca 12. Jens Petter Nilssen 28. Christian Moritz 44. Brenda Hahn 13. Judith Dayton 29. Wolfgang Hofmann 45. Laszlo Forro 14. Esa Ranta 30. Steven S. Schwartz 46. Henri J. Dumont 15. Miguel Alonso Garcia -Amilibia 31. Vladimir Korinek 47. David G. Frey 16. Geoffrey Fryer 32. Pirjo Uimonen-Simola Contents

Preface...... V

TAXONOMY & SYSTEMATICS

Dr. Jene Daday de Dees and the Hungarian hydrobiological research by B. Entz & D. Lukacs ...... 1 The Taxonomy and biogeography of the Cladocera by D. G. Frey ...... 5 Morphology and the classification of the so-called Cladocera by G. Fryer...... 19 Review of taxonomic studies on freshwater cladocera from India with remarks on Biogeography by B. K. Sharma & R. G. Michael ...... 29 Revision of three species on the genus Diaphanosoma Fischer, 1850 by V. Korinek ...... 35 A paedomorphic form type of Daphnia triquetra Sars 1903 (Cladocera, Daphniidae) from Mongolia by D. FleBner ...... 47 The biogeography of Australian Daphnia: clues of an ancient (> 70 m.y.) origin for the genus by J. A. H. Benzie ...... 51 Is ephippial morphology a useful taxonomic descriptor in the Cladocera? An examination based on a study of Daphniopsis (Daphniidae) from Australian salt lakes by M. J. Kokkinn & W. D. Williams ...... 67 Significance of head and carapace pores in Ceriodaphnia (Crustacea, Cladocera) by D. B. Berner...... 75 Inter- and Intraspecific variations in limb morphology of some south-east Australian cladocerans: Im• plications for particle capture by R. J. Shiel & G. G. Ganf ...... 85 A Far East Moina, M. weismanni Ishikawa, 1896 found in an Italian ricefield by F. G. Margaritora, 1. Ferrari & D. Crosetti ...... 93 Scapholeberis erinaceus Daday, 1903 (syn S. echinulata Sars, 1903) and Ilyocryptus alexandrinae sp. nov. in Roumania by S. Negrea ...... 105 A contribution to the taxonomy of C. sphaericus sens. lat. (Cladocera, Chydoridae) by C. A. Duigan & D. A. Murray ...... 113 Chydol'us arcticus n.sp., a new cladoceran crustacean (Chydoridae: Chydorinae) from the North Atlan- tic Arctic and Subarctic areas by.U. R0en ...... 125 x

Morphological Differentiation of two new Ephemeroporus species (Cladocera, Chydoridae) belonging to the barroisi complex: E. margalefi and E. epiaphantoii, in Spain by M. Alonso...... 131 Moina oryzae n.sp. (Cladocera, Moinidae) from Tamil Nadu (South India) by 1. Hudec...... 147

DISTRmUTION & GENETICS

The distribution of chydorid Cladocera in Canada by R. Chengalath ...... 157 Salinity and the distribution of Cladocera in Warri River, Nigera by A. B. M. Egborge ...... 159 Groundwater Cladocera: A synopsis by H. J. Dumont...... 169 The ultrastructure of the so-called olfactory setae on the antennula of Daphnia magna Straus (Crustacea, Cladocera) by N. Rieder...... 175 Genotypic characteristics of the Cladocera by P. D. N. Hebert...... 183 Breeding system of Daphniopsis ephemeralis: adaptions to a transient environment by S. S. Schwartz & P. D. N. Hebert...... 195 The possible origin of Daphnia cucullata procurva Poppe 1887 in the lakes of the Pomeranian Lakeland by hybridization in the past by U. Lieder ...... 201 Interspecific hybridization between Daphnia hyalina D. galeata and D. cucullata and seasonal abun• dances of these species and their hybrids by H. G. Wolf...... 213 Naturally occurring interspecific hybridization in Simocephalus (Cladocera, Daphniidae): its potential significance by B. J. Hann ...... 218

ECOWGY, POPULATION STUDffiS & HISTORY

Cladocera: Predators and prey by R. de Bernardi, G. Giussani & M. Manca ...... 225 Competitive dominance among Cladocera: Are single-factor explanations enough? An examination of the experimental evidence by J. Bengtsson ...... 245 Size and shape of Daphnia Longispina in rock-pools by E. Ranta & S. Tjossem ...... 259 Feeding and energy budget estimations in Daphnia obtusa by M. Manca & R. de Bernardi ...... 269 A population study of Scapholeberis rammneri Dumont and Pensaert (Cladocera: Daphniidae) by H. J. Dumont ...... 275 Estimation of production and mortality of Bosmina longirostris (0. F. Muller) in the River Danube (Danubiala Hungarica, CIX) by A. Botha! ...... 285 XI

Reproductive variations of Cladocerans in grasslands periodically flooded for irrigation in Mantecal, Venezuela by B. Jimenez & E. Zoppi de Roa ...... 293 Observations on the zooplankton in the reedbelt area of the Neusiedlersee by L. Forro & H. Metz ...... 299 A note on the hatching and viability of Ceriodaphnia ephippia collected from lake sediment by C. Moritz ...... 309 Cladocera in space and time: Analysis of lake sediments by W. Hofmann ...... 315 Alga consumption of our dominant planktonis crustaceans in Lake Balaton (Hungary) by L. G. Toth, N. P. Zankai & O. M. Messner ...... 323 The dead-arm evolution of river systems: A comparison between the information provided by living Copepoda and Cladocera populations and by Bosminidae and Chydoridae remains by C. Amoros & C. Jacquet ...... Effects of recent acidification on Cladocera in small clear-water Lakes studied by means of sedimen• tary remains by P. Uimonen Simola & P. K. Tolonen ...... 343