ECOLOGY OF FOREST INSECTS

Transcription

1 ECOLOGY OF FOREST INSECTS

2 SERIES ENTOMOLOGICA EDITOR K. A. SPENCER VOLUME 26

3 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

4 Distributors: for the United States and Canada Kluwer Boston Inc. 190 Old Derby Street Hingham, MA USA for Albania, Bulgaria, Cuba, Czechoslovakia, German Democratic Republic, Hungary, 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 Warszawa, Poland for all other countries Kluwer Academic Publishers Group Distribution Center P.O. Box 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.S ' ISBN-13: e-isbn-13: DOl: / 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 retrieval system or transmitted in any form or by any means, electronic, mechanic, photocopying, recording or otherwise without prior permission of Polish Scientific Publishers.

5 To my Professor, doctor honoris causa of the Agricultural University of Warsaw, MARIAN NUNBERG, on the 60th anniversary of His scientific work The Author

6 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 homeostatic 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 determining 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 species, in dissipating their enormous reproductive potential, may rapidly reach such enormous numbers that their nutrient requirements exceed VII

7 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 understanding 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 influence 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 multispecific 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 planning and realization of protective methods as well as biological recultivation 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 entomocenotic consequences of forest management as well as the influence of other anthropogenic factors. VIII

8 Contents Introduction VII 1. SPECIES AND POPULATIONS OF INSECTS AND T:-IEIR BASIC ASSOCIATIONS WITH THE FOREST ENVIRON- MENT INFLUENCE OF ABIOTIC ENVIRONMENTAL FACTORS ON FOREST INSECTS Influence of solar radiation and atmospheric factors Light Influence of light on behaviour.... Influence of light on reproduction and devejopment Temperature Regulation of body temperature Thermal tolerance Influence of temperature on activity and orientation Influence of infra-red radiation on behaviour and orientation Dependence of growth and development on temperature Humidity and precipitation Water in the insect body Tolerance and adaptatio!l Activity, orientation and preference phenomena Influence of humidity on reproduction and development Precipitation IX

9 CONTENTS Air, atmospheric pressure and wind Integrated action of climatic factors Climograms, bioclimograms and hygrothermic coefficient Influence of the forest mesoclimate on the distribution of insects Influence of soil factors on insects General characteristics of soil insects Insects and physical soil properties Structure and mechanical composition Humidity Thermal conditions Light Soil air Humus content Influence of chemical properties of the soil THE ROLE OF NUTRIENT FACTORS IN THE LIFE OF FOREST INSECTS General characteristics of trophic relationships Nutrient relationships of phytophages Nutrient specialization Selection of the host plant Food value Influence of food on reproduction and develop- ment Influence of the physiological condition of the host plant Feeding relationships among zoophages Predators Parasites 3.4. Characteristics of saprophages INTRASPECIFIC RELATIONSHIPS OF FOREST INSECTS 112 x Integration of populations Sexual life and protection of progeny Collective life of insects

10 CONTENTS Structure and function of the population Abundance and density 4.4. Spatial distribution 4.5. Age distribution Sex structure Reproductive potential Mortality Population dynamics Level of abundance and its variation Coefficient r and curves of population increase Oscillations Fluctuations and outbreaks (gradations) Phasic character and spread of outbreaks (gradations) CAUSES AND COURSE OF CHANGES IN THE ABUN- DANCE OF INSECT POPULATIONS Development of opinions on the causes of changes in abundance Concepts not accepting the notion of automatic regulation of the abundance of populations...., Concept of the decisive importance of one factor Concept of the complex influence of the components of the ecosystem Concepts based on the automatic regulation of population size Concepts of self-regulation with respect to the decisive role of parasites and of overcrowding Concept of the automatic regulation of abundance Facts justifying the concept of the automatic regulation of abundance Milne's integrating concept Concept based on the influence of genetic mechanisms Concept of modifying and regulating factors 196 Factors determining the course of changes in abundance Undirectionally acting (random) factors Direct influence of meteorological conditions. 203 XI

11 CONTENTS Indirect influence of meteorological conditions Cyclic character of culmination of outbreak phenomena Factors acting (regulating) according to the rule of feedback Types of mechanisms regulating abundance Intraspecific regulating mechanisms Biocenotic regulating mechanism:; Mechanism of genetic feedback INSECTS IN FOREST BIOCENOSES XII Entomocenoses, associations and communities of insects Descriptive characteristic:; of r..s30ciations and communities Characteristics of quantitative proportions Characteristics of the structure of associations and communities Methods of comparison of cenos;:s and communities. 226 Insects in the spatial and time structure of forest biocenoses Spatial differentiation Insect communities in various fore:>t habitat type Insects in the layers structui"0 of the forest Differentiation in time Diurnal rhythm of activity Seasonal rhythm of activity Changes in insect communities occurring with the growth of forest stands Insects in the functioning of forest biocenoses Biotic associations of forest inse:::ts Contribution of insects to the flow of energy in the ecosystem Energy budget of insects The role of phytophagous insects in the productivity of forest ecosystems The role of saprophagous soil insects Succession of insect associations and communities 309

12 CONTENTS 7. CHARACTERISTICS OF THE FOREST ENTOMOFAUNA OF POLAND History of the forest entomofauna of Poland Problems of regionalization Characteristics of the entomofauna of Polish forests HUMAN INFLUENCE ON FOREST ENTOMOCENOSES. 433 General regularities of the processes of anthropization of the forest entomofauna Influence of antropogenous factors not associated with forest management on the entomocenosis Changes in soil water conditions Destruction of the forest herb layer and litter Forest fires Industrial air pollution and its accumulation in the soil Mining Influence of forest management on the entomofauna Specific composition of the forest stand Cutting systems and improvement fellings Mechanical soil cultivation Fertilization and lupin manuring of forest soils PROBLEMS OF THE DEVELOPMENT OF ENTOMOCE- NOSES UNDER REFORESTATION CONDITIONS Waste-heaps and dumping grounds Areas poisoned by industrial air pollution 9.3. Dune areas Afforested old farmland.. References Index of latin names of Invertebrata Su bject index XIII