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The Spatial Distribution of Wood-Nesting Ants in the Central

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The Spatial Distribution of Wood-Nesting Ants in the Central AN ABSTRACT OF THE THESIS OF Gary R. Nielsen for the degree of Doctor of Philosophy in Botany and Plant Pathology presented on March 6, 1986. Title: The Spatial Distribution of Wood-Nesting Ants in the Central Coast Range of Oregon / 4 Abstractapproved: Redacted for privacy Fred okickson Two coniferous forests in the central Coast Range of western Oregon were surveyed for nests of wood inhabiting ants.Nineteen species and 825 ant nests were found, corresponding to an average nest density of 0.079 nests/m2 (maximum 0.38/m2) and a mean species density of 0.026 species/m2 (maximum 0.08/m2). The spatial distribution of all species was random within the study areas In contrast, the nest distribution patterns of the six most common species and all ants combined were found to be highly clumped (contagious) due to high nest densities on a few favorable sites. Most ants achieved greatest nest densities on high imsdlatim, early successional plots such as clear- cuts. The nest abundances of 15 species were negatively correlated with tree canopy cover. However, Lasius pallitarsis and Leptothorax nevadensis had higher nest densities in woody debrison forested plots. Furthermore, the nest densities of all ants combined, and of nine individual species were greater in stumps than logs. Within stumps, the nests of all species combined, as well as Camponotus modoc, Tapinoma sessile, and Lasius pallitarsis were concentratedon the south sides of stumps.The bark, cambial zone, and wood of woody debris in all stages of decompositionwere exploited by ants for nest sites. Leptothorax nevadensis, Tapinoma sessile, and Aphaenogaster subterranea occupied bark significantly more often than other tissues. Lasius pallitarsis and Camponotus modoc occupied wood significantly more often than other tissues although large nests extended through all tissues. The Spatial Distribution of Wood-Nesting Ants in the Central Coast Range of Oregon by Gary R. Nielsen A THESIS submitted to Oregon State University in partial fulfillment of the requirements for the degree of Doctor of Philosophy Completed March 6, 1986 Commencement June 1986 Redacted for privacy r Professor of Botany and Plant Pathology in Charge of major Redacted for privacy Head ofdelD'aitinentof Botarii and Plant Pathology Redacted for privacy School dr Date thesis is presented March 6, 1986 Typed by Gary Nielsen for Gary R. Nielsen Aclawaledgements I would like to thank Dylan Bulesco, Rose McKeown, Kendrick Moho lt, and Bob Truitt for their help one or two daysa week during the summers of 1982 and 1983. locating and sampling nests of these ants was very a difficult and labor-intensive task that could not have been accomplished in the ten odd months of sampling without these volunteers. The curiosity and interest of Annette Olson and Susan Holdeman is also gratefully acknowledged for it led them to assist me in the field on two separate occasions. I am indebted to Dr. George Ferguson for teaching me the basics of ant taxonomy and natural history. Many thanks to Steve Shattuck for his expert identification of the ants collected and for the natural history information he provided. Thanks to Brad Smith's vast knowledge of the Cornell Ecology Series Programs (of which Detrended Correspondence Analysis isa part) and patience with computer neophytes I was able to explore the patterns in the data and overcome the inevitable "bugs". A special thank you to Dave Swofford (Illinois Natural History Survey) who taught me CMS, showed me innumerable data manipulation and analysis shortcuts, and greatly facilitated my data analyses after I left Oregon. Mark Harmon introduced me to circular statistical analyses which made the analysis of my stump aspect data possible. Thanks to OSU's Forestry School for the use of McDonald Forest and to Marvin Rowley, manager of McDonald and Paul Dunn Forests, for his consultations on the history and management practices ofmy study area. I also gratefully acknowledge the U.S. Forest Service Alsea Ranger District for their permission to use the Corvallis Watershed site and for the management information they provided. I would like to thank Dr. Everett Hansen and Dr. Donald Zobel for loaning field and laboratory equipmenttome. I am indebted to my entire committee for their support in what started out to be a very interdisciplinary study. Dr. Peter McEvoy provided valuable insect ecology information; Dr. EverettHansen and Dr. Kermit Cromack helped metoexplore the implications of this study to forest management practices and forest ecology in general (especially wood pathology and the dynamics of wood decomposition); discussions with Dr. George Fergusonon the natural history, taxonomy, and physiology of ants were rewarding andvery much appreciated. I would like to thank my advisor, Dr. Fred Rickson, for givingme "free rein"to choosea project of my fancy, for his photographic assistance, and for his review ofmy dissertation. I also wish to express my appreciation to all my fellow graduate students and colleagues with whom I have had many rewarding discussions. I want to especially acknowledge the advice, support, and manuscript reviews of Joe Antos, Charles Halpern, Hiram Larew, Teresa Magee, and Annette Olson. Finally, I wish to thank my wife, Connie Rinaldo, for her help in all aspects of this project and without whose support, encouragement, and love this work would not have been possible. Table of Contents Introduction 1 Chapter I. The Spatial Distribution of Wood-Nesting Ants 3 in the Central Coast Range of Oregon. I. The Macrosite Introduction 4 Methods and Materials 6 Results 9 Discussion 38 Chapter II. The Spatial Distribution of Wood-Nesting Ants 46 in the Central Coast Range of Oregon. II. The Microsite Introduction 47 Methods and Materials 48 Results 51 Discussion 79 Chapter III. The Spatial Distribution of Wood-Nesting Ants 82a in the Central Coast Range of Oregon: General Discussion and Summary Ant Distribution and Forest Ecology 83 Spatial Distribution of Common Species 86 Nest Distribution of All Species 91 Ant Succession 92 Literature Cited 94 Appendix A Number of Queens in Incipient Colonies of Three 98 Carpenter Ant Species Appendix B Ant Nests in the Bark of Living Tree Boles 101 List of Tables Table Page 1.1 Frequency of occurrence of ant nests from both 11 study areas arranged in descending order of Abundance and overall mean nest and species densities. 1.2 Indices of dispersion and test of fit to the 12 negative binomial function for the eight most abundant species and all ant nests and species combined. 1.3 Numbers of ant nests in logs and stumps from 14 clearcut and forested areas of both study sites. 1.4 Correlation of plot scores on the first two 17 ordination axes with environmental variables for the combined study areas. 1.5 Correlation matrix for the combined study areas' 18 environmental variables. 1.6 Correlation matrix for site variables and 20 species of both study areas (McDonald Forest and Mary's Peak). 1.7 Correlation matrix for ant species of both study 21 areas (McDonald Forest and Mary's Peak). 1.8 Correlation of plot scores on first two ordin- 25 ation axes with environmental variables for McDonald Forest. I.9 Correlation matrix for McDonald Forest site 27 variables. 1.10 Correlation matrix for McDonald Forest site 29 variables and ant species. 1.11 Correlation matrix for McDonald Forest ants. 30 1.12 Frequency of occurrence, mean nest and species 33 density of ants of the Mary's Peak site and comparison of forest and clearcut habitats. 1.13 Frequency of occurrence, mean nest and species 34 density of ants of the McDonald Forest site. 1.14 Correlation of plot scores on the first two 36 ordination axes with environmental variables for Mary's Peak. Table Page 1.15 Correlation matrix for Mary's Peak site 37 variables. 1.16 Correlation matrix for Mary's Peak site 40 variables and ant species. 1.17 COrrelation matrix for Mary's Peak ants. 41 II.1 Mean nest density (standard deviation) of all 52 ants in woody debris from forest and clearcut habitats. 11.2 Mean nest density in woody debris from forest 53 and clearcut habitats (standard deviation). 11.3 Mean nest density of ants in stumps and logs 55 from all habitats (standard deviation). 11.4 Correlation of ordination scores for McDonald 56 Forest logs with log variables. 11.5 Correlation matrix for McDonald Forest log 59 microsite variables. 11.6 Mean nest densities (standard deviation) in 61 McDonald Forest logs from clearcut and forest habitats. 11.7 Correlation of Mary's Peak log ordination scores 63 with log variables. 11.8 Correlation matrix for Mary's Peak log microsite 64 variables. 11.9 Mean nest densities (standard deviation) in 67 Mary's Peak logs from clearcut and forest habitats. II. 10 Correlation matrix for McDonald Forest stump 70 microsite variables. II.11 Correlation matrix for Mary's Peak stump 73 microsite variables. 11.12 Comparison of numbers of nests indifferent 76 regions of woody debris. 11.13 Comparison of moisture content of woody debris 78 with (species name) and without (habitat name) ants. B. 1 Frequency of occurrence and mean nest density (in 102 parentheses) of ants found in live trees of both study areas. List of Figures Figure Page I.1 Detrended correspondence analysis ordination of 15 combined study area samples. 1.2 Detrended correspondence analysis ordination for 23 the ant species of the combined study areas. 1.3 Detrended correspondence analysis ordination of 24 McDonald Forest samples. 1.4 Detrended correspondence analysis ordination of 28 McDonald Forest species. 1.5 Detrended correspondence analysis ordination of 32 Mary's Peak samples. 1.6 Detrended correspondence analysis ordination of 39 Mary's Peak species. II.1 Detrended correspondence analysis ordination of 58 McDonald Forest log samples.
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