Table Of ContentCopepoda: Developments in Ecology,
Biology and Systematics
Developments in Hydrobiology 156
Series editor
H. J. Dumont
Copepoda: Developments in Ecology,
Biology and Systematics
Proceedings of the Seventh International Conference on Copepoda,
held in Curitiba, Brazil, 25–31 July 1999
Edited by
Rubens M. Lopes1, Janet W. Reid2 & Carlos E. F. Rocha3
1
State University of Santa Cruz, llhéus, Brazil
2
National Museum of Natural History, Washington, DC, U.S.A.
3
University of São Paulo, Brazil
Reprinted from Hydrobiologia, volumes 453/454 (2001)
KLUWER ACADEMIC PUBLISHERS
NEW YORK, BOSTON, DORDRECHT, LONDON, MOSCOW
eBookISBN: 0-306-47537-5
Print ISBN: 0-7923-7048-1
©2002 Kluwer Academic Publishers
NewYork, Boston, Dordrecht, London, Moscow
Print ©2001 Kluwer Academic Publishers
Dordrecht
All rights reserved
No part of this eBook maybe reproducedor transmitted inanyform or byanymeans,electronic,
mechanical, recording, or otherwise,withoutwritten consent from the Publisher
Createdin the UnitedStates of America
Visit Kluwer Online at: http://kluweronline.com
and Kluwer's eBookstoreat: http://ebooks.kluweronline.com
v
TABLE OF CONTENTS
Preface ix–xi
Group photograph xii
List of participants xiii
Maxilliped Lecture
Why do symbiotic copepods matter?
Ju-shey Ho 1–7
Biology and ecology of Antarctic and Subantarctic copepods
Aspects of the study of the life cycles of Antarctic copepods
Sigrid B. Schnack-Schiel 9–24
Feeding patterns of dominant Antarctic copepods: an interplay of diapause, selectivity,
and availability of food
Anna F. Pasternak, Sigrid B. Schnack-Schiel 25–36
The deep-sea copepod fauna of the Southern Ocean: patterns and processes
Peter Ward, Rachael S. Shreeve 37–54
Production and fate of faecal pellets during summer in an East Antarcticfjord
K.L. Beaumont, A.J. Plummer, G.W. Hosie, D.A. Ritz 55–65
Effects of the ice-edge bloom and season on the metabolism of copepods in the Weddell
Sea, Antarctica
Helena G. Kawall, Joseph J. Torres, Stephen P. Geiger 67–77
The effect of the receding ice edge on the condition of copepods in the northwestern
Weddell Sea: results from biochemical assays
Stephen P. Geiger, Helena G. Kawall, Joseph J. Torres 79–90
Ecology, distribution and biology of marine copepods
Patterns in stage duration and development among marine and freshwater calanoid
and cyclopoid copepods: a review of rules, physiological constraints, and evolutionary
significance
William T. Peterson 91–105
vi
Feeding, eggproduction, andegghatchingsuccessofthecopepodsAcartia tonsaand
Temora longicornis on diets of the toxic diatom Pseudo-nitzschia multiseries and the
non-toxic diatomPseudo-nitzschia pungens
Jean A. Lincoln, Jefferson T. Turner, Stephen S. Bates, Claude Léger, David A. Gauthier 107–120
Summer egg production rates of paracalanid copepods in subtropical waters adjacent
to Australia’s North West Cape
A.D. McKinnon, S. Duggan 121–132
Size paradigms in copepod communities: a re-examination
R.R. Hopcroft, J.C. Roff, F.P. Chavez 133–141
The influence of coastal upwelling on the distribution of Calanus chilensis in the
Mejillones Peninsula (northern Chile): implications for its population dynamics
Ruben Escribano, Victor H. Marin, Pamela Hidalgo 143–151
Succession of pelagic copepod species in coastal waters off northern Chile: the
influence ofthe1997–98 ElNiño
Pamela Hidalgo, Ruben Escribano 153–160
Spatial distributions of copepod genera along the Atlantic Meridional Transect
Rachel S. Woodd-Walker 161–170
Biogeographyof the familyAcartiidae(Calanoida)in the Ponto-MediterraneanProvince
G. Belmonte, D. Potenza 171–176
Temporal variability and production ofEuterpina acutifrons (Copepoda: Harpacticoida)
in the Cananéia Lagoon estuarine system, São Paulo, Brazil
Koichi Ara 177–187
Harpacticoid copepod communities of floating seaweed: controlling factors and implic-
ations for dispersal
Emil Ólafsson, Agnar Ingólfsson, María Björk Steinarsdóttir 189–200
Copepods in challenging environments
A human challenge: discovering and understanding continental copepod habitats
Janet W. Reid 201–226
Groundwater copepods: diversity patterns over ecological and evolutionary scales
Diana M.P. Galassi 227–253
Harpacticoid copepods aresuccessful in thesoft-bottom deep sea
David Thistle 255–259
Ecology of freshwater copepods
Effect of the cyclopoid copepod Mesocyclops thermocyclopoides on the interac-
tions between the predatory rotifer Asplanchna intermedia and its prey Brachionus
calyciflorus andB. angularis
Ram Kumar, T. Ramakrishna Rao 261–268
Two calanoids,twolakes, andadecade ortwo.Anupdated record and evaluation ofoc-
currence and periodicity ofTropodiaptomus spectabilisandMetadiaptomus meridianus
(Copepoda: Calanoida), and alternative stable states in two cascading impoundments
Rob C. Hart 269–283
Life cycles of the two freshwater copepods Cyclops strenuus Fischer and Cyclops
insignisClaus (Cyclopoida, Copepoda) in an amphibious floodplain habitat
Dagmar Frisch 285–293
vii
Diapause in copepods (Crustacea) from ephemeral habitats with different hydroperiods
in Everglades National Park (Florida, U.S.A.)
Maria Cristina Bruno, William F. Loftus, Janet W. Reid, Sue A. Perry 295–308
Distribution and ecology of copepods in mountainous regions of the Eastern Alps
C.D. Jersabek, A. Brancelj, F. Stoch, R. Schabetsberger 309–324
Copepod communities in karstic mediterranean lakes along the eastern Adriatic coast
M. Kerovec, 325–333
Cyclopoid diversity in the basin of Lake Hula (Israel), after its partial reflooding
F.D. Por, Ch. Dimentman 335–339
Biogeography ofcopepods in lakes and ponds ofsubarctic Québec, Canada
Kerrie M. Swadling, John A.E. Gibson, Reinhard Pienitz, Warwick F. Vincent 341–350
The distribution of calanoid copepods in the plankton of Wisconsin Lakes
Byron Torke 351–365
Planktonic Copepoda of the Upper Paraná River Floodplain lakes (São Paulo and Mato
Grosso do Sul, Brazil)
Suzana Sendacz 367–374
The temporal distribution pattern of copepods in Corumbá Reservoir, State of Goiás,
Brazil
C.C.Bonecker,F.A.Lansac-Tôha,L.F.M.Velho,D.C.Rossa 375–384
The longitudinal distribution of copepods in Corumbá Reservoir, State of Goiás, Brazil
L.F.M. Velho, F.A. Lansac-Tôha, C.C. Bonecker, L.M. Bini, D.C. Rossa 385–391
Effects of food quality on growth and biochemical composition of a calanoid copepod,
Argyrodiaptomus furcatus, and its importance as a natural food source for larvae of two
tropical fishes
Lúcia Helena Sipaúba-Tavares, Maria Adriana Bachion, Francisco Manoel de Souza Braga 393–401
The influence of water and sediment properties on the occurrence of Potamocaris
Dussart, 1979 (Harpacticoida) in the upper Paraná River (Brazil)
Alice Michiyo Takeda, Carlos Eduardo, F. Rocha, José Cândido Stevaux 403–409
Symbiotic copepods
Annual changes in the population size of the salmon louse Lepeophtheirus sal-
monis (Copepoda: Caligidae) on high-seas Pacific Salmon (Oncorhynchus spp.), and
relationship to host abundance
Kazuya Nagasawa 411–416
Two ascidicolous copepods, Haplostomides otagoensis n. sp. and Botryllophilus cf.
banyulensis Brément, living in compound ascidians from Otago Harbor, NewZealand
Shigeko Ooishi 417–426
A new Species of Acontiophorus Brady, 1880 (Copepoda: Siphonostomatoida) from
Ushuaia, Argentina
R. Johnsson 427–430
Two new Artotrogids (Copepoda: Siphonostomatoida) from Madeira Island, Portugal
R. Johnsson 431–440
Systematics and biology
viii
Sexual dimorphism in calanoid copepods: morphology and function
Susumu Ohtsuka, Rony Huys 441–466
Taxonomy of Oncaeidae (Copepoda, Poecilostomatoida) from the Red Sea. III.Morpho-
logy and phylogenetic position ofOncaea subtilis Giesbrecht, 1892
Ruth Böttger-Schnack, Rony Huys 467–481
Longipedia corteziensis sp. nov. (Copepoda, Harpacticoida, Longipediidae) from a
coastal lagoon in northwestern Mexico, with the definition of thehelgolandica species-
group of the genusLongipedia Claus, 1863
Samuel Gómez 483–496
The naupliar stages ofCyclopina yutimaete Lotufo (Cyclopinidae, Cyclopoida)
Tagea K.S. Björnberg 497–501
Post-embryonic development ofPsammopsyllus maricaeCottarelli, Saporito & Puccetti,
1983(Copepoda,Harpacticoida)
Raffaella Berera, Vezio Cottarelli 503–512
Male ofMoraria radovnae Brancelj, 1988 (Copepoda: Crustacea), andnotes on endemic
and rare copepod species from Slovenia and neighbouring countries
Anton Brancelj 513–524
How many species ofDiacyclops? New taxonomic characters and species richness in a
freshwatercyclopid genus (Copepoda, Cyclopoida)
Fabio Stoch 525–531
A newDiacyclops (Copepoda, Cyclopoida, Cyclopidae) fromnorthwestern Argentina
Cecilia Locascio de Mitrovich, Silvina Menu-Marque 533–538
A new species of Notodiaptomus Kiefer, 1936 (Copepoda, Diaptomidae) fromthe
Amazon and Orinoco River Basins
Graciela Cicchino, Edinaldo N. Santos Silva, Barbara Robertson 539–548
Clarification of the taxonomic status of Notodiaptomus anisitsi (Daday, 1905) and re-
lated species, with description of a new species from Argentina(Crustacea: Copepoda:
Diaptomidae)
J.C. Paggi 549–564
Distributions of three Eodiaptomus species (Copepoda: Calanoida) in Thailand, with a
redescription ofE. draconisignivomi Brehm, 1952
La-orsri Sanoamuang 565–576
Hydrobiologia 453/454:ix–xiii, 2001.
R.M. Lopes, J.W. Reid & C.E.F. Rocha (eds), Copepoda: Developments in Ecology, Biology and Systematics. ix
Preface
This volume comprises the Proceedings of the Seventh International Conference on Copepoda,
held in Curitiba, Brazil, during July 1999, under the sponsorship of the Federal University of
Paraná and the World Association of Copepodologists (WAC).
Copepods are a major group of metazoans. They usually occupy a central place in aquatic
food webs, and are of direct importance to humans in their roles as food source for juvenile
fish, parasites of commercial fish and shellfish, and intermediate hosts of certain parasites and
pathogenic organisms.
A total of 228 specialists, including many graduate students, attended the Curitiba Confer-
ence (Fig. 1). There was a strong representation from the Southern Hemisphere, and particularly
Latin America. The main topics of the conference included the ecology, biology, and systemat-
ics of copepods. Special symposia were held on the following subjects (chairperson’s name in
parentheses): copepod development (Hans-Uwe Dahms), copepods in Antarctic and Subantarc-
tic ecosystems (Sigrid Schiel), copepods in challenging environments (Janet Reid and Carlos
Rocha), sexual dimorphism in copepods (Geoff Boxshall), and biogeography of copepods (Dov
Por). Pre- and post-conference workshops were held on Neotropical copepods (Carlos Rocha),
chromatin diminution (Grace Wyngaard), and benthic harpacticoid copepods (Dov Por). Of the
49 articles in this volume, 17 deal with the ecology of marine copepods, 16 treat freshwater and
semi-terrestrial copepods, and 5 report on the systematics and biology of parasitic copepods.
An additional 11 deal with taxonomy and systematics of free-living species, from both marine
and freshwater habitats.
Prof. Ju-shey Ho, the outgoing President of the WAC at the time of the Conference, presented
the traditional “Maxilliped Lecture,” which is reproduced here in full as the first article in this
volume. Prof. Ho offered a general view of the status of research on parasitic and associated
copepods, and strongly encouraged younger researchers to enter the field. Certainly, the scarcity
of young specialists in certain areas of investigation, such as parasitology and taxonomy, and in
some specific topics such as small pelagic copepods and ecophysiology of tropical species, was
a pervasive subcurrent and often discussed during the Conference.
Dr Tagea Björnberg, the “mother” of copepod studies in Brazil, was acknowledged in a
special presentation by Carlos Rocha (on behalf of the Local Organizing Committee) during the
Plenary Session. During nearly her entire career, Dr Björnberg has encouraged research on the
morphology, ecology, and behavior of nauplii, still a seriously neglected field.
The efficient organization, the comfortable venue at the Barigui Park Convention Center,
and the active cultural program provided ample opportunities for personal interactions and –
obviously – intensive discussions. As is traditional at copepod conferences, the organizers made
a serious effort to minimize the number of simultaneous oral sessions, and scheduled ample time
to study the many posters. Participants were thoroughly informed before, during, and after the
Conference by the well-considered and diverse website (http://www.ufpr.br/eventos/icoc).
x
The Local Organizing Committee consisted of Rubens Lopes, Carlos Rocha, Dov Por and
Monica Montú. Several colleagues and students from the Federal University of Paraná, and
particularly from the Center of Marine Studies, formed the competent “army” of local helpers.
Their assistance to the Local Organizing Committee both before and during the meeting was
instrumental for the success of the Conference. The collaboration of the International Organ-
izing Committee, composed by Geoffrey Boxshall (U.K.), Hans-Uwe Dahms (Germany), M.
Madhupratap (India), Anna Pasternak (Russia), Janet Reid (U.S.A.), Jefferson Turner (U.S.A.),
Shin-ichi Uye (Japan), and Hans Verheye (South Africa), was critical during the initial stages of
the organization. We thank the special session organizers, and also the session chairpersons who
conducted their work with efficiency and enthusiasm. The Conference received financial support
from the Brazilian funding agency CNPq (Conselho Nacional de Desenvolvimento Científico e
Tecnológico), the Federal University of Paraná, and the World Association of Copepodologists.
We also fully acknowledge the support given by the State University of Santa Cruz (Ilhéus,
Brazil) during the process of editing this volume.
All manuscripts submitted at the Conference were reviewed and edited following the Hydro-
biologia style. The articles are organized approximately according to the major themes of the
Conference. We are indebted to the many reviewers for their careful and supportive evaluations:
Andrade, C. F. S. (Brazil), Ashjian, C. (U.S.A.), Atkinson, A. (U.K.), Bamstedt, U. (Norway),
Bathmann, U. V. (Germany), Beaumont, K. (Australia), Benfield, M. (U.S.A.), Björnberg, T. K.
S. (Brazil), Bodin, P. (France), Boltovskoy, D. (Argentina), Bradford-Grieve, J. (New Zealand),
Brancelj, A. (Slovenia), Brandorff, G.O. (Germany), Bruno, M. C. (U.S.A.),
I. (Croatia), Buskey, E. (U.S.A.), Christou, E. (Greece), Cicchino, G. (Venezuela), Corkett,
C. (Canada), Coull, B. (U.S.A.), Dagg, M. (U.S.A.), Dahms, H.-U. (Germany), Dam, H.
G. (U.S.A.), Defaye, D. (France), DeMott, W. (U.S.A.), Dodson, S. (U.S.A.), Dutz, J. (The
Netherlands), Escribano, R. (Chile), Fernandez-Puelles, M. L. (Spain), Ferrari, F. D. (U.S.A.),
Fosshagen, A. (Norway), Fransz, G. (The Netherlands), Frisch, D. (Germany), Fussmann, G.
(U.S.A.), Galassi, D. (Italy), Gannon, J. E. (U.S.A.), Gilbert, J. (U.S.A.), Gomez, S. (Mexico),
Gotto, R. V. (U.K.), Greenwood, J. (Australia), Hagen, W. (Germany), Hardy, E. (Brazil),
Hart, R. (South Africa), Havel, J. (U.S.A.), Hirst, A. (U.K.), Ho, J.-s. (U.S.A.), Hogans, W.
E. (Canada), Holsinger, J. (U.S.A.), Hopcroft, R. (U.S.A.), Hwang, J.-S. (Taiwan), Ianora,
A. (Italy), Ishida, T. (Japan), Izawa, H. (Japan), Jaume, D. (Spain), Jersabek, C. (Austria),
Johnsson, R. (Brazil), Karanovic, T. (Italy), Kikuchi, Y. (Japan), Kimmerer, W. (U.S.A.), Klein
Breteler, W. (The Netherlands), Lakkis, S. (Lebanon), Luque, J. L. (Brazil), Maly, E. (Canada),
Marin, V. (Chile), Marten, G. G. (U.S.A.), Martinez Arbizu, P. (Brazil), Matsumura-Tundisi,
T. (Brazil), McKinnon, D. (Australia), Menu-Marque, S. (Argentina), Mielke, W. (Germany),
Montú, M. (Brazil), Morgan, C. (U.S.A.), Nierhoff, B. (Germany), Olafsson, E. (Sweden),
Omori, M. (Japan), Pace, M. (U.S.A.), Paffenhöfer, G.-A. (U.S.A.), Pagano, M. (France), Paggi,
J. C. (Argentina), Pasternak, A. (Russia), Pesce, G. (Italy), Peterson, W. T. (U.S.A.), Piaseki, W.
(Poland), Pinel-Alloul, B. (Canada), Pinto-Coelho, R. (Brazil), Por, F. D. (Israel), Ranga-Reddy,
Y. (India), Rayner, N. (South Africa), Roff, J. (Canada), Roman, M. (U.S.A.), Saiz, E. (Spain),
Santer, B. (Germany), Santos-Silva, E. N. (Brazil), Saunders III, J. (U.S.A.), Schnack-Schiel,
S. (Germany), Schram, T. A. (Norway), Schriever, G. (Germany), Sendacz, S. (Brazil), Siokou-
Frangou, I. (Greece), Sipauba Tavares, L. H. (Brazil), Smith, S. (U.S.A.), Snell, T. (U.S.A.),
Souissi, S. (France), Sprules, G. (U.S.A.), Stoch, F. (Italy), Strayer, D. (U.S.A.), Strickler, J.
