Advances in Marine Biology, Volume 16

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  • Advances in

    MARINE BIOLOGY VOLUME 16

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  • Advances in MARINE

    BIOLOGY VOLUME 16

    Edited by

    SIR FREDERICK S. RUSSELL Plymouth, England

    and

    SIR MAURICE YONGE Edinburgh, Scotland

    Academic Press London New York San Francisco 1979 A Sulsidiay of Harcourt Brace Jovanovich, Publishers

  • ACADEMIC PRESS INC. (LONDON) LTD.

    24-28 OVAL ROAD

    LONDON NW1 7DX

    U.S. Edition published by ACADEMIC PRESS INC.

    111 FIFTH AVENUE

    NEW YORK, NEW YORK 10003

    Copyright 0 1979 by Aca.demic Press Inc. (London) Ltd.

    All rights reserved

    NO PART OF THIS BOOK MAY BE REPRODUCED IN ANY FORM BY PHOTOSTAT,

    MICROFILM, OR ANY OTHER MEANS, WITHOUT WRITTEN PERMISSION FROM

    THE PUBLISHERS

    Library of Congress Catalog Card Number: 63-14040 ISBN: 0-12-02611&2

    PRINTED IN QEUCAT BRITAIN BY TEE WHITEPRIARS PRESS LTD. LONDON AND TONBRIDQE

  • CONTRIBUTORS TO VOLUME 16

    R. V. GOTTO, Department of Zoology, Queens University, Belfast, United Kingdum.

    ROGER P. HARRIS, Plymouth Laboratory of the Marine Biological Association of the United Kingdom, Plymouth, England.

    G. Y. KENNEDY, The University of Shefield, England.

    GUSTAV-ADOLF PAFFENHOFER, Skidaway Institute of Oceanogrqhy, Savannah, Georgia, U.S.A.

    A. J. UNDERWOOD, Department of Zoology, School of Biological Sciences, University of Sydney, N.S. W . 2006, Australia.

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  • CONTENTS

    COXTRIBWTORS TO VOLUME 16 . . . . . . . . . .

    The Association of Copepods with Marine lnverte brates

    R. V. GOTTO

    .. .. . . .. .. I. Introduction .. . . .. .. .. 11. Previous Reviews . . . .

    Considerations . . . . 111. Classification of Associated Forms and Other Systematic

    . . . . .. . . IV.

    V.

    VI.

    VII.

    VIII.

    IX.

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    XI.

    XII.

    XIII.

    General Studies of Single Species . . .. Anatomical and Functional Aspects . . ..

    A. Integument . . .. .. .. .. B. Sensory Structures . . . . . . .. C. FoodandFeeding . . . . . . . .

    E. Reproduction and Allied Topics . . .. Host Specificity . . .. . . . . ..

    D. Structural Studies of the Alimentary Canal

    Attraction to Host . . . . . . . . Preferential Host Niche . . . . . . . . Effect on Host and Host Reaction . . .. Morphological Variability at Infraspecific Level

    Sibling Speciation . . . . .. .. . . Population Studies . . .. .. .. Larval Studies . . . . . . .. . .

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  • viii OONTENTS

    XIV. Associated Harpacticoids and Calanoids . . A. Harpacticoids .. I . .. ..

    XV. Future Investigations . . .. .. . . .. . . . . B. Calanoids . . . .

    .. .. .. .. XVI. References . . .. XVII. Addenda . . .. .. . . .. ..

    The Ecology of intertidal Gastropods A. J. UNDERWOOD

    I. Introduction .. .. .. .. ..

    . . 84

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    .. 89 . . 90 .. 91

    .. 107

    .. 111

    11. Factors Affecting the Establishment of Patterns of Distribution . . 1 . .. .. .. .. 113

    A. Large-scale Patterns .. .. . . . . 113

    C. Summary and Conclusions . . .. . . 136 B. Local Patterns . . . . . . . . . . 120

    111. Ma,intenance of Patterns of Distribution by Behavioural Adaptations . . .. . . . . . . . . 137

    A. Patterns of Zonation .. .. .. .. 137 B. Dispersion within Zones: Homing Behaviour . . 142 C. Migrations and Aggregations . . .. .. 147 D. Summary and Conclusions . . . . .. 157

    IV. Maintenance of Patterns of Distribution by Physiological Stress . . .. .. .. . . .. . . 159

    A. Temperature and Desiccation . . . . . . 169 B. Salinity and Osmoregulation . . . . . . 164 C. Other Factors . . . . . . . . . . 166 D. Summary and Conclusions . . . . . . 168

    V. Competition and the Distribution and Abundance of Populations . . .. . . . . . . .. 170

    VI. Predation and the Distribution and Abundance of Populationa . . .. .. .. .. . . 179

    VII. Reproductit-e Biology and Geographical Distribution. . 183

  • CONTENTS

    VIII. Influences of Gastropods on the Structure of Intertidal ~

    Communities . . . . . . I . . . A. The Effects of Grazers on Sessile Animals B. The Effects of Grazers on Algae . . .. C. The Effects of Predators on Sessile Animals D. Summary and Conclusions . . ..

    X. Acknowledgements . . .. . . .. .. .. .. .. IX. Conclusions ..

    .. . . . . .. XI. References . . ..

    I.

    11.

    111.

    Laboratory Culture of Marine Holozooplankton and its Contribution

    t o Studies of Marine Planktonic Food Webs

    GUSTAV-ADOLF PAFFENROFER AND ROGER P. HARRIS

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    Introduction .. Cultivation Techniques

    A. Protozoa . . B. Cnidaria . . C. Ctenophora .. D. Rotifera . . E. Chaetognatha F. Mollusca . . G. Amphipoda .. H. Mysidacea . , I. Euphausiacea J. Ostracoda . . K. Decapoda . . L. Copepoda . . 13. Cladocera . . N. Tunicata . .

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    Contribution of Cultivation to the Study of Plankton . . . . .. Ecology . . .. . .

    A. Taxonomy and Morphology . . .. . . B. Experimental Studies Relating to Secondary

    Production .. . . C. Simulation Studies . , . . . .

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    IV . V .

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    CONTENTS

    Conclusions .. .. .. .. Acknowledgements .. .. .. References . . .. .. .. ..

    Pigments of Marine Invertebrates

    G . Y . KENNEDY Introduction I .

    Protozoa . . . . Porifera . . .. Coelenterata .. Ctenophora .. Platyhelminthes . . Nemathelrninthes . .

    A . Nematoda . . B . Acanthocephala

    Rotifera . . .. Nemertini .. . .

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    .. Annelida. Echiuroidea. Sipunculoidea. Priapuloidea and

    Phoronidea

    A . Crustacea B . Arachnida C . Myriapoda

    Arthropoda

    Mollusca . . Chaetognatha

    Brachiopoda

    Polyzoa . . Echinodermata

    Pogonophora

    Tunicata . .

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  • CONTENTS xi

    XIX. Comment . . . . .. .. .. .. .. 364 XX. Acknowledgements .. .. . . .. .. 366

    XXI. References . . .. . . .. .. .. .. 366 Taxonomic Index . . . . .. .. .. .. 383 Cumulative Index of Titles . . .. .. .. 423 Cumulative Index of Authors . . .. .. .. 425 Subject Index . . .. .. .. .. .. 399

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  • Adv. mar. BioZ., Vol. 16 1979 pp. 1-109.

    THE ASSOCIATION OF COPEPODS WITH MARINE INVERTEBRATES

    R. V. GOTTO Department of ZooJogy, Queen's University,

    Belfast, United Kingdom

    I. Introduotion . . .. .. .. .. .. .. .. .. 1 11. PreviousReviews .. .. .. .. .. .. .. .. 3

    111. Classification of Associated Forms and Other Systematio Considerations 3 IV. V.

    VI. VII.

    VIII. IX. X.

    XI. XII.

    XIII. XIV.

    xv. XVI.

    XVII.

    General Studies of Single Speoies .. . .. .. Anatomical and Functional Aspects . . ..

    A. Integument . . .. .. .. .. B. Sensory Struotures . . .. .. . . C. Food and Feeding . . .. .. ..

    E. Reproduction and Allied Topics . . .. Host Speoifioity . . .. .. .. .. Attrmtion to Host . . .. .. . . . . Preferentiml Host Niohe . . .. . . .. Effeot on Host and Host Reaction .. .. Morphological Variability at Infraspeoifio Level

    D. Struotural Studies of the Alimentary Canal

    Sibling Speoiation . . .. . . .. .. Population Studies . . .. .. . . .. Larval Studies .. .. .. . . .. Assooiated Harpacticoids and Calanoids . . ..

    A. Harpacticoids . . .. .. .. .. B. Calanoids .. .. .. .. ..

    Future Investigations. . .. .. .. .. Referenoes . . .. .. .. .. .. Addenda . , .. .. .. .. ..

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    I. INTRODUOTION

    In contrast to the fish parasites, those copepods which habitually partner marine invertebrates have received scant attention until fairly recent times. This discriminatory treatment is hardly surprising, for while the fish associates are frequently conspicuous, often bizarre and,

    1

  • 2 R. V. QOTTO

    above all, linked with hosts of economic importance, the latter are mainly unobtrusive forms, their ecology clandestine and their economic significance limited. It might indeed be fair to say that, up to the last half century, the handful of such species then known were regarded almost as aberrations on the part of an otherwise well-ordered Nature- a few curious types generated in odd moments of evolutionary whimsy.

    Although described and recorded as occasion offered by the carci- nologists of the nineteenth century, sustained study of these copepods attracted comparatively few workers. Thorell, Hesse, Claus, Gies- brecht, Canu and the Sars may be numbered among those who contributed more significantly to our knowledge in this earlier epoch. They were followed by such researchers as de Zulueta, working on the lamippids associated with alcyonarians, and Chatton who, in collabora- tion with Br6ment and Harant, concentrated on ascidicolous species. It is, however, only within the past thirty years that the immense variety of such copepods has become apparent, and, in particular, the full extent of their host spectrum realized. It is now, indeed, difficult to name a marine phylum some a t least of whose members do not harbour these versatile and little-known associates.

    At this point, we should perhaps define two terms rather more closely. The word " associate " was suggested by Gooding (1957) to describe those copepods which habitually partner other organisms, but whose precise ecological relationship with the host may be currently obscure. This neatly avoids the use of such vague terms as " semi- parasite " but does not prejudge future application of more rigidly defined categories when further information becomes available. On the host side, the term 'L invertebrate " is here taken to include not only the phyla universally recognized as such, but also certain of the " acraniate chordates "-in practice, ascidians, salps, enteropneusts and ptero- branchs.

    Since the present paper is a review rather than a monograph, some constraints in treatment at once become operative. There would seem little point, for example, in repeating lengthy morphological descriptions (which in fact represent the bulk of recent work) when these are readily available elsewhere. Again, certain aspects of structure, function and indeed general biology have remained virtually unexplored and thus unreviewable. Finally, hard information on many taxa of associated copepods is so sparse that it is difficult to treat such scanty material in an organized manner under appropriate headings. It is for this latter reason that I have referred symbiotic harpacticoids and calanoids to a single section, rather than including them with the more extensively studied associated cyclopoids.

  • THE ASSOCIATION OF COPEPODS WITH MARINE INVERTEBRATES 3

    11. PREVIOUS REVIEWS

    Only one, rather brief review of a wide ranging nature has been devoted largely to the copepod partners of marine invertebrates within recent years. Bocquet and Stock (1963) have discussed the inter- relationships between the major groups of parasitic copepods, showing that the Copepoda purasitica of earlier workers certainly constitutes no monophyletic unit, but should probably be apportioned between the poecilostomatous and the siphonostomatous cyclopoids. In like manner the old order, or sub-order, Notodelphyoida is a completely artificial assemblage, most of its members belonging by right to the gnathosto- matous cyclopoids while others should more properly be referred to the poecilostomes. As regards the latter, Bocquet and Stock have reviewed the old argument as to the presence or absence of a mandible in these copepods. They point out that there is general agreement amongst recent workers that the following mouth-parts can be attributed to all cyclopoids: mandible, maxillule (= Grst maxilla), maxilla (= second maxilla) and maxilliped-although any or all of these may be subject to reductions or specializations of varying degree. The same paper incorporates a discussion on the origin of parasitism and it5 specificity in copepods and concludes with some brief observations on sexuality, ecology, behaviour, development and larval cycles.

    The only other papers of note in this context are those of Bouligand (1966a), who has contributed a useful review of copepods found in association with coelenterates, and Cheng (1967), who has dealt with the copepod parasites and commensals of commercially important marine molluscs. To some extent, this last paper updates and amplifies the earlier work of Monod and Dollfus (1932a, b, 1934) on the copepods associated with molluscs in general.

    111. CLASSIF1:CATION OF ASSOCIATED FORMS AND OTHER SYSTEMATIC CONSIDERATIONS

    Some sort of taxonomic framework, however skeletal, must now be attempted. Recent discoveries of new families and genera, many of them clearly annectant, make this task somewhat easier than it would have been even a few years ago, but it nonetheless remains a daunting proposition.

    Let us take the broad view first. Although opinions still differ as to the circumscription of major divisions within the Copepoda, eight groupings are often cited as meriting recognition at ordinal or sub-

  • 4 R. V. QOTTO

    ordinal level : the Caligoida, Lernaeoida, Calanoida, Harpacticoida, Mons t rilloida , Not odelph yoida , Herp yllobioida and C y clopoida. Of these, the first two, as fish parasites, are largely outside the scope of this paper.* Such harpacticoids and calanoids as are known to be associated forms will be dealt with, as already mentioned, in a separate section. The monstrilloids are associated with other animals only in their larval instars, and the notodelphyoids, by general agreement, retain no further claim to ordinal or sub-ordinal status. Despite recent

    TABLE I. SIPIiONOSTOME CYCLOPOIDS AND THEIR HOSTS* _.

    Hosta f I . ;

    CYCLOPOIDA SIPHONOSTOWA

    -Calvocheridae Echinoids Stellicomi...

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