OTTO KINNE Editor Robert T. Paine MARINE ROCKY SHORES AND COMMUNITY ECOLOGY: AN EXPERIMENTALIST’S PERSPECTIVE Introduction (Otto Kinne) Robert T. Paine: A Laudatio (Tom Fenchel) O L O G C Y Publisher: Ecology Institute E Nordbünte 23, D-21385 Oldendorf/Luhe I N E Germany S T T I T U Robert T. Paine Department of Zoology NJ-15 University of Washington Seattle, Washington 98195, USA ISSN 0932-2205 Copyright © 1994, by Ecology Institute, D-21385 Oldendorf/Luhe, Germany All rights reserved No part of this book may be reproduced by any means, or transmitted, or translated without written permission of the publisher Printed in Germany Typesetting by Ecology Institute, Oldendorf Printing and bookbinding by Konrad Triltsch, Graphischer Betrieb, Würzburg Printed on acid-free paper Contents Introduction (Otto Kinne) .......................................... VII Robert T. Paine: Recipient of the Ecology Institute Prize 1989 in Marine Ecology. A Laudatio (Tom Fenchel) .......................... XIX Preface (Robert T. Paine)........................................... XXI I GENERAL CONSIDERATIONS ................................. 1 II HISTORICAL ORIGINS OF COMMUNITY ECOLOGY.............. 11 (1) Natural History/Observational Ecology ........................ 12 (2) Quantitative Methodologies ................................. 16 (3) Experimental Manipulation ................................. 19 (4) Mathematical Ecology ..................................... 28 (5) A Marine, Benthic Perspective on “Roots” ..................... 30 III COMPETITIVE INTERACTIONS ON HARD SURFACES ............ 37 (1) Kinds of Competition ...................................... 39 (2) General Statement of the Problem ............................ 40 (3) Competition for Space in the Presence of Consumers ............. 42 (4) Competition for Space when Predation has been Substantially Reduced 54 IV CONSTRUCTING COMMUNITIES FROM POPULATIONS .......... 77 (1) Models and Monocultures .................................. 77 Prediction in ecology ..................................... 79 Reference states and ecological experiments .................. 80 Competitively formed monocultures ......................... 81 (2) Unifying Dynamics in Intertidal Communities .................. 86 Disturbance ............................................ 88 Disturbance dynamics .................................. 90 Patch or gap traits ..................................... 91 Life history events in a patchy environment ................... 94 Patch size............................................ 95 Growth plasticity and population density ................... 96 Patch characteristics and individual performance............. 99 Source-sink populations ................................ 103 Patchily distributed individuals and metapopulations ......... 104 Dispersal ............................................ 106 Dispersal in marine environments...................... 107 Recruitment variation or supply-side ecology ............ 109 Trophic interactions ...................................... 114 Static or descriptive food webs ........................... 115 Dynamics and real food webs ............................ 117 The future of food web research .......................... 121 V RETROSPECTIVE ............................................ 123 (1) Human Exclusions from the Rocky Shoreline of Chile ............ 125 (2) Generalization of the Human Impact .......................... 131 (3) The Value of “Pristine” Environments ......................... 133 References ...................................................... 135 Introduction Otto Kinne Ecology Institute, Nordbünte 23, D-21385 Oldendorf/Luhe, Germany The book series “Excellence in Ecology” (EE) is published by the Interna- tional Ecology Institute (ECI) and made available at cost price.* EE books may also be donated to scientific libraries in Third-World countries. EE books are authored by recipients of the Ecology Institute Prize, which honors the sustained high performance of outstanding research ecologists. Prizes are awarded annually, in a rotating pattern, for the fields of marine, terrestrial and limnetic ecology. Prize winners are selected by a jury of seven ECI members appointed by the director. EE books offer the laureates the chance to publish their personal views on the state of the art of their fields of expertise and to bring to the attention of a world-wide audience their insights into the knowledge, problems and realities that form the biological basis for human existence. EE books address fellow scientists, teachers, students and decision makers who must translate ecological information into practicable rules and laws for the benefit of nature and mankind. The aims and activities of the Ecology Institute have been outlined in EE Book 3 (pp. VIII–IX). The ECI Prize carries a stipend of US $5000. In addition to this prize, the ECI awards an annual IRPE Prize (International Recognition of Profes- sional Excellence) which carries a stipend of US $750 and which honors a young (not more than 40 years of age) research ecologist who has pub- lished uniquely independent and/or challenging papers representing an important scientific breakthrough and/or who must work under particularly *Address orders for EE books to: *Ecology Institute, Nordbünte 23, D-21385 Oldendorf/Luhe, Germany Tel: (+49) (0) 4132 7127; Fax: (+49) (0) 4132 8883; E-mail: [email protected]. Payment may be made via credit card (American Express, Visa, Euro/ Mastercard; please give account number and expiration date). Standing orders for the whole series are accepted at 10% reduced prices. For book authors, titles and prices, consult pp. VIII–XI. VIII INTRODUCTION difficult conditions. The ECI also supports, via the Otto Kinne Foundation (OKF), promising postgraduates in environmental sciences in East European countries — especially in the fields of ecology, diseases of organisms, and climate research. The OKF aids postgraduates — without distinction of race, religion, nationality or sex — by providing financial assistance for profes- sional travel and/or scientific equipment. For details write to the ECI. Nominations for ECI and IRPE Prizes (accompanied by CV, list of publi- cations, and a statement why, in the opinion of the nominator, the nominee qualifies for the prize) are invited from research ecologists on a global scale. They should be sent to the chairperson of the respective ECI Jury, or, alter- natively, to the ECI’s director, who will then forward them to the chairper- son. Eligible are all ecologists engaged in scientific research (except the ECI’s director, the Jury’s chairperson, and previous Laureates; Jury mem- bers nominated will be replaced by other ECI members). The Jury selects prize winners using the nominations received as well as their own knowl- edge of top performers and their own professional judgement. Nominations for OKF Fellows, addressed to the ECI and accompanied by a letter of support as well as a documentation of the nominees’ performance, are invited from ECI members and members of the Editorial Staffs of the three international Inter-Research journals. ECI Prize Winners, Their Major Scientific Achievements and Their Books Tom Fenchel (Helsingør, Denmark), ECI Prize winner 1986 in marine ecology. Quotation of the Jury (Chairman: John Gray, Oslo, Norway) The Jury found Professor T. Fenchel’s contribution to ecological knowledge in a variety of research fields to be of the highest international class. In particular, the Jury cites his brilliant and uniquely important studies on the microbial loop which have opened up a fundamentally new research field. Professor Fenchel is, in addition, an excellent publicizer in his field of research with authorship of a number of standard works in marine ecology. Book 1: Ecology – Potentials and Limitations. (Published 1987; price DM 67 plus postage and handling) ECI PRIZE WINNERS IX Edward O. Wilson (Cambridge, MA, USA), ECI Prize winner 1987 in terrestrial ecology. Quotation of the Jury (Chairman: Sir Richard Southwood, Oxford, UK) Professor E. O. Wilson is distinguished for his many contributions to differ- ent aspects of ecology and evolutionary biology. His life-time love of Nature, a theme explored in his book “Biophilia”, has been particularized in his study of ants leading to major new insights on the evolution of castes and the operation of social systems. His seminal “Sociobiology”, derived from this work, has founded a new branch of science, between ecology and the social sciences. With the late Robert MacArthur he was the originator of the mod- ern theories of island biogeography that have contributed not only to the understanding of island biota, but to community and population ecology. Book 2: Success and Dominance in Ecosystems: The Case of the Social Insects. (Published 1990; price DM 49 plus postage and handling) Gene E. Likens (Millbrook, NY, USA), ECI Prize winner 1988 in limnetic ecology. Quotation of the Jury (Chairman: William D. Williams, Adelaide, Australia) Gene Likens is a distinguished limnologist who has made salient contribu- tions to many fields of limnology. In 1962 he initiated and developed (with F. H. Bormann) the Hubbard Brook Ecosystem Study in New Hampshire. Com- prehensive investigations in this study provided a model for ecological and biogeochemical studies worldwide. A major finding of the study was that rain and snow are highly acidic. “Acid rain” is now recognized as one of the major environmental hazards in North America, Europe and elsewhere.