Predators and prey in fishes Developments in environmental biology of fishes 2 Series Editor EUGENE K. BALON Predators and prey in fishes Proceedings of the 3rd biennial conference on the ethology and behavioral ecology of fishes, held at Normal, Illinois, U.S.A., May 19-22, 1981 Edited by DA VID L.G. NOAKES, DAVID G. LINDQUIST, GENE S. HELFMAN & JACK A. WARD Reprinted from EnI'ironmental biology of fishes 8 (3/4), 1983 with addition of more material from the conference Dr W. JUNK PUBLISHERS THE HAGUE - BOSTON - LONDON 1983 Distributors: for the United States and Canada Kluwer Boston, Inc. 190 Old Derby Street Hingham, MA 02043 U.S.A. for all other countries Kluwer Academic Publishers Group Distribution Center P.O. Box '322 3300 AH Dordrecht The Netherlands Library or Congress Cataloging in Publication Data Conference On the Ethology and Behavioral Ecology of Fishes (3rd : 1981 : Normal, Ill.) Predators and prey in fishes. (Developments in environmental biology of fishes ; 2) "Reprinted from Environmental biology of fishes 8 (3/4), 1983 with addition of more material from the conference." Includes bibliographies and index. 1. Fishes--Behavior--Congresses. 2. Fishes-- Ecology--Congresses. 3. Predation (Biology)--Congresses. I. Noakes, David L. G. II. Environmental biology of fishes. III. Title. IV. Series. QL639.3.C65 1981 597' .051 83-238 ISBN-13: 978-94-009-7298-8 e-ISBN-13:978-94-009-7296-4 DOl: 10.1007/978-94-009-7296-4 Copyright <l=:; 1983. Dr W. Junk Publishers, The Hague Softcover reprint of the hardcover 1s t edition 1983 All rights resen·ed. No part of this publication may be reproduced, stored in a retriel'al system, or transmitted in any form or by any means, mechanical, photocopying, recording, or otherll"ise, lI"ithow the prior lITillen permission of the publishers, Dr W. Junk Publishers, P.O. Box 13713. 2501 ES The Hague, The Netherlands. Predators and Prey in Fishes This, the second volume in the series Developments reflect this central theme, as this volume will show. in Environmental Biology of Fishes, combines two Studies of fishes not only hold promise in this special issues of the journal Environmental Biology regard, they are clearly at the forefront in many of Fishes resulting from the 3rd biennial conference cases. on the Ethology and Behavioral Ecology of Fishes. The 3rd biennial conference on the Ethology and While one of those special issues drew attention to Behavioral Ecology of Fishes was also the 2nd the symposium on Predator-Prey Interactions in North American meeting of the International As- Fishes, it may not be so readily apparent that sociation of Fish Ethologists. The conference was another symposium, Applied Aspects of Fish Be- held May 19-22, 1981 at the Illinois State U ni- havior, was also held during the conference. The versity campus in Normal, Illinois, the site of the papers published here came from those two sym- two previous conferences (1977 and 1979) and of posia, and from contributed papers presented dur- the 4th conference in May, 1983. The organiza- ing the plenary session of the conference. We have tional committee for the 3rd conference was Jack A. presented the volume with its title, as that is clearly Ward, Chairman (Biology, Illinois State University, the central theme of most contributions, whether or Normal, Illinois), Jeffrey R. Baylis (Zoology, Uni- not they happen to have been presented in the versity of Wisconsin, Madison, Wisconsin), Luther predation symposium. We would also like to em- Brown (Biology, George Mason University, Fair- phasize the general significance of predation in the fax, Virginia), J. Michael Fitzsimons (Zoology, life of fishes by this volume. Perhaps it will be Louisiana State University, Baton Rouge, Louis- worthwhile if this leads us somewhat beyond the iana), Gene S. Helfman (Zoology, University of usual bounds of considerations of predator-prey Georgia, Athens, Georgia), Mary G. Henry (Ani- interactions. We want to draw attention to the mal Ecology, Iowa State University, Ames, Iowa), examples presented here illustrating the diversity David L.G. Noakes (Zoology, University of Guelph, and variety among fishes, both as model systems for Guelph, Ontario), Benoni Seghers (Kananaskis those interested in purely theoretical issues and as Centre for Environmental Research, Seebe, Alberta), practical systems for those concerned with empiri- Chris L. Thoms (Biology, Mt. Mercy College, Cedar cal testing of their hypotheses. We also want to Rapids, Iowa), Richard L. Wyman (Biology, Hart- suggest that predation (whether as predator or wick College, Oneonta, New York). There were 129 prey) might reasonably be taken as a central theme registered participants, 69 contributed papers in a for many descriptions or analyses of behavior and plenary session, and three invited speakers, George ecology. Visual pigments, colors and color patterns, W. Barlow (University of California, Berkeley), morphology, physiology, and social systems can all Arthur A. Myrberg, Jr. (Rosenstiel School of 5 Atmospheric and Marine Sciences, University of Mark Matus, Mark Lamon, Mark Zoran, Carrie Miami) and Carl D. Hopkins (University of Minne- Stafford, Gary Ostrander, Darin Collins, Sherry sota, now at Cornell University). Nevius, Dona Milinkovich, Greg Yarnick, Sue We would like to express our special appreciation Schaus, Elaine McGuffin, Tom Bei!, Kevin Deets to Gene S. Helfman who organized the symposium and Vicki Miller. They organized and coordinated Behavioral Tactics of Predators and Prey, and to all projection of visual aids, catered every coffee Mary G. Henry who organized the symposium on break, as well as the evening wine and cheese party Applied Aspects of Fish Behavior. We also thank and the opening beer bust, they were receptionists, the College of Continuing Education at Illinois guides, and operators of a continual taxi service to State University for making all the physical ar- and from the local airport. We give them a special rangements for the conference. In particular, we vote of thanks and appreciation. give special thanks to Dean Edward Anderson, Those who contributed to the editorial and re" Olive Aldrich, Roger Herberts, Rita Kohn, Marie view processes of this special issue are acknowled- Malinauskas and Nancy Park, all staff members of ged in the list included at the end of this volume. the College. We also thank Howard Hetzel, Biology, Articles published in this volume with an Environ,., Illinois State University, who served as the Chair- mental Biology of Fishes bibliographic note above man for the Committee for Local Arrangements. their titles were also published in the special issue of A conference such as this would be impossible the journal. Articles without that bibliographic note without the untiring efforts of interested students. are published only in this volume of Developments We thank the following who were members of the in Environmental Biology of Fishes. Committee for Local Arrangements: Thorn Ardelt, 6 Behavioral Tactics of Predators and Prey Papers from a symposium held during the third biennial conference on the Ethology and Behavioral Ecology of fishes, Illinois State University, Normal, 19~22 May 1981 Special Co-editors: David L.G. Noakes, David G. Lindquist & Gene S. Helfman 7 Bf HAVIORAL TACTICS O~ PREDATORS & PRE1 £ 8 ·£ 01 f . '81 Collector's T-shirt produced by the Symposium organizer for each speaker only. By permission of Johnny Hart and Field Enter- prises, Inc. (left): drawing by Mankoff (t. 1981 The New Yorker Magazine, Inc. (right). 8 Preface The potential consequences of a predator-prey predators tend to do that increase or maximize prey interaction are probably more serious, especially capture and that prey tend to do to avoid being for the prey, than are the consequences of most captured? The second theme emphasized the eco- competitive or parasite-host interactions. For this logical approach to predator-prey interactions. reason, the adaptations and tactics that prey show What are the environmental constraints that in- to the foraging manoeuvers of their predators, and fluence the evolution of structures and behaviors the counteradaptations of their predators, are often involved in predation and its avoidance? How do pronounced and even spectacular. Predation may in these factors combine to produce particular ,fssem- fact be a critical determinant of both the behavior blages of predators and prey with common adapta- of individuals and of the functional and taxonomic tions? The 18 papers presented at the symposium, composition of fish assemblages. This possibility and the ones published in these proceedings, re- was a major factor leading to the organization of present the varied approaches that researchers have the Behavioral Tactics symposium. The obvious taken in addressing these questions. necessity of feeding, the wealth of information Several topics were common to many presenta- available on food and energy budgets of animals tions: four of these in particular deserve at least both in the laboratory and field, and the approach brief mention. The first topic involved the question of optimality arguments to foraging situations have of individual variation and its prevalence and pos- all been contributing factors leading to the con- sible importance in predator interactions. Treat- siderable emphasis on feeding behavior in fishes. ment of this topic went beyond the traditional We were interested in discussing the behaviors discussion of the inapplicability of the idea of involved in predator-prey interactions and in the 'species-typical behavior' or of the 'specialist-gener- evolution of predator foraging and prey escape alist' dichotomy. Implied was the possible adaptive- tactics. The likelihood that many potential contri- ness of plasticity in the behavioral repertoire of a butors and participants already planned to attend predator or prey: by switching among tactics one's the third biennial symposium on Ethology and prey or predators may have difficulty adjusting Behavioral Ecology of Fishes made the arranging of rapidly to such behavioral changes. A possible speakers and ensuring an interested audience all the consequence could be the 'fatal errors' that often easier. Our symposium was entitled Behavioral appear to characterize successful (from the preda- Tactics of Predators and Prey to attract researchers tor's viewpoint) predator-prey interactions. One who were working on the actual behaviors associ- question that arises is whether such individual ated with such interactions in fishes. variation is more characteristic of predators or We considered two general themes in our selec- prey. This topic involves the investigation of me- tion of speakers. These were intended to reflect the chanisms at the individual, population and com- major categories of interest of the persons that have munity levels and is conducive to observation and regularly attended the biennial symposia. The first manipulation in both the field and laboratory. theme concerned the ethological context of The second topic involved the subject of mecha- predator-prey interactions. Could we identify pat- nisms. Researchers are increasingly interested in not terns of tactics and make genaralizations about just the consequences or outcomes of behavioral attack and escape behavior? Are there things that interactions, but in the causes of those outcomes. David L.G. Noakes et al. (eds.), Predators and prey in lishes. 9 ([ 1983. Dr W. Junk Publishers. The Hague. While this is certainly not a new subject, one now conditions as a result of their small size, but also the hears the term mechanism more frequently in both relative simplicity of their behavioral repertoire behavioral and ecological circles. This increasing should simplify observation and recording proce- interest will likely prove to be an important objec- dures. tive around which to formulate investigations. The The fourth topic is almost a negative one, in that appreciation that behavior may itself serve as an it concerns a lack of emphasis. There was a conspi- explanatory mechanism for phenomena at the levels cuous absence of the use of sociobiological termi- of populations or communities already seems well nology and reasoning during the presentations and established. The focus now seems to be shifting to a in the resulting papers. What this might say about more detailed analysis of the behavior, in turn. the discipline of sociobiology or about those who The third topic of emphasis, particularly in some study the behavior and ecology is difficult to inter- of the talks, was the ontogenetic implications of pret. Perhaps it is an emphasis deserving more con- predation. Several speakers discussed predation by sideration, even if it is to be rejected, for those or on early life history stages of fishes, as well as the planning research on predator-prey interactions. responses of young individuals or their parents that These are only four of a number of related may reduce the impact of predation during those subjects covered by the various speakers and au- particularly vulnerable intervals. Some of this inte- thors. Their cooperation during the organization rest probably stems from observations of the repea- and presentation of the symposium, and during the ted, often ineffective feeding efforts of young preda- subsequent editorial processes is gratefully ac- tors. Some also results from the realization of the knowledged. We would also like to thank a number greater predatory impact on younger fish, and of individuals for their special contributions to the especially from the extensive body of literature symposium and this special issue by way of advice dealing with the concept of the critical period for and assistance in various ways; E.K. Balon, J .R. first feeding and the presumed problem of star- Baylis, L. Dill, W. Rowland, R. Stein, and J.A. vation in larvae and young juveniles. A researcher Ward and his many assistants and volunteers. Those intent on obtaining quantifiable observations of who contributed to this issue during the review and predation should consider studying schools of editorial processes are listed at the end of the young fish. Not only would they be more amenable volume. to detailed studies under controlled laboratory 10