Ecology and Morphology of Copepods Developments in Hydrobiology 102

Series editor H. J. Dumont Ecology and Morphology of Copepods

Proceedings of the 5th International Conference on Copepoda, Baltimore, USA, June 6-13, 1993

Edited by Frank D. Ferrari & Brian P. Bradley

Reprinted from Hydrobiologia, vo/s 2921293 (1994)

Springer-Science+Business Media, BV. Library of Congress Cataloging-in-Publication Data

A C.I.P. Catalogue record for this book is available from the Library of Congress.

ISBN 978-90-481-4490-7 ISBN 978-94-017-1347-4 (eBook) DOI 10.1007/978-94-017-1347-4

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AII Rights Reserved © 1994 Springer Science+Business Media Dordrecht Originally published by Kluwer Academic Publishers in 1994 No part of the material protected by this copyright notice may be reproduced or utilized in any form or by any means, electronic or mechanical including photocopying, recording or by any information storage and retrieval system, without written permission from the copyright owner. v

Contents

Preface...... ix

Photograph and List of Participants x

Maxilliped lecture How many copepods? by A.G. Humes 1

Systematics Acartia tonsa: a species new for the Black Sea fauna by G. Belmonte, M.G. Mazzocchi, I.Y. Prusova & N.V. Shadrin ...... 9 A new species of Erebonectes (Copepoda, Calanoida) from marine caves on Caicos Islands, West Indies by A. Fosshagen & T.M. Iliffe ...... 17 Nomenclature, redescription, and new record from Okinawa of Cymbasoma morii Sekiguchi, 1982 (Monstrilloida) by M.J. Grygier ...... 23 Copepod phylogeny: a reconsideration of Huys & Boxshall's 'parsimony versus homology' by J-S. Ho ...... 31 Mesocyclops thermocyclopoides species-group: redefinition and content by M. Holynski & F. Fiers ...... 41 Variation in the species of freshwater harpacticoid copepods in Japan. II. Attheyella nakaii (Brehm) by T. Ishida ...... 53 Glaciella, a new genus of freshwater Canthocamptidae (Copepoda, Harpacticoida) from a glacier in Nepal, Himalayas by Y. Kikuchi...... 59 Descriptions ofAcartia (Euacartia) southwelli Sewell 1914 and Acartia (Euacartia) sarojus n.sp. from India and status of the subgenus Euacartia Steuer 1923 by M. Madhupratap & P. Haridas ...... 67 Morphometric study of Calanus chilensis males along the Chilean coast by V. Marin, S. Espinoza & A. Fleminger ...... 75 Boeckella diamantina n.sp. (Callanoida, Centropagidae), from a high Andean lake in Mendoza, Argentina by S.A. Menu-Marque & L.R. Zuftiga ...... 81 Haplostomides hawaiiensis, new species (Copepoda: Cyclopoida: Ascidicolidae), associated with the ascidian Polyclinum constellatum at Honolulu, Hawaii by S. Ooishi ...... 89 Tropodiaptomus zambeziensis, T. bhangazii and T. capriviensis, three new species of Tropodiaptomus (Copepoda, Calanoida) from southern Africa by N.A. Rayner ...... 97 vi

Troglocyclops janstocki, new genus, new species, a very primitive cyclopid (Copepoda: Cyclopoida) from an anchialine cave in the Bahamas by C.E.F. da Rocha & T.M. !liffe ...... 105 Senecella siberica n.sp. and the position of the genus Senecella in Calanoida classification by N.V. Vyshkvartzeva ...... 113 A clarification of two congeners, Pseudodiaptomus lobipes and P. binghami (Calanoida, Pseudodi• aptomidae) from India, with description of P. mixtus sp.n. from Bangladesh by T.C. Walter ...... 123

Morphology Morphological aspects of subitaneous and resting eggs from Acartia josephinae (Calanoida) by G. Belmonte & M. Puce ...... 131 The integumental ultrastructure of Parathalestris harpactoides (Claus, 1863) (copepoda, harpacti• coida) by J. Bresciani & H.-U. Dahms...... 137 Importance of the genital operculum in the taxonomy of South American Diaptomidae (Copepoda, Calanoida) by G. Cicchino ...... 143 Morphological variability of geographically distinct populations of the estuarine copepod Acartia Tonsa by T.G. Garmew, S. Hammond, A. Mercantini, J. Morgan, C. Neunert & J.A. Fornshell. 149 Speculations on enigmatic structures in relation to associated copepod phylogeny by R.V. Gotto ...... 157 Copepodid stages of Conchyliurus quintus Tanaka, 1961 (Poecilostomatoida, Clausidiidae) asso• ciated with bivalve mollusks by I-H. Kim ...... 161 External morphology of the male and female of Sphyrion lumpi (Kmyer, 1845) (Copepoda; Siphonostomatoida; S phyriidae) by J.D.W. Moran & W. Piasecki ...... 171 Variability of Acartia clausi in the Black Sea by N.V. Shadrin & E.V. Popova ...... 179

Feeding and reproduction Induction of diapause egg production in Eurytemora affinis by their own metabolites by S. Ban & T. Minoda ...... 185 Seasonal feeding and fecundity of the calanoid copepod Acartia tonsa in Long Island Sound: is omnivory important to egg production? by H.G. Dam, W.T. Peterson & D.C. Bellantoni...... 191 Physiological characteristics of the antarctic copepod Calanoides acutus during late summer in the Weddell Sea . by A.V. Drits, A.F. Pasternak & K.N. Kosobokova ...... 201 Diapause dynamics of two diaptomid copepod species in a large lake by N.G. Hairston, Jr. & R.A. van Brunt ...... 209 Reproduction of the calanoid copepod Calanus propinquus in the southern Weddell Sea, Antarc- tica: observations in laboratory by K.N. Kosobokova ...... 219 VB

Reproductive behaviour in the harpacticoid copepod Tigriopus fulvus by 1. Lazzaretto, F. Franco & B. Battaglia ...... 229 Ingestion of the cyanobacterium Trichodesmium spp. by pelagic harpacticoid copepods Macrose- tel/a, Mirada and Oculosetella by J.M. O'Neill & M.R. Roman ...... 235 Gut fluorescence in herbivorous copepods: an attempt to justify the method by A.F. Pasternak ...... 241 Copepod grazing in a summer cyanobacteria bloom in the Gulf of Finland by K.G. Sellner, M.M. Olson & K. Kononen ...... 249 Surface glycoproteins in copepods: potential signals for mate recognition by T.W. Snell & M.J. Carmona ...... 255

Distributions in time and space Labidocera aestiva and L. scotti in Tamiahua Lagoon, Veracruz, Mexico by C. Alvarez-Silva & S. Gomez-Aguirre ...... 265 The microcopepod fauna in the Eastern Mediterranean and Arabian Seas: a comparison with the Red Sea Fauna by R. Bottger-Schnack ...... 271 Seasonal variations in carbon and nitrogen content of Acartia clausi (Copepoda, Calanoida) in the Gulf of Trieste (Northern Adriatic Sea) by B. Cataletto & S.F. Vmani ...... 283 Life cycle of Calanus chilensis Brodsky in Bay of San Jorge, Antofagasta, Chile by R. Escribano & L. Rodriguez ...... 289 Copepod nauplii from the barrier reef of Belize by J.A. Fornshell ...... 295 Seasonal succession of Copepoda in the Bahia Blanca estuary by M.S. Hoffmeyer ...... 303 Spermatophore transfer in Euchaeta species in a 2000 m water column by J. Mauchline ...... 309 Geographic distribution of the bathypelagic genus paraeuchaeta (Coepoda, Calanoida) byT.Park ...... 317 Diel changes in copepod respiration rates by E.V. Pavlova ...... 333 Latitudinal diversity patterns of continental benthic copepod species assemblages in the Americas by J.W. Reid...... 341 A calanoid copepod Gladioferens imparipes, holding to surfaces by R.J. Rippingale ...... 351 Autecology of an intertidal Alaskan harpacticoid copepod Apolethon sp. by N.V. Schizas & T.C. Shirley ...... 361 Diversity and structure of pelagic copepod populations in the frontal zone of the eastern Alboran Sea by G. Seguin, A. Errhif & S. Dallot ...... 369 Freshwater littoral copepods of British Columbia and Alberta (Canada) by C.-T. Shih & R. Chengalath ...... 379 viii

Diel vertical migration and gut pigment rhythm of Paracalanus parvus, P. crassirostris, Acattia erythraea and Eucalanus subcrassus (Copepoda, Calanoida) in Tolo harbour, Hong Kong by K.W. Tang, Q.c. Chen & c.K. Wong ...... 389 Seasonal variability in the individual carbon content of the calanoid copepod Acartia bifilosa from the northern Baltic Sea by S. Tanskanen ...... 397 Planktonic copepods of Boston Harbor, Massachusetts Bay and Cape Cod Bay, 1992 by J.T. Turner ...... 405 Seasonal dynamics of Acartia bifilosa and Eurytemora affinis (Coepoda: Calanoida) in relation to abiotic factors in the northern Baltic Sea by M. Viitasalo, T. Katajisto & I. Vuorinen ...... 415 Life history traits of the tropical freshwater copepod Mesocyclops longisetus (Crustacea: Copepoda) by G.A. Wyngaard, C.E. Goulden & A. Nourbakhsh ...... 423 Species assemblages of cyclopoid Copepoda from Mantecal, Venezuela by E.Z. de Roa ...... 429

Environmental relationships Trophic roles of particle feeders and Detritus in a mangrove island prop root ecosystem by J.W. Ambler, J. Alcala-Herrara & R. Burke...... 437 Factors affecting feeding selectivity of visual predators on the copepod Acartia tonsa: locomotion, visibility and escape responses by E.J. Buskey ...... 447 Commensal life, sexual dimorphism, and handedness in the canuellid harpacticoid Parasunaristes chelicerata (Por & Marcus, 1972) by C.L. Falck & T.E. Bowman ...... 455 Salinity stress proteins in Eurytemora affinis by C.R.M. Gonzalez & B.P. Bradley ...... 461 Development of Acartia clausi (Copepoda, Calanoida) cultured at different conditions of tempera- ture and food by W.C.M. Klein Breteler & N. Schogt ...... 469 Coexistence and competition within Acartia (Copepoda, Calanoida) congeners from Lebanese coastal water: nice overlap measurements by S. Lakkis ...... 481 Use of cyclopoid copepods for mosquito control by G.G. Marten, E.S. Bordes & M. Nguyen ...... 491 Effects of impounding coastal salt marsh for mosquito control on microcrustacean populations by E. Ruber, A. Gilbert, P.A. Montagna, G. Gillis & E. Cummings ...... 497 Sublethal effects of cupric ion activity on the phototaxis of three calanoid copepods by D.E. Stearns & A.A. Sharp...... 505 Replacement of large copepods by small ones with eutrophication of embayments: cause and consequence by S.-I. Uye ...... 513 The role of copepods in the planktonic ecosystems of mixed and stratified waters of the European shelf seas by R. Williams, D.V.P. Conway & H.G. Hunt ...... 521 Hydrobiologia 292/293: 1994. F.D. Ferrari & B,P. Bradley (eds), Ecology and Morphology of Copepods. IX

Preface

The Fifth International Conference on Copepoda was held on the Baltimore County Campus of the University of Maryland in Baltimore, USA, from June 6 to June 13, 1993. About 170 copepodologists from 35 countries took part. The Conference was sponsored jointly by the University of Maryland• Baltimore County (UMBC) and the Smithsonian Institution (SI). The Local Organizing Committee consisted of Thomas Bowman (SI), Brian Bradley (UMBC), Frank Ferrari (SI), Elvie Fornshell (SI), John Fornshell (Thomas Jefferson High School for Science and Technology), Janet Reid (SI), Chad Walter (SI), and Grace Wyngaard (James Madison University). Financial support from the University of Maryland-Baltimore County, the Smithsonian Institution, the International Science Foundation, the Maryland Agricultural Experiment Station, the Chesapeake Research Consortium, the National Aquarium in Baltimore, and the World Association of Copepodologists was critical to the implemen• tation of the Conference. Rosel Halle and Carolyn Harriger of the Office of Continuing Education (UMBC), as well as staff and volunteers from both the University of Maryland-Baltimore County and the Smithsonian Institution, helped make the conference a success. The program of the Conference included both platform and poster sessions on a variety of research subjects currently being pursued by copepodologists. The presentations were organized under the following general topics: Behavior, Feeding, Genetics, Horizontal Variations, Morphology, Phylogeny, Reproduction, Seasonal Changes, Vertical Distribution plus two special sessions on copepods of the genus Acartia and cyclopidjmosquito interactions. The Maxilliped Lecture, given by Dr Arthur G. Humes of Boston University, clearly establishes the importance of copepods to the earth's organic diversity. The Proceedings of the Conference consists of selected research articles from the different sessions. We wish to thank our editorial committee (Elvie Fornshell, John Fornshell, Janet Reid, Chad Walter, Grace Wyngaard) and over 150 colleagues throughout the world who reviewed the contents of these Proceedings. The articles published here reflect the diverse research interests of copepodologists today, and are distinguished by their high quality. Their impact will ensure that this volume is consulted by a wide range of research biologists.

FRANK FERRARI & BRIAN BRADLEY Editors

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1 Patricia A. Tester 27 Donald E. Stearns 52 Terry W. Snell 77 Ksenia N. Kosobokova 102 Mark D. Ohman 2 Ludmila Sazhina 28 Jens Christian Nejstgaard 53 Alessandra Comas chi 78 Frank Kurbjeweit 103 Markku Viitasalo 3 Maria Auxiliadora Amado 29 Carlos Eduardo F. da Rocha 54 Bruno Cataletto 79 John R. McConaugha 104 Nelson G. Hairston, Jr. 4 Ju-shey Ho 30 C. Kim Wong 55 Serena Fonda-Umani 80 Cornelia Metz 105 Nancy A. Rayner 5 Brian Bradley 31 Makoto Terazaki 56 Teruo Ishida 81 Anna F. Pasternak 106 Pamela Arnofsky 6 Arthur G. Humes 32 Unknown 57 Wojciech Piasecki 82 Gerd Schriever 107 Elke Willen 7 Geoffrey A. Boxshall 33 Marie H. Bundy 58 Wen-Been Chang 83 Robert J. Rippingale 108 Hans-U. Dahms 8 Horst Kurt Schminke 34 Mari Butler 59 Lisa A. Drake 84 Gunther Steinbruck 109 Darcy J. Lonsdale 9 Ian A. McLaren 35 T. Chad Walter 60 Heidi Dorward 85 Thomas Glatzel 110 Gustav-A. PaffenMfer 10 Suzanne Razouls 36 Won Choel Lee 61 Mai D.G. Lopez 86 Syuhei Ban 111 J. Rudi Strickler 11 Sami Lakkis 37 Nicolas V. Shadrin 62 Liana Talaue McManus 87 Tomohiko Kikuchi 112 Paul Trygve Solberg 12 Karen Gresty 38 Bruno Battaglia 63 Ruth Bottger-Schnack 88 Pamela Blades-Eckelbarger 113 Klaus Kohlhage 13 Stewart C. Johnson 39 Evelyn Zoppi de Roa 64 Hans Verheye 89 Jeffery R. Cordell 114 Barbara Hosfeld 14 Nelson Santos-Silva 40 Almeida Prado-Por 65 W. Klein Breteler 90 Nikolaos Schizas 115 Ulf Bamstedt 15 Gerard Seguin 41 Francis Dov Por 66 Constantine H. Fernando 91 John F offishell 116 John Mauchline 16 Shin-Ichi Uye 42 Elizabeth Pavlova 67 Yoshiaki Kikuchi 92 Jack G. Greenwood 117 Angus Atkinson 17 Shuhei Nishida 43 Chintala Rama Devi 68 Tatsuki Toda 93 Robert Williams 118 Isobel Hopkins Mauchline 18 Xinsheng Zhang 44 Jeanette Yen 69 Nancy H. Marcus 94 Jefferson T. Turner 119 Ernest Ruber 19 Sigrid B. Schnack Schiel 45 Unknown 70 Elvie Fornshell 95 Genuario Belmonte 120 Gisela Moura 20 Shigeko Ooishi 46 Hans G. Dam 71 Karen F. Wishner 96 Sanna Tanskanen 121 Audun Fosshagen 21 Chang-Tai Shih 47 Peter P. Jaros 72 Ed Buskey 97 Ulrich Einsle 122 Elena Lvovna Markhaseva 22 Rubens M. Lopes 48 Silvina A. Menu-Marque 73 Serge Andre Poulet 98 Taisoo Park 123 M. Madhupratap 23 Guilherme Ribeiro Lotufo 49 Janet W. Reid 74 Adrianna Ianora 99 John B.J. Wells 124 Jonathan Moran 24 Grace A. Wyngaard 50 Maria K. Holynski 75 Massimiliano Cervelli 100 B.H. Ross Othman 125 Bong-Choel Oh 25 Ruben Escribano 51 Julie Ambler 76 Judith M. O'Neil 101 Eduardo Suarez Morales 126 Pedro Martinez-Arbizu

26 Victor H. Marin ~.