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Ecology Of Forest Insects PDF

615 Pages·1986·10.1 MB·English
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ECOLOGY OF FOREST INSECTS SERIES ENTOMOLOGICA EDITOR K. A. SPENCER VOLUME 26 ECOLOGY OF FOREST INSECTS ANDRZEJ SZUJECKI Department of Forest Protection and Ecology Agricultural University of Warsaw, Poland Translated from the Polish by ALBERT RADZIWTLL 1987 DR W. JUNK PUBLISHERS a member of the KLUWER ACADEMIC PUBLISHERS GROUP DORDRECHT/BOSTON/LANCASTER PWN -POLISH SCIENTIFIC PUBLISHERS WARSZAWA Distributors: for the United States and Canada Kluwer Boston Inc. 190 Old Derby Street Hingham, MA 02043 USA for Albania, Bulgaria, Cuba, Czechoslovakia, German Democratic Republic, Hun gary, Korean People's Democratic Republic, Mongolia, People's Republic of China, Poland, Rumania, the U.S.S.R., Vietnam and Yugoslavia Ars Polona Krakowskie Przedmiescie 7 00-068 Warszawa, Poland for all other countries Kluwer Academic Publishers Group Distribution Center P.O. Box 322 3300 AH Dordrecht The Netherlands First English Edition 1987 Translation from the Polish original Ekologia owadow ldnych published by Panstwowe Wydawnictwo Naukowe, Warszawa 1980 Library of Congress Cataloging in Publication Data Szujecki, Andrzej. Ecology of forest insects. (Series entomologica ; v. 26) Bibliography: p. I. Forest insects - Ecology. I. Title. II. Series. QL463.S96 1984 595.7'052642 83-17602 ISBN-13: 978-94-010-8630-1 e-ISBN-13: 978-94-009-4804-4 DOl: 10.1007/978-94-009-4804-4 Cover design: Zygmunt Ziemka Copyright © by PWN - Polish Scientific Publishers, Warszawa 1987 Softcover reprint of the hardcover I st edition 1987 All rights reserved. No part of this publication may be reproduced, stored in a re trieval system or transmitted in any form or by any means, electronic, mechanic, photocopying, recording or otherwise without prior permission of Polish Scientific Publishers. To my Professor, doctor honoris causa of the Agricultural University of Warsaw, MARIAN NUNBERG, on the 60th anniversary of His scientific work The Author Introduction Few branches of the biological sciences have developed to such an extent as has ecology in the recent decades. The successful development of this science is directly related to need to create a sound scientific basis by which we can control our diminishing natural resources and integrate this with the control of biological systems of the component species. Studies on the bioenergetics of ecosystems, as well as on the home ostatic mechanisms functioning at the populations and biocenoses levels are of great importance in this respect. The results of these studies are very significant in forestry management which deals with multiannual tree communities - forest stands. It is particularly important in this respect for economic planning, in view of the necessity for the possible prolonged maintenance of forest biocenoses as stable systems. Neither in the present nor in the immediate future can be protection of forest ecosystems consisting of natural plant and animal communities be achieved by the cultivation of resistant forms or the intensification of chemical intervention. Historical experience indicates that, besides the main factors deter mining the turnover of matter and energy flow in forest ecosystems, the mechanisms regulating the size of populations of various insect species are the most decisive element of stability of exploited forest associations securing also a favourable course of timber production, lasting forest exploitation and permanence of the forest as a component of the natural landscape. This is due to the fact that communities of forest insects, formed in the course of many centuries within the process of development of forest formations, are particularly susceptible to external stimuli and readily react to changing environmental conditions in the form of changes in their numbers. Under definite conditions many phytophagous spe cies, in dissipating their enormous reproductive potential, may rapidly reach such enormous numbers that their nutrient requirements exceed VII INTRODUCTION the efficiency of the given phytocenosis. This leads to a change in the rate and direction of the circulation of matter and energy and even to a collapse of the entire ecosystem. These circumstances seem to justify the need for a detailed under standing of the ecology of forest insects which should supply certain data for correct forest management, particularly in the field of forest protection and shaping of the environment. The ecology of forest insects as a scientific branch connected with forest ecology and entomology includes: 1) studies on the dependences of individual insect species on various factors of the forest environn:c nt, which frequently change under the in fluence of management practices, as well as the elucidation of the causes and regularities of occurrence of definite insects in the forest due to these dependences; 2) studies of the influence of the forest environment on the changes in the abundance of individual species and their contribution to the turnover of matter and energy flow in forest ecosystems; 3) studies on the conditions and regularities of the structure of multi specific communities of organisms, particularly insects in various forest types under different geographical conditions, and upon the application of various forest management methods as well as the recognition of the homeostatic mechanism of these communities. The influence on communities and functions of forest insects of anthropogenic activities not connected with forest management, as for example industrial pollution or agricultural utilization of forest soils, etc., is an important aspect of the study of the ecology of forest insects. The aim of these studies is to create an (cological basis for the plan ning and realization of protective methods as well as biological recultiva tion of areas requiring afforestation. A study of ecology of forest insects should establish the rules governing insect numbers and by forming the forest environment should raise the resistance of forest stands. Such a study should also indicate the natural and, in particular, the ento mocenotic consequences of forest management as well as the influence of other anthropogenic factors. VIII Contents Introduction . . . . . . . . . . . . . . . . . . . . . . . . VII 1. SPECIES AND POPULATIONS OF INSECTS AND T:-IEIR BASIC ASSOCIATIONS WITH THE FOREST ENVIRON- MENT ........................ . 2. INFLUENCE OF ABIOTIC ENVIRONMENTAL FACTORS ON FOREST INSECTS . . . . . . . . . . . . . 13 Influence of solar radiation and atmospheric factors. 13 2.1. Light 13 2.1.1. Influence of light on behaviour . . . . 13 2.1.2. Influence of light on reproduction and deveJop- ment ............ . 17 2.2. Temperature . . . . . . . . . . . . . 21 2.2.1. Regulation of body temperature . 22 2.2.2. Thermal tolerance . . . . . . . 26 2.2.3. Influence of temperature on activity and orienta- tion. . . . . . . . . . . . . . . . . . . . . 30 2.2.4. Influence of infra-red radiation on behaviour and orientation. . . . . . . . . . . . . . . . . . 32 2.2.5. Dependence of growth and development on tem- perature. . . . . . . . 33 2.3. Humidity and precipitation . . . 39 2.3.1. Water in the insect body. 39 2.3.2. Tolerance and adaptatio!l . 41 2.3.3. Activity, orientation and preference phenomena. 47 2.3.4. Influence of humidity on reproduction and devel- opment . . . 49 2.3.5. Precipitation. . . . . . . . . . . . . . . . . 51 IX CONTENTS 2.4. Air, atmospheric pressure and wind 52 2.5. Integrated action of climatic factors 54 2.5.1. Climograms, bioclimograms and hygrothermic coefficient . . . . . . . . . . . . . . . . . . 55 2.5.2. Influence of the forest mesoclimate on the distribu- tion of insects. . . . 59 Influence of soil factors on insects. 63 2.6. General characteristics of soil insects . 63 2.7. Insects and physical soil properties. . 64 2.7.1. Structure and mechanical composition 65 2.7.2. Humidity ..... 68 2.7.3. Thermal conditions. 69 2.7.4. Light .... . 73 2.7.5. Soil air ..... . 73 2.7.6. Humus content .. 74 2.8. Influence of chemical properties of the soil 76 3. THE ROLE OF NUTRIENT FACTORS IN THE LIFE OF FOREST INSECTS . . . . . . . . . . . . . . . 78 3.1. General characteristics of trophic relationships. 78 3.2. Nutrient relationships of phytophages 82 3.2.1. Nutrient specialization . . 82 3.2.2. Selection of the host plant . . 85 3.2.3. Food value . . . . . . . . . 91 3.2.4. Influence of food on reproduction and develop- ment . . . . . . . . . . . . . . . . . . . . 95 3.2.5. Influence of the physiological condition of the host plant . . . . . . . . . . . . . 97 3.3. Feeding relationships among zoophages . 100 3.3.1. Predators 102 3.3.2. Parasites 104 3.4. Characteristics of saprophages 110 4. INTRASPECIFIC RELATIONSHIPS OF FOREST INSECTS 112 Integration of populations . . . . . . . . 112 4.1. Sexual life and protection of progeny. 113 4.2. Collective life of insects. . . . . . . 118 x CONTENTS Structure and function of the population. 120 4.3. Abundance and density 120 4.4. Spatial distribution 122 4.5. Age distribution . . . 124 4.6. Sex structure. . . . . 127 4.7. Reproductive potential. 129 4.8. Mortality. . . . 131 Population dynamics. 141 4.9. Level of abundance and its variation. 141 4.10. Coefficient r and curves of population increase 144 4.10.1. Oscillations . . . . . . . . . . . . . 149 4.10.2. Fluctuations and outbreaks (gradations) 152 4.11. Phasic character and spread of outbreaks (gradations). 155 5. CAUSES AND COURSE OF CHANGES IN THE ABUN- DANCE OF INSECT POPULATIONS .......... 162 Development of opinions on the causes of changes in abundance. 163 5.1. Concepts not accepting the notion of automatic regulation of the abundance of populations. . . . , . . . . . . 163 5.1.1. Concept of the decisive importance of one factor. . 163 5.1.2. Concept of the complex influence of the components of the ecosystem. . . . . . . . . . . . . . . 172 5.2. Concepts based on the automatic regulation of popula- tion size . . . . . . . . . . . . . . . . . . . 180 5.2.1. Concepts of self-regulation with respect to the decisive role of parasites and of overcrowding. . 180 5.2.2. Concept of the automatic regulation of abundance. 181 5.2.3. Facts justifying the concept of the automatic regu lation of abundance . . . . . . . . . . . . . 185 5.2.4. Milne's integrating concept .......... 191 5.2.5. Concept based on the influence of genetic mecha- nisms . . . . . . . . . . . . . . . . .. 193 5.2.6. Concept of modifying and regulating factors 196 Factors determining the course of changes in abundance 202 5.3. Undirectionally acting (random) factors. . . . .. 202 5.3.1. Direct influence of meteorological conditions. 203 XI

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Few branches of the biological sciences have developed to such an extent as has ecology in the recent decades. The successful development of this science is directly related to need to create a sound scientific basis by which we can control our diminishing natural resources and integrate this with t
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