ATTACK DYNAMICS, IMPACT AND BIOLOGY OF PISSODES TERMINALIS HOPPING, IN REGENERATING LODGEPOLE PINE STANDS L.E. Maclauchlan B. Sc., University of Victoria, 1980 M.P.M., Simon Fraser University, 1986 THESIS SUBMITTED IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE DEGREE OF DOCTOR OF PHILOSOPHY in the Department of Biological Sciences Lorraine E. Maclauchlan 1992 SIMON FRASER UNIVERSITY August, 1992 All right reserved. This work may not be reproduced in whole or in part, by photocopy or other means, without permissiofi of the author. APPROVAL Name: LORPtAHNE E. MIACLAUCHLAN Degree: Doctor of Philosophy Title of Thesis: ATTACK DYNAMICS, IMPACT AND BIOLOGY OF PIISSODES TERMIXALXS HOPPING IN REGENERATING STANDS OF LODGEPOLE PIPIE Examining Committee: Chair: Dr. N.1-I. Haunerland, Assistant Professor m~orden,Professor, Senior ~uperviso< Department of Biological Sciences, SFU Dr. B. Roitberg, Associate Professor, Departmat nf @io$gical Sciences, SFU Dr. R.k! Alfaro, Resea For ry Canada, Paci V'A t ia, B.C. Public &+niner br. E.A. ~arnero6~~rofes6r,. Department of Entomology, Penn State University, University Park, Pennsylvania External Examiner PARTIAL.COPYRIGHT LICENSE I hereby grant to Slmon Fraser Unlverslty the right to lend my thesis, proJect or extended essay'(the ?ltle of whlch Is shown below) to users of the Slmon Fraser Unlverslty ~lbrir~,and to make partial or single coples on1 y for such usors or In response to a request f tom tho library of any othor unlverslty, or other educational Instltutlon, on its own behalf or for, one of Its users. I further agree that permission for multlple copylng of thls work for scholarly purposes may be granted by me or the Doan of Graduate Studios. It is understood that,copylng or publlcatlon of thl s work for flnanclal galn shall not be allowed without my wrltten permlsslon. Tltle of Thesls/Project/Extended Essay Attack dynamics, impact and biology of Pissodes terminal i s Howina. in regenerating lodgepole pine stands Author: - ., (s lgnature) ~:Ix. 1992 (date) ABSTRACT The impact of Pissodes terminalis Hopping on immature lodgepole pine, Rnus contorta var. latifolia Engelm. is greatest in the dry, low elevation Interior Douglas-fir zone (IDF), where up to 31 and 17% of the annual height growth is lost in the first and second years after attack, respectively. In the Montane spruce (MS) zone 25 and 14% of annual height growth is lost in the same two years. An increase in the area potentially available (APA) to a tree can cause impact of the weevil in the MS zone to approach that in the IDF. The cool, high-elevation Engelmann spruce-subalpine fir zone has the lowest hazard. The natural distribution of lodgepole pine approaches an aggregated pattern, as described by negative values of the Clark-Evans-Donnelly statistic (CED), as does the spatial pattern of attack by P. terminalis, with CED values ranging from -7.127 (P < 0.001) in the lowest density, IDF plots, to -3.98 1 (I?< 0.00 1) in the highest density, MS plots. As density decreases in the IDF and MS zones, the probability of individual stems being attacked by P. terminalis increases, with one stand in the MS having a density of < 800 stems per ha sustaining > 70% of stems attacked. Increases in the APA lead to increasingly severe defects after attack. For 161 defects caused by P. terminalis, followed for three growing seasons, the Tau-b statistic was 0.602, indicating little change in defect category, although 11.8 % increased in severity and 16.8 % decreased in severity. Various phenomena were disclosed that could affect the impact of P. terminalis, including variations in the life cycle between biogeoclirnatic zones, apparent resistance against the weevil by vigorous trees, and the possible existence of an oviposition deterrent pheromone. A hazard rating system was developed based on biogeoclimatic zone, stand age and density, APA, and tree girth. In high hazard stands that are thinned, 70% of stems could be weevilled. Silvicultural recommendations are to "feather" clumps of lodgepole pine during spacing, and in stands aged 5 to 20 years, to increase the APA gradually, removing weevilled stems at each entry. DEDICATION To Connie Maclauchlan and W. M. Maclauchlan . ACKNOWLEDGEMENTS I would like to thank Dr. J.H.Borden for his patience, guidance and encouragement during the course of this study. I appreciate and thank my committee members Drs. R.1 Alfaro and B.D. Roitberg for their advice and review of various drafts and manuscripts. I also thank: M.C.M. Matteau, A.L. Carroll, T.L. McMullan, S. Sirvio and especially I. Moe (and all three dogs), for field and laboratory assistance; F. Bellavance for statistical assistance; and L.J. Chong, Dr. G. Gries and others in B6220 for helpful discussions; and Dr. George Harvey, and Mrs. P.M. Roden, Forest Pest Management Institute, Forestry Canada, Sault Ste. Marie, Ontario, for the isozyme collaboration work. I would also like to thank D.W. Hutcheson for his tolerance at work over the duration of this study. In particular, I thank J.R. Thompson, my husband, for his help measuring all those "little" trees, and for his understanding and support. The research was supported in part by a G.R.E.A.T. Award from the Science Council of B.C. and by the FederalIProvincial Forest Resource Development Agreement (FRDA I) Grant No. F-52-41-107. Dr. M.A. Hulme served as Liason Officer for this grant, and his guidance and advice is appreciated. TABLE OF CONTENTS .. Approval ..........................................................................................ii ... ABSTRACT ......................................................................................iii DEDICATION ..................................................................................v ACKNOWLEDGEMENTS ....................................................................vi List of Tables ....................................................................................ix .. List of Figures ................................................................................... xi1 I . INTRODUCTION ...........................................................................1 A . Life history ........................................................................... -3 B . Damage.............................................................................. -6 C . Host selection in the genus Pissodes .............................................7 D . Pheromones in the genus Pissodes ...............................................8 I1. OBJECTIVES ................................................................................10 I11 . DESCRIPTION OF BIOGEOCLIMATIC ZONES ...................................13 IV . DETERMINATION OF THE INCIDENCE, IMPACT AND SPATIAL DYNAMICS OF P . TERMINALIS ........................................................21 A . Sampling methodology ............................................................21 B . Okanagan Falls: a case study .....................................................28 1. Attack dynamics and impact in relation to stand age and tree size ......31 2 . Spatial attack dynamics ........................................................50 C . Lac le Jeune: a case study .........................................................62 1. Spatial attack dynamics and impact of P. terminalis in different stand densities ..........................................................................-64 2 . Attack dynamics in relation to stand age ....................................86 V . SPATIAL ATTACK DYNAMICS AND IMPACT OF P. TERMINALIS IN DIFFERENT BIOGEOCLIMATIC ZONES ............................................94 vii VI . INFESTATION PHENOLOGY. EMERGENCE PATTERNS AND FECUNDITY ................................................................................ 115 A . Methods ...............................................................................115 B . Results and Discussion ............................................................. 116 VII . THE SEARCH FOR A PHEROMONE MARKER .................................. 137 VIII . HAZARD RATING ....................................................................-142 IX . CONCLUSIONS ..........................................................................-161 X . APPENDIX I ............................................................................... -162 XI . APPENDIX I1 .............................................................................-173 XI1. APPENDIX 111 ...........................................................................-180 XI11. APPENDIX IV ..........................................................................-188 XIV . REFERENCES ..........................................................................-205 viii LIST OF TABLES Table 1. Description of biogeoclimatic zones sampled in this study. ....................19 Table 2. Location, year surveyed and summary of strip surveys done in fifty regenerating stands in the Kamloops Forest Region from 1987-1989. Surveys are grouped into general geographic areas, by biogeoclimatic zone and subzone, and the range of stand density, stand age and P. terminalis attack incidence is noted for each subzone. ..............................................................-22 Table 3. Location, biogeoclimatic zone classification, and summary statistics for 2 1 permanent sample plots established in the Kamloops Forest Region for long term monitoring of P. terminalis attack dynamics and impact. ..................23 Table 4. Location, size and attributes of eleven stem-mapped plots established