Proceedings in Life Sciences

Transcription

1 Proceedings in Life Sciences

2 The Role of Arthropods in Forest Ecosystems Edited by 'V. J. Mattson With Contributions by G. D. Amman J. T. Callahan B. W. Comaby K. Cromack, Jr. D. A. Crossley, Jr.. D. L. Dindal W. M. Fender R. Fogel B. Heinrich D. Inouye J. C. M. Jonkman J. F. McBrayer L. J. Metz P. A. Morrow P. SoIlins A. W. Todd R. L. Todd D. P. Webb With 28 Figures Springer-Verlag New York Heidelberg Berlin

3 WILLIAM J. MATTSON U.S. Department of Agriculture, Forest Service North Central Forest Experiment Station 1992 Folwell Avenue St. Paul, Minnesota 5510S/USA ISBN-13: e-isbn-13: : / Library of Congress Cataloging in Publication Data. International Congress of Entomology, 15th, Washington, D.C., The role of arthropods in forest ecosystems. (Proceedings in life sciences). Includes index. 1. Forest insects-congresses. 2. Arthropoda-Congresses. 3. Forest ecology-congresses. I. Mattson, William J., II. Title. QLA '.2' This work is subject to copyright. All rights are reserved, whether the whole or part of the material is concerned, specifically those of translation, reprinting, re-use of illustrations, broadcasting, reproduction by photocopying machine or similar means, and storage in data banks. Under 54 of the German Copyright Law, where copies are made for other than private use, a fee is payable to the publisher, the amount of the fee to be determined by agreement with the publisher. by Springer-Verlag, New York, Inc Softcover reprint of the hardcover 1 st edition 1977 The use of registered names, trademarks, etc. in this publication does not imply, even in the absence of a specific statement, that such names are exempt from the relevant protective laws and regulations and therefore free for general use.

4 Preface The role of arthropods in forest ecosystems is poorly understood. Yet such knowledge may be critical in order to explain fully the fundamental forces that shape the structure and regulate the functioning of such ecosystems. There are numerous hypotheses about the roles of various arthropods, but few, if any, of these hypotheses have been rigorously tested. Some, however, have been repeated so often and so widely that they are now accepted by many as unequivocal fact. Nothing could be further from the truth. Forest arthropods which derive most of their sustenance from plants are usually specially adapted for feeding in one of three subsystems-the above-ground plant system, the soil-litter system, or the aquatic stream system. Plant-feeding arthropods in the soil-litter and stream systems are primarily saprophous although many consume significant amounts of microorganisms. Research on the role of arthropods in each of these three subsystems has historically been provincial. Until very recently there has been little effort to collate, assimilate, and synthesize the plethora of findings in even one of these systems-rnuch less all three. This Symposium (at the 15th International Congress of Entomology, Washington, D.C. August 19-27, 1976) was organized for the specific purpose of promoting scientific synthesis. It fulfills one of the first requirements in such endeavors; namely, the juxtapositioning of current knowledge and hypotheses so that similarities can be perceived, insights can be derived, and more elaborate conceptual constructs can be built. These Proceedings have two Divisions-terrestrial phytophagous arthropods and terrestrial saprophagous arthropods. The Division on aquatic arthropods, which was given at the 15th International Congress of Entomology, is unfortunately not included here. The papers present a variety of perspectives on the roles of a variety of arthropods. However, there are some dominant recurring themes-one is the significant impact of arthropods on quantitative and qualitative aspects of system structure. For example: bark beetles affect the frequency distribution of tree size classes and plant succession; phyllophagous and sucking insects can differentially affect the competitive ability of coexisting eucalypt trees, bumblebees and other pollinating organisms can influence plant species composition as well as the evolution of certain qualitative characteristics of plants and plant communities, ants build nests which become ideal sites for the colonization and establishment of certain plants, soil arthropods alter the physicochemical structure of the soil-litter milieu through pelletizing plant

5 vi debris and converting some refractory substances (often with the aid of microorganisms) into more mobile, elementary forms, soil Collembola are not merely random assemblages of species but interact with one another forming definite associations. Another obvious theme which reinforces and is a corollary of the first is the sensitivity and responsiveness of arthropods to system structure-broadly speaking, cover and food. For example, it seems that organisms have evolved an "energy fit" to their normal hosts in their usual environment. This fit evolves in relation to the concentration and distribution of energy, nutrients and plant defenses through space and time. Environmental "patchiness" describes such variation. Although the three subsystems are different in many ways, they do have some similar structural features to which arthropod consumers are sensitive. In all three systems it is apparent that there is (1) a scarcity of Nand perhaps other elements like P and Na: (2) an abundance of nutritionally impoverished, nearly indigestible substrates like celluloses and lignins: (3) a pervasive presence of secondary plant compounds that are repellent, deterrent or toxic: and (4) an abundance of microorganisms which are intimately associated with arthropod microhabitats, foods, and bodies. An organism's life history strategy may vary in relation to these as well as other components of system structure. For example, where food substrates are nutritionally pauperate, many consumers rely heavily on microorganisms (as gut symbionts or as food) to enable them to gain adequate nutrition. Secondary compounds usually slow utilization rates of plant tissue by inhibiting microorganisms and arthropod consumers. Arthropods either avoid the compounds by attacking tissues before or after significant concentrations occur or evolve enzymes or other mechanisms to cope with such substances. The evidence suggests that arthropods, along with microorganisms, significantly interact with system structure; that is, they affect and are affected by it. It is a continuous, dynamic feed-back loop with rates of change usually subtle, but occasionally rapid and sweeping. St. Paul, July 1977 w. J. MATTSON

6 Contents Terrestrial Phytophagous Arthropods 1. The Role of the Mountain Pine Beetle in Lodgepole Pine Ecosystems: Impact on Succession (With 11 Figures) G. D. AMMAN 3 2. The Significance of Phytophagous Insects in the EucaZyptus Forests of Australia P. A. MORROW Resource Utilization by Colonial Lepidoptera Defoliators J. T. CALLAHAN Species Structure of Bumblebee Communities in North America and Europe (With 2 Figures) D. INOUYE Pollination Energetics: An Ecosystem Approach B. HEINRICH 41 Terrestrial Saprophagous Arthropods 6. The Roles of Terrestrial Saprophagous Arthropods in Forest Soils: Current Status of Concepts (With 1 Figure) D. A. CROSSLEY, JR Regulation of Deciduous Forest Litter Decomposition by Soil Arthropod Feces (With 6 Figures) D. P. WEBB Contributions of Cryptozoa to Forest Nutrient Cycles J. F. McBRAYER. 9. Soil Microorganism-Arthropod Interactions: Fungi as Major Calcium and Sodium Sources (With 1 Figure) K. CROMACK, JR., P. SOLLINS, R. L. TODD, D. A. CROSSLEY, JR., W. M. FENDER, R. FOGEL, and A. W. TODD 10. Ant Nests as Accelerators of Succession in Paraguayan Pastures (With 1 Figure) J. C. M. JONKMAN. 11. Community Structure of Collembola Affected by Fire Frequency (With 6 Figures) D. L. DINDAL and L. J. METZ

7 viii 12. Saprophagous Organisms and Problems in Applied Resource Partitioning B. W. CORNABY 96 Subject Index

8 List of Contributors AMMAN, G. D., Intermountain Forest and Range Experiment Station, Forest Service, U.S.D.A. Ogden, UT 84401, USA CALLAHAN, J. T., National Science Foundation, Washington, DC 20550, USA CORNABY, B. W., Battelle Columbus Laboratories, Columbus, OH 43201, USA CROMACK, K., JR., Department of Forest Science, Oregon State University, Corvallis, OR 97331, USA CROSSLEY, D. A., JR., Department of Entomology, Institute of Ecology, University of Georgia, Athens, GA 30602, USA DINDAL, D. L., Department of Forest Zoology SUNY, College of Environmental Science and Forestry, Syracuse, NY 13210, USA FENDER, W. M., Department of Entomology, Oregon State University, Corvallis, OR 97331, USA FOGEL, R., Department of Forest Science, Oregon State University, Corvallis, OR 97331, USA HEINRICH, B., Department of Entomological Sciences, University of California, Berkeley, CA 94720, USA INOUYE, D., Department of Zoology, University of Maryland, College Park, MD 20740, USA JONKMAN, J. C. M., Royal Netherlands Embassy, Scientific Office, 4200 Linnean Avenue NW, Washington, DC 20008, USA McBRAYER, J. F., Environmental Sciences Division, Building 2024, Oak Ridge National Lab., P. O. Box X, Oak Ridge, TN 37830, USA METZ, L. J., Southeastern Forest Experiment Station, Forest Service, U.S.D.A., Research Triangle Park, NC 27709, USA MORROW, P. A., Department of Ecology and Behavioral Biology, University of Minnesota, Minneapolis, MN 55455, USA SOLLINS, P., Department of Forest Science, Oregon State University, Corvallis, OR 97331, USA

9 x TODD, A. W., Department of Botany and Plant Pathology, Oregon State University, Corvallis, OR 97331, USA TODD, R. L., Institute of Ecology, Department of Agronomy, University of Georgia, Athens, GA USA WEBB, D. P., Department of Entomology, Texas A & M University, College Station, TX 77843, USA