SABROXYLIC BEETLES and DEADWOOD STRUCTURE in MANAGED and NATURALLY DISTURBED SPRUCE FOWSTS in NOVA SCOTTA Delancey J. Bishop, B
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SABROXYLIC BEETLES AND DEADWOOD STRUCTURE IN MANAGED AND NATURALLY DISTURBED SPRUCE FOWSTS IN NOVA SCOTTA by DeLancey J. Bishop, B.Sc.H. A thesis submitted to the Facuity of Graduate Studies and Research in partial ~fi~hentof the requirements for the degree of Master of Science Department of Biology Carleton Universiv Ottawa, Ontario, Canada 28 May 1998 O DeLancey J. Bishop 1998 National Library Bibliothèque nationale I*m of Canada du Canada Acquisitions and Acquisitions et Bibliographie Services services bibliographiques 395 Wellington Street 395, rue Wellington Ottawa ON KIA ON4 Olrawa ON KiA ON4 Canada Canada Your file Voire référence Our iYe Notre réltirencB The author has granted a non- L'auteur a accordé une licence non exclusive licence allowing the exclusive pennethnt à la National Library of Canada to Bibliothèque nationale du Canada de reproduce, loan, distribute or seil reproduire, prêter, distribuer ou copies of t thesis in microfonn, vendre des copies de cette thèse sous paper or electronic formats. la fome de microfichelfilm, de reproduction sur papier ou sur format électronique. The author retains ownership of the L'auteur conserve Ia propriete du copyright in this thesis. Neither the droit d'auteur qui protège cette thèse. thesis nor substantial extracts fiom it Ni la thèse ni des extraits substantiels may be printed or otherwise de celle-ci ne doivent être imprimés reproduced without the author's ou autrement reproduits sans son pmission. autorisation. ABSTRACT Modem forestry alters invertebrate assemblages by altering the microhabitats of invertebrate species. 1studied relationships between saproxylic beeties and deadwood structures in spruce forests with natural (wind and fire) and forestry (clearcut and clearcut with thinning) disturbance histones. 1tested the predictions that species assemblages would ciiffer in composition and be less diverse in managed forests due to a different and lower diversity of deadwood structures. Little evidence supported the prediction of lower species diversity, eirher locally or across an array of forests. However, analysis revealed diEerent assemblage compositions between managed and natiirally disturbed forests, and lower assemblage turnover between managed forests. I discerned species that may indicate different disturbance histones, species correlated with different forest structures, and the forest structures that best explain differences in species assemblages between farests of different disturbance histories. 1make research and management recornmendations regarding the preservation of saproxylic species in managed forests. ACKNO WLEDGMENTS If1 have approached scientific rationaliv during these past two years it is iargely because of my supe~sor,Stewart Peck; his wisdom, patience, and kindness have allowed my time at Carleton to be a pleasant and productive learning experience. My advisors M. Forbes and F. Chapleau provided usefui criticism dong the way. S. Bondnip-Nielsen of Acadia University enthiisiastically contrïbuted to the study design and provided nearly dl sampling matenals. R. Cameron kindly and greatly assisted the forest selection process and offered laboriously-collected forest data without hesitation. K. Potter aiso went out of her way to provide previously collected forest data- L. Parriag, L. Maddison, and C. Bishop provided fuie field and laboratory assistance. The following coleopterists kindly identified whatever form of beetle 1 could not: J. Cook (Carleton University), F. Andrews (USDFA Analysis and Identification, California), D. Chandler (University of New Hampshire), J. Klirnaszewski (BC Research, British Columbia), Y. Bousquet, D. Bright, A. Davies, S. Laplante, and A. Smetana (al1 fiom the Agriculture Canada Biosystematics Research Center, Ottawa). Without the support of these individuals and the availability of the invaluable Canadian National Collection this work would be less than what it is- hundreds of species less. A scholarship fiom the Natural Sciences and Engineering Research Couocil funded this aspect of my education. The Nova Scotia Department of Naturai Resources supported a summer assistant, thanks to T. Duke and J.S. Boates. Another keystone of this project is J. Kukalova-Peck, who's motherly kindness perhaps saved this oft homesick boy. And 1am never without the love of Mom, Dad, Linda, and Cohen. Finally, 1thank the saproxylic beetles; 1hope this helps you! TABLE OF CONTENTS - * LIST OF TPLBLES ..~........-.......~........,~,............-.~.~.,,..,,,,........,...................~.....~+...............~vil LIST OF FIGURES.........~...~...~.~.-......,....~...-...~-.......~..~..~......,..~...~.-.-.-.-....~.--....-......-........ ix LIST OF APPENDICES ....................... ,., ......-.-............-S...-..-.........................-..--............--.. x CKA-PTER ONE DEADWOOD DIVERSITY AND SAPROXYLIC BEETLE DIVERSlTY IN MANAGED AND NATURALLY DISTURBED SPRUCE FOREST'S: TESTIh'G RELATIONSHIPS AND ASKING WHETHER ECOLOGICAL INTEGRZTY 1s VTOLATED BY MODERN FORESTRY PRACTICES ..-....... .. .. ... -.... ..--... ,........ .......--1 INTRODUCTION.. .. - ., , . - .. .. METHO D S... -. .+. .. .. , . -. - .. .-. .. -. - --... -. -. - -. - -. -. .- -... -. -. .-. .. - - -..- -.. -. .. - .- -..-6 Study forests..... ............ .... ....--....--...-.......... .. .. ..---.......-.... ..-.. ...-.-..--..--. .... ... .. .. .....--6 Forest habitat structure.. ... .. .. .--. .. -.- .. ... .. , . .. .... .. .. -7 Saproxylic beetle sampling and identification....... - ...-................... ................... 1 1 -. Statist~calanalyses .............-..----... .-.-... .... --.-..-.....-...... ....-..-....... .. .... .-................... 13 RESULTS ............ - ..... - ............................................................................19 Local diversity scale............................................................................................ 73 Beta diversity scaie............................................................................................ 26 Gamma diversity scale.............. ...-..-......--. .......................................................... 29 DISCUSSION ...................................... ................................... ......................................... -3 1 The beta-scale: spatial turnover in forest composition and ecological integrity.3 3 More of fewer species? A potential sampling enor with dire consequences..... 36 Future research and forest management in Nova Scotia............................. .......-- 3 8 CHAPTER TWO SAPROXYLIC BEETLE SPECIES AS INDICATORS OF DISTURBANCE HISTORY AND FOREST STRUCTURE IN NOVA SCOTIA SPRUCE FORESTS............ ..... 43 INTRODUCTION...................................................................................................... $3 METHODS ................................................................................................................. 46 .. Statistical analyses .......... ... ........................................................................... -46 Species indicators of forest disturbance history............................. .... ................. 49 Species-structure models.................... ..... .................................................. 30 Fauna differentiating forest disturbance classes and structural correlates .......... 51 DISCUSSION............................................................................................................. 55 Indicators of forest disturbance and indicators of forest structures......... .. ....... 56 Observation and hypothesis testiog towards confirming or rejecting species as . mdicators ............................................................................................................. 58 L1TERA.TUR.E CITED ...................................................................................................... 61 LIST OF TABLES CHAPTER 1 Table 1. Mean age of dominant canopy trees and generd disturbance history of each forest................. .-..... ..... ..-.. ..............................................................................-9 Table 2. Distribution of forests delineated by age and disturbance history classes ............ 9 Table 3. Description of forest structure variables measured in each forest ...................... 10 Table 4. Description of calculated habitat (A) and beetle (B) diversity amibutes ............ L 5 Table 5. Spearman"srank correlations between forest structure variables ............ .. .....20 Table 6- Mean and standard deviation of each forest structure within forest disturbance classes, and Kniskal-Wallis results for test of differences between disturbance classes ...20 Table 7. Mean and standard deviation of each forest structure diversity attribute within forest disturbance classes, and ANOVA results for test of differences between disturbance classes ............................................................................................................. 24 Table 8. Mean and standard deviation of each beetle assemblage diversity amibute within forest disturbance classes, and ANOVA results for test of differences between disturbance classes ........................................................................................................... ..25 Table 9. Significant multiple linear regcession models of beetle assemblage amibutes regressed against forest structure diversity amibutes.. .............. ... ...............................-25 Table 10. Results of multi-response permutation procedures analysis, testing