University of Alberta Retention Patch Characteristics and Ground Dwelling Beetle Diversity: Implications for Natural Disturbance-Based Management by Matthew Paull Pyper \Sv A thesis submitted to the Faculty of Graduate Studies and Research in partial fulfillment of the requirements for the degree of Master of Science in Forest Biology and Management Department of Renewable Resources Edmonton, Alberta Spring 2009 Library and Archives Bibliotheque et 1*1 Canada Archives Canada Published Heritage Direction du Branch Patrimoine de I'edition 395 Wellington Street 395, rue Wellington Ottawa ON K1A 0N4 OttawaONK1A0N4 Canada Canada Your file Votre reference ISBN: 978-0-494-54998-8 Our file Notre reference ISBN: 978-0-494-54998-8 NOTICE: AVIS: The author has granted a non­ L'auteur a accorde une licence non exclusive exclusive license allowing Library and permettant a la Bibliotheque et Archives Archives Canada to reproduce, Canada de reproduire, publier, archiver, publish, archive, preserve, conserve, sauvegarder, conserver, transmettre au public communicate to the public by par telecommunication ou par I'lnternet, preter, telecommunication or on the Internet, distribuer et vendre des theses partout dans le loan, distribute and sell theses monde, a des fins commerciales ou autres, sur worldwide, for commercial or non­ support microforme, papier, electronique et/ou commercial purposes, in microform, autres formats. paper, electronic and/or any other formats. The author retains copyright L'auteur conserve la propriete du droit d'auteur ownership and moral rights in this et des droits moraux qui protege cette these. Ni thesis. Neither the thesis nor la these ni des extraits substantiels de celle-ci substantial extracts from it may be ne doivent etre imprimes ou autrement printed or otherwise reproduced reproduits sans son autorisation. without the author's permission. In compliance with the Canadian Conformement a la loi canadienne sur la Privacy Act some supporting forms protection de la vie privee, quelques may have been removed from this formulaires secondaires ont ete enleves de thesis. cette these. While these forms may be included Bien que ces formulaires aient inclus dans in the document page count, their la pagination, il n'y aura aucun contenu removal does not represent any loss manquant. of content from the thesis. •+• Canada To my wife, and best friend Shelagh Marie Pyper, for supporting me in everything I do. Your love and patience has meant so much. Abstract Scientific evaluation of natural disturbance-based forest management requires fine-scale studies of biodiversity. I studied the role of retention patch size and isolation from the un-cut forest matrix in preserving communities of ground beetles (Coleoptera: Carabidae) and rove beetles (Coleoptera: Staphylinidae) in North-western Alberta. Patch size was an important driver of beetle community structure, with beetle assemblages in patches greater than two hectares most similar to the mature forest. Patch isolation had little effect on common species but resulted in decreased catches of rare species, emphasizing the need to assess rare species in forest management research. I also studied edge effects along forest-harvest edges and found narrow, abrupt edge effects. Beetle assemblages of edges began to recover by 15 years post-harvest in deciduous stands. My work suggests that increasing the frequency of large retention patches in managed forests has clear benefits for biodiversity, and that large patches can be used to reduce the negative influences of isolation. Acknowledgements First and foremost I would like to thank my supervisors, Dr. John Spence and Dr. David Langor. To Dave, for his encouragement, unique perspectives, and extensive support during initiation of this project. To John, for taking a wildlife enthusiast under his wings, providing endless opportunities, and mentoring my passion for science communication; for this, I am truly grateful. I also thank my committee members Dr. Fangliang He and Dr. Heather Proctor for comments, advice, and encouragement throughout this project; as well as members of the examining committee. I have also been fortunate to have encountered many amazing people along the road to completing this thesis. First, to my field assistants: Tomona Morita, and Wallis Johnson, for showing grit and determination. To Wallis, I hope you too develop an appreciation of our amazing wolf encounter. To Jim Witiw, for the endless hours spent helping with field site selection, and continual interaction during this project. To the Spence lab (C.Wood, J. Pinzon, T. Cobb, C. Bergeron, J. Jacobs, E. Esch, S. Lee, S. Abele, S. Bourassa, E. Kamunya, M. Koivula and C. MacQuarrie) for influential discussions, statistical advice, daily humour, constructive criticism, and lifelong friendships; it has been a good time, but not a long enough time. I also thank L. Gray and J. Fitzpatrick for spurts of field assistance, and Brett Bodeux for his enthusiasm and ideas during countless conversations. To Dustin Hartley, for the rove beetle identifications in this thesis, your help was invaluable. Jason Edwards and Charlene Halm ran the best field station a student could ask for, thanks for your discussions and support. Special thanks to Shelagh Pyper, for the many hours committed to this thesis; whether in the wee hours of the night or on trains in Bolivia, your help was unwavering. To my parents, Brian and Kathy Pyper, for instilling in me a passion for hard work, a love of the outdoors, and a desire to make a difference. To Andrew Murphy and Susan Crump for support, encouragement, and advice throughout this thesis. Funding for this work was provided by the Sustainable Forest Management Network- Networks for Centres of Excellence, ACA Grants in Biodiversity- supported by the Alberta Conservation Association, National Science and Engineering Research Council, Daishowa Marubeni International Ltd., Canadian Forest Products Ltd., and the Canadian Forest Service Graduate Supplement program. Table of Contents Chapter 1: Introduction 1 Background and Rationale 1 The Role of Community Ecology in Forest Management 2 Thesis Objectives 4 Literature Cited 5 Chapter 2: Aggregated retention patch size and beetle conservation in the boreal mixedwood forest of Alberta, Canada 8 Introduction 8 Materials and Methods 9 Study Sites 9 Study Design 10 Data Collection // Data Analyses 12 Results 14 Beetle Community 14 Single Species Comparisons 16 Indicator Species 16 Temperature 17 Discussion 17 Species Richness and Abundance 17 Impacts on the Beetle Community 18 Temperature 20 Indicator Species 21 Management Implications 22 Literature Cited 23 Appendices 33 Chapter 3: Ground and rove beetle responses to retention patch isolation: the value in considering rarity 37 Introduction 37 Materials and Methods 39 Study Design 39 Data Collection 40 Data Analyses 40 Results 43 General Results 43 Beetle Catch and Species Richness 43 Community Analyses 43 Single Species Comparisons 45 Discussion 45 Impact of Patch Isolation 46 Challenges of Analyzing Rarity 48 Conclusions 49 Literature Cited 49 Chapter 4: Spatial and temporal response of ground beetles to recent and regenerating harvest edges in the boreal forest of Alberta, Canada 60 Introduction 60 Methods 61 Study Site 61 Study Design 61 Beetle Sampling 62 Data Analyses 63 Results 65 Cover-type Effect 65 Edge Recovery 67 Discussion 68 Cover-type Effect 68 Indicator Species 70 Edge Age 71 Conclusions 72 Literature Cited 72 Chapter 5: Discussion 84 Main Results 84 Implications for Forest Management 86 Future Research and Limitations of the Dissertation 87 Literature Cited 89 List of Tables Table: 3.1: Known habitat affinities of ground and rove beetles exhibiting strong (>0.2) correlation with canonical analysis of principle co-ordinates (CAP) axis 2 55 Table 4.1: Ground beetle indicator species that exhibited the highest Indicator Values to two forest zones, identified by previous analyses, and to the two cover-types studied (deciduous dominated and conifer dominated) 77 List of Figures Figure 2.1: Map of study area showing relative proximity of harvest blocks 26 Figure 2.2: Mean (+1 S.E.) standardized catch of ground beetles and rove beetles within clear- cuts, mature forest, and three sizes of retention patches: small (<1.4 ha); medium (1.8-4.4 ha); and large (>4.5 ha); within two forest cover-types. Bars with the same letters are not significantly different (a =0.05) 27 Figure 2.3: Rarefaction-estimated species richness of ground and rove beetles within clear-cuts, mature forest, and three sizes of retention patches: small (<1.4 ha); medium (1.8-4.4 ha); and large (>4.5 ha). Two cover-types were sampled: a) deciduous dominated stands; b) conifer dominated stands. Arrows indicate sample size used to compare estimates across treatments... 27 Figure 2.4: Hierarchical clustering analysis of Bray-Curtis similarity coefficients for_ground beetles and rove beetles sampled in clear-cuts, mature forest, and three sizes of retention patches: small (<1.4 ha); medium (1.8-4.4 ha); and large (>4.5 ha). Two cover-types were sampled: a) deciduous dominated stands; b) conifer dominated stands. Numbers 1-9 represent node labels for comparison to indicator species analysis (ISA) 28 Figure 2.5: Non-metric multidimensional scaling (NMDS) ordinations of: a) ground and rove beetles; b) rove beetles; and c) ground beetles, sampled from: clear-cuts; mature forest; and small (<1.4 ha); medium (1.8-4.4 ha); and large (>4.5 ha) aggregated retention patches within two dominant forest cover-types of the boreal region 29 Figure 2.6: Standardized catch of the six most common species collected via standard pitfall trapping within clear-cuts, mature forest, and three sizes of retention patches: small (<1.4 ha); medium (1.8-4.4 ha); and large (>4.5 ha); and in two of the dominant cover-types (deciduous dominated and conifer dominated) 30 Figure 2.7: Significant ground beetle and rove beetle indicator species (p<0.05) for each node of a hierarchical clustering analysis.