Sensory Ecology Review and Perspectives NATO ADVANCED STUDY INSTITUTES SERIES A series of edited volumes comprising multifaceted studies of contemporary scientific issues by some of the best scientific minds in the world, as- sembled in cooperation with NATO Scientific Mfairs Division. Series A: life Sciences Recent Volumes in this Series Volume 11 - Surface Membrane Receptors: Interface Between Cells and Their Environment edited by Ralph A. Bradshaw, William A. Frazier, Ronald C. Merrell, David I. Gottlieb, and Ruth A. Hogue-Angeletti Volume 12 - Nucleic Acids and Protein Synthesis in Plants edited by L. Bogorad and J. H. Well Volume 13 - Prostaglandins and Thromboxanes edited by F. Berti, B. Samuelsson, and G. P. Velo Volume 14 - Major Patterns in Vertebrate Evolution edited by Max K. Hecht, Peter C. Goody, and Bessie M. Hecht Volume 15 - The Lipoprotein Molecule edited by Hubert Peeters Volume 16 - Amino Acids as Chemical Transmitters edited by Frode Fonnum Volume 17 - DNA Synthesis: Present and Future edited by Ian Molineux and Masamichi Kohiyama Volume 18 - Sensory Ecology: Review and Perspectives edited by M. A. Ali Volume 19 - Animal Learning: Survey and Analysis M. E. Bitterman, V. M. LoLordo, J. B. Overmier, and M. E. Rashotte Volume 20 - Antiviral Mechanisms in the Control of Neoplasia edited by P. Chandra The series is published by an international board of publishers in con- junction with NATO Scientific Affairs Division A Life Sciences Plenum Publishing Corporation B Physics New York and London C Mathematical and D. Reidel Publishing Company Physical Sciences Dordrecht and Boston 0 Behavioral and Sijthoff International Publishing Company Social Sciences Leiden E Applied Sciences Noordhoff International Publishing Leiden Sensory Ecology Review and Perspectives Edited by M.A.Ali Departement de Biologie Universite de Montreal Montreal, Canada PLENUM PRESS • NEW YORK AND LONDON Published in cooperation with NATO Scientific Affairs Division Library of Congress Cataloging in Publication Data NATO Advanced Study Institute on Perspectives in Sensory Ecology, Bishop's University, 1977. Sensory ecology. NATO advanced study institutes series: Series A, Life sciences; v. 18) "Lectures presented at the 1977 NATO Advanced Study Institute on Perspectives in Sensory Ecology, held at Bishop's University, Lennoxville, Quebec, Canada, July 10-22, 1977, co-sponsored by the Universite de Montreal and the National Research Council of Canada." "Published in cooperation with NATO Scientific Affairs Division." Includes index. 1. Animal ecology-Congresses. 2. Senses and sensation-Congresses. I. Ali, Mohamed Ather, 1932- II. Universite de Montreal. III. National Research Council of Canada. IV. North Atlantic Treaty Organization. Division of Scientific Affairs. V. Title. VI. Series. QH540.N17 1977 591.5 78-17597 ISBN 978-1-4684-3365-4 ISBN 978-1-4684-3363-0 (eBook) DOI 10.1007/978-1-4684-3363-0 Lectures presented at the 1977 NATO Advanced Study Institute on Perspectives in Sensory Ecology held at Bishop's University, Lennoxville, Quebec, Canada, July 10-22, 1977 Co-sponsored by the Universite de Montreat and the National Research Council of Canada Director: M. A. Ali, Montreal Advisory Committee: H.-J. Autrum, Munich P. Couillard, Montreal F. E. J. Fry, Toronto W. R. A. Muntz, Stirling © 1978 Plenum Press, New York Softcover reprint of the hardcover 1st edition 1978 A Division of Plenum Publishing Corporation 227 West 17th Street, New York, N.Y. 10011 All rights reserved No part of this book may be reproduced, stored in a retrieval system, or transmitted, in any form or by any means, electronic, mechanical, photocopying, microfilming, recording, or otherwise, without written permission from the Publisher Preface Never so pleased, sir. 'Twas an excellent dance, And for a preface, I never heard a better. Two Noble Kinsmen, Act III, Sc.S This volume is based mostly on the lectures delivered at an Advanced Study Institute (ASI) of the same title held in July 1977. One lecture given is not in the volume and three chapters, although not based on lectures delivered, have been added to better balance the book. A chapter on the ecosensory functions in crustaceans could not be put in due to time contingency. This absence is deeply regretted. The idea to hold an ASI on Sensory Ecology evolved slowly, main- ly due to my own research interest in the past and partly to the discussions I had with a number of colleagues, particularly Dr. John Lythgoe of the University of Sussex. The purpose was to interface Sensory Physiology with Ecology so that workers in those fields will develop a greater awareness for each other. Sense organs have of course evolved to keep their possessors.~ware of the environment and changes in it. Thus, normally one could expect that a study of their functions will be undertaken in relation to environmental parameters. But this has not been the case. Many if not most studies of sense organs have been undertaken either to study their structure or, some aspect of their function in a restricted way. Ecology, of course, deals with the interrelationship between the organism and its environment. This interrelationship obviously cannot exist without some kind of sensory input. However, one sees very little awareness of the importance of sensory functions in Ecology. Even ethologists often tend to ignore the importance of the interrelationship between sensory modalities and environmental factors in moulding the organism's total reaction. Thus, since sensation of some sort is essential to the detection of environmental factors, particularly changes in them, it appears that an imaginative approach to the v PREFACE study of sensory functions with regard to ecological factors on the one hand, and the investigation of environmental parameters in relation of the sensory functions in various taxonomic groups on the other, will provide a great deal of interesting information. The interplay between the physico-chemical organism and the physico- chemical environment is a delicate one and leads to complex behavioural responses such as migrations, rhythms etc. These will have to be examined in relation to sensory functions and their interactions with environmental cues in the light of information obtained by modern methods in both sensory physiology and ecology. The ASI was structured with the volume in mind. It was divided into two parts, one dealing with ecosensory functions in the various animal groups and the other concerning the adaptive radiation of sensory functions across the animal kingdom. The animal groups, particularly the invertebrate ones, were arbitrarily drawn up on the basis of convenience, availability of lecturers/authors and time. In most cases, the lecturer~/authors were chosen and invited on the basis of suggestions made by the members of the advisory committee and some others. Being a NATO-ASI, we also had to make sure that the lecturers were drawn from as many different countries as possible. I asked the authors to prepare lectures of a general nature and encouraged them to be as provocative and speculative as they wanted. Many accepted to do so and indeed during the ASI many hours were spent in lively discussions. On the basis of these discussions, the general reaction of the participants and the content of the others' lectures, the lecturers prepared their final drafts after the ASI. Obviously, in a volume of this sort a certain amount of overlap or repetition are unavoidable, even desirable. I have tried to keep them to a minimum but may not have succeeded entirely since I am certainly not expert in all aspects that this volume deals with. In my capacity as the director of the ASI, I attended every lecture and seminar given and, as the editor I have had to read every chapter in this volume carefully. This has considerably broadened my knowledge of Sensory Ecology and I appreciate having had this opportunity and hope that the readers of this volume will find it worthwhile. I am grateful to Dr. T. Kester of the NATO-Scientific Affairs Division for the advice .and encouragement he gave, without which the ASI could never have been held. I thank Dr. Maurice L'Abbe, Vice-Rector for Research in my university and Professor Jean-Guy Pilon, Chairman of my department for their moral and material support. Financial help was also provided by the National Research Council of Canada. The authorities of Bishop's University cooperated in every possible way, making our stay enjoyable. I am especially thankful to Monsieur J.L. Gregoire, Vice-Principal and Mr. Ivan Saunders, PREFACE Director of Buildings and Grounds for all their help. I thank also the members of my advisory committee for the help they gave and the suggestions they made concerning the choice of lecturers. I particularly thank my colleague Pierre Couillard for the more than the expected role he played in the organisation of the ASI and especially arranging the excursion to Quebec city. I wish to note how greatly the assistance of my librarian Miss Margaret Pertwee in putting the volume together is appreciated. I thank Mademoiselle Marielle Chevrefils for typing most of the camera-ready manuscript. On behalf of the authors and myself I wish to acknowledge the permission given by various authors and publishers for the reproduction of figures. Last but not the least, my thanks to Steve Dyer, Production Editor at Plenum for his patience and for his efficiency in getting the volume out quickly. Montreal February, 1978 Contents 10 GENERAL INTRODUCTION M.A. Ali •••• 3 2. SURVEY OF ECOSENSORY FUNCTIONS Phylogenetic Survey of Sensory Functions 11 M.A. Ali, R.P. Croll and R. Jaeger Taxes in Unice11s, Especially Protozoa 31 P. Couillard Ecosensory Functions in Lower Invertebrates 55 E.A. Ferrero Coe1omate Invertebrates (Except Crustacea, Arachnida and Insecta) . . . . . . . . • . • • . . . . . . . . 91 M.A. Ali Ecosensory Functions in Insects (With Remarks on Arachida) • • • • • • • • • • • • • • 123 M. Goga1a Fishes: Vision in Dim Light and Surrogate Senses • • • •• 155 J.N. Lythgoe Ecological Niche Dimensions and Sensory Functions in Amphibians .. . • . • . . . • . . . . . • . • 169 R.G. Jaeger ix x CONTENTS Reptile Sensory Systems and the Electromagnetic Spec trum • . . • . . . . . • . . . . . . 197 W.R.A. Muntz Sensory Ecology of Birds • • • • • • • • • • • • • • • •. 217 R.A. Suthers Sensory Ecology of Mammals • • • • • • • • • . • • . • • • 253 R.A. Suthers 3. ADAPTIVE RADIATION OF SENSORY MODALITIES Functional Adaptations in Chemosensory Systems 291 R.A. Gleeson Adaptive Radiation of Mechanoreception • • . • • • • •• 319 H. Markl Sound Reception in Different Environments 345 A. Michelsen Baroreception • • • • • . • • • • • • • • • • • • • •• 375 J.H.S. Blaxter The Adaptive Radiation of Proprioceptors • • • • • • •• 411 W. Wales Peripheral Thermal Receptors • • • • • • • • • • • • •• 439 R. Loftus Photoreception • • • •• • • • • • • 467 M.A. Ali, M. Anctil, and L. Cervetto A Survey of Vertebrate Strategies for Vision in Air and Water • . • . . . . . . . . . . . . 503 J.G. Sivak Ecological Aspects of Electroreception • • • • • • • •• 521 H.O. Schwassmann 4. INDICES Author Index 537 Subject Index 569 1. GENERAL INTRODUCTION