Conservation of Saproxylic Insect Diversity Under Variable Retention Harvesting

  • Seung-Il LeeEmail author
  • John R. Spence
  • David W. Langor
Part of the Zoological Monographs book series (ZM, volume 1)


Saproxylic (i.e., deadwood-associated) insects are important functional components of biological diversity in forest ecosystems; however, they depend on microhabitats associated with dying, dead, and decaying wood that are dramatically altered by industrial forestry. Habitat loss and fragmentation by activities, such as clear-cutting, salvage logging, and bioenergy extraction, threaten saproxylic biodiversity on forested landscapes through changes in quantity, quality, and dynamics of deadwood. Retention forestry has been proposed and widely applied to support conservation and recovery of biodiversity and the associated ecological function on managed landscapes. In spite of its short history, retention forestry has undoubtedly had positive effects on biodiversity compared to conventional clear-cut harvest. The amount and pattern of retention are two important factors that determine biotic and abiotic responses and thereby influence success of retention approach. We review major findings from several large-scale variable retention experiments that have considered impacts on saproxylic insects. General conclusions from these experiments include the following three points: (1) aggregated retention conserves saproxylic insect faunas better than dispersed retention; (2) mixes of aggregated and dispersed retention have greater conservation value for saproxylic beetles than a single retention type; and (3) inclusion of prescribed burning will improve the conservation performance of retention forestry approaches. Successful conservation of saproxylic insect populations will likely depend on management to ensure availability of a full range of deadwood quantities and qualities in harvested and regenerating forest stands. We argue that variable retention harvesting, which includes sufficient amounts of retained trees as combinations of aggregated and dispersed retention, will best support diversity of deadwood habitats in the harvested matrix and promote conservation and local recovery of saproxylic assemblages.



We thank Susan Baker and Jari Kouki for providing valuable information about retention forestry practices in Australia and northern Europe. Their efforts greatly improved this manuscript.


  1. Abrahamsson M, Lindbladh M (2006) A comparison of saproxylic beetle occurrence between man-made high- and low-stumps of spruce (Picea abies). For Ecol Manag 226:230–237CrossRefGoogle Scholar
  2. Alexander KNA (2008) Tree biology and saproxylic Coleoptera: issues of definitions and conservation language. Revue d’écologie – La Terre et la Vie 63:1–5Google Scholar
  3. Andison DW (2004) Island remnants on foothills and mountain landscapes of Alberta: part 2 on residuals. Alberta Foothills Disturbance Ecology Research Series, Report No. 6, November 1994. Foothills Model Forest, Hinton, AlbertaGoogle Scholar
  4. Aubry KB, Amaranthus MP, Halpern CB et al (1999) Evaluating the effects of varying levels and patterns of green-tree retention: experimental design of the DEMO Study. Northwest Sci 73:12–26Google Scholar
  5. Aubry KB, Halpern CB, Peterson CE (2009) Variable-retention harvests in the Pacific Northwest: a review of short-term findings from the DEMO study. For Ecol Manag 258:398–408CrossRefGoogle Scholar
  6. Baker SC (2011) Seeking a balance between forestry and biodiversity – the role of variable retention silviculture. Insights from western USA and Canada. Forest & Wood Products Australia, Melbourne, p 60Google Scholar
  7. Baker SC, Read SM (2011) Variable retention silviculture in Tasmania’s wet forests: ecological rationale, adaptive management and synthesis of biodiversity benefits. Aust For 74:218–232CrossRefGoogle Scholar
  8. Baker SC, Barmuta LA, McQuillan PB et al (2007) Estimating edge effects on ground-dwelling beetles at clearfelled non-riparian stand edges in Tasmanian wet eucalypt forest. For Ecol Manag 239:92–101CrossRefGoogle Scholar
  9. Baker SC, Grove SJ, Forster L et al (2009) Short-term responses of ground-active beetles to alternative silvicultural systems in the Warra Silvicultural Systems Trial, Tasmania, Australia. For Ecol Manag 258:444–459CrossRefGoogle Scholar
  10. Baker SC, Spies TA, Wardlaw TJ et al (2013) The harvested side of edges: effect of retained forests on the re-establishment of biodiversity in adjacent harvested areas. For Ecol Manag 302:107–121CrossRefGoogle Scholar
  11. Baker SC, Halpern CB, Wardlaw TJ et al (2015) Short- and long-term benefits for forest biodiversity of retaining unlogged patches in harvested area. For Ecol Manag 353:187–195CrossRefGoogle Scholar
  12. Baker SC, Halpern CB, Wardlaw TJ et al (2016) A cross-continental comparison of plant and beetle responses to retention of forest patches during timber harvest. Ecol Appl 26:2493–2504PubMedPubMedCentralGoogle Scholar
  13. Baker SC, Grove SJ, Wardlaw TJ et al (2017) Monitoring the implementation of variable retention silviculture in wet eucalypt forest: a key element of successful adaptive management. For Ecol Manag 394:27–41CrossRefGoogle Scholar
  14. Bergeron Y, Leduc A, Harvey BD et al (2002) Natural fire regime: a guide for sustainable management of the Canadian boreal forest. Silva Fenn 36:81–95CrossRefGoogle Scholar
  15. Bergeron JAC, Pinzon J, Odsen S et al (2017) Ecosystem memory of wildfires affects resilience of boreal mixedwood biodiversity after retention harvest. Oikos 126:1738–1747CrossRefGoogle Scholar
  16. Bouget C, Lassauce A, Jonsell M (2012a) Effects of fuelwood harvesting on biodiversity – a review focused on the situation in Europe. Can J For Res 42:1421–1432CrossRefGoogle Scholar
  17. Bouget C, Nusillard B, Pineau X et al (2012b) Effect of deadwood position on saproxylic beetles in temperate forests and conservation interest of oak snags. Insect Conserv Diver 5:264–278CrossRefGoogle Scholar
  18. Bouget C, Larrieu L, Brin A (2014) Key features for saproxylic beetle diversity derived from rapid habitat assessment in temperate forests. Ecol Indic 36:656–664CrossRefGoogle Scholar
  19. Bunnell FL, Kremsater LL, Wind E (1999) Managing to sustain vertebrate richness in forests of the Pacific Northwest: relationships within stands. Environ Rev 7:97–146CrossRefGoogle Scholar
  20. Cobb TP, Morissette JL, Jacobs JM et al (2011) Effects of postfire salvage logging on deadwood-associated beetles. Conserv Biol 25:94–104PubMedCrossRefGoogle Scholar
  21. Daishowa-Marubeni International Ltd (2013) FMA operating ground rules. Daishowa-Marubeni International Ltd., Peace River Pulp Division, Alberta, p 88Google Scholar
  22. den Herder M, Kouki J, Ruusila V (2009) The effects of timber harvest, forest fire, and herbivores on regeneration of deciduous trees in boreal pine-dominated forests. Can J For Res 39:712–722CrossRefGoogle Scholar
  23. Diamond JM (1976) Island biogeography and conservation: strategy and limitations. Science 193:1027–1029PubMedCrossRefGoogle Scholar
  24. Djupström LB, Weslien J, Schroeder LM (2008) Dead wood and saproxylic beetles in set-aside and non set-aside forests in a boreal region. For Ecol Manag 255:3340–3350CrossRefGoogle Scholar
  25. Dollin PE, Majka CG, Duinker PN (2008) Saproxylic beetle (Coleoptera) communities and forest management practices in coniferous stands in southwestern Nova Scotia, Canada. In: Majka CG, Klimaszewski J (eds) Biodiversity, biosystematics, and ecology of Canadian Coleoptera, ZooKeys, vol 2, pp 291–336Google Scholar
  26. Enrong Y, Xihua W, Jianjun H (2006) Concept and classification of coarse woody debris in forest ecosystems. Front Biol (Beijing) 1:76–84Google Scholar
  27. Esseen PA, Ehnström B, Ericson L et al (1997) Boreal forests. Ecol Bull 46:16–47Google Scholar
  28. Ewers RM, Didham RK (2008) Pervasive impact of large-scale edge effects on a beetle community. PNAS 105:5426–5429PubMedPubMedCentralCrossRefGoogle Scholar
  29. Fedrowitz K, Koricheva J, Baker SC et al (2014) Can retention forestry help conserve biodiversity? A meta-analysis. J Appl Ecol 51:1669–1679PubMedPubMedCentralCrossRefGoogle Scholar
  30. Ferro ML, Gimmel ML, Harms KE et al (2012) Comparison of Coleoptera emergent from various decay classes of downed coarse woody debris in Great Smoky Mountains National Park, USA. Insecta Mundi 0260:1–8Google Scholar
  31. Fogel RM, Ogawa M, Trappe JM (1973) Terrestrial decomposition: a synopsis. US IBP Coniferous Forest Biome Report 135. Coniferous Forest Biome, College of Forest Resources, University of Washington, Seattle, p 12Google Scholar
  32. Franc N (2007) Standing or downed dead trees – does it matter for saproxylic beetles in temperate oak-rich forest? Can J For Res 37:2494–2507CrossRefGoogle Scholar
  33. Franklin JF, Shugart HH, Harmon ME (1987) Tree death as an ecological process. Bioscience 37:550–556CrossRefGoogle Scholar
  34. Franklin JF, Berg DR, Thornburgh DA et al (1997) Alternative silvicultural approaches to timber harvesting: variable retention harvest systems. In: Kohm KA, Franklin JF (eds) Creating a forestry for the 21st century: the science of ecosystem management. Island Press, Washington, DC, pp 111–139Google Scholar
  35. Gibb H, Ball JP, Johansson T et al (2005) Effects of management on coarse woody debris volume and composition in boreal forests in northern Sweden. Scand J For Res 20:213–222CrossRefGoogle Scholar
  36. Grove SJ (2002) Saproxylic insect ecology and the sustainable management of forests. Annu Rev Ecol Syst 33:1–23CrossRefGoogle Scholar
  37. Grove SJ, Forster L (2011a) A decade of change in the saproxylic beetle fauna of eucalypt logs in the Warra long-term log-decay experiment, Tasmania. 1. Description of the fauna and seasonality patterns. Biodivers Conserv 20:2167–2188CrossRefGoogle Scholar
  38. Grove SJ, Forster L (2011b) A decade of change in the saproxylic beetle fauna of eucalypt logs in the Warra long-term log-decay experiment, Tasmania. 2. Log-size effects, succession, and the functional significance of rare species. Biodivers Conserv 20:2167–2188CrossRefGoogle Scholar
  39. Grove SJ, Stamm L (2011) Downed woody debris in Tasmanian eucalypt forest: modelling the effects of stand-replacing disturbance dynamics. Division of Forest Research and Development Technical Report 15/2011. Forestry Tasmania, Hobart, p 28Google Scholar
  40. Grove SJ, Stamm L, Barry C (2009) Log decomposition rates in Tasmanian Eucalyptus obliqua determined using an indirect chronosequence approach. For Ecol Manag 258:389–397CrossRefGoogle Scholar
  41. Grove SJ, Stamm L, Wardlaw TJ (2011) How well does a log decay-class system capture the ecology of decomposition? – a case-study from Tasmanian Eucalyptus obliqua forest. For Ecol Manag 262:692–700CrossRefGoogle Scholar
  42. Gustafsson L, Kouki J, Sverdrup-Thygeson A (2010) Tree retention as a conservation measure in clear-cut forests of northern Europe: a review of ecological consequences. Scand J For Res 25:295–308CrossRefGoogle Scholar
  43. Gustafsson L, Baker SC, Bauhus J et al (2012) Retention forestry to maintain multifunctional forests: a world perspective. Bioscience 62:633–645CrossRefGoogle Scholar
  44. Hagan JM, Grove SL (1999) Coarse woody debris. J For 97:6–11Google Scholar
  45. Halaj J, Halpern CB, Yi H (2008) Responses of litter-dwelling spiders and carabid beetles to varying levels and patterns of green-tree retention. For Ecol Manag 255:887–900CrossRefGoogle Scholar
  46. Halaj J, Halpern CB, Yi H (2009) Effects of green-tree retention on abundance and guild composition of corticolous arthropods. For Ecol Manag 258:850–859CrossRefGoogle Scholar
  47. Hale CM, Pastor J (1998) Nitrogen content, decay rates, and decompositional dynamics of hollow versus solid hardwood logs in hardwood forests of Minnesota, U.S.A. Can J For Res 28:1276–1285CrossRefGoogle Scholar
  48. Halme E, Niemelä J (1993) Carabid beetles in fragments of coniferous forest. Ann Zool Fenn 30:17–30Google Scholar
  49. Halpern CB, McKenzie D, Evans SA et al (2005) Initial responses of forest understories to varying levels and patterns of green-tree retention. Ecol Appl 15:175–195CrossRefGoogle Scholar
  50. Hämäläinen A, Hujo M, Heikkala O et al (2016) Retention tree characteristics have major influence on the post-harvest tree mortality and availability of coarse woody debris in clear-cut areas. For Ecol Manag 369:66–73CrossRefGoogle Scholar
  51. Hammond HEJ, Langor DW, Spence JR (2004) Saproxylic beetles (Coleoptera) using Populus in boreal aspen stands of western Canada: spatiotemporal variation and conservation of assemblages. Can J For Res 34:1–19CrossRefGoogle Scholar
  52. Hammond HEJ, Langor DW, Spence JR (2017) Changes in saproxylic beetle (Insecta: Coleoptera) assemblages following wildfire and harvest in boreal Populus forests. For Ecol Manag 401:319–329CrossRefGoogle Scholar
  53. Harmon ME, Franklin JF, Swanson FJ (1986) Ecology of coarse woody debris in temperate ecosystems. Adv Ecol Res 15:133–302CrossRefGoogle Scholar
  54. Harmon ME, Nadelhoffer KJ, Blair JM (1999) Measuring decomposition, nutrient turnover, and stores in plant litter. In: Robertson GP, Coleman DC, Bledsoe CS, Sollins P (eds) Standard soil methods for long-term ecological research. Oxford University Press, New York, pp 202–240Google Scholar
  55. Heikkala O, Suominen M, Junninen K (2014) Effects of retention level and fire on retention tree dynamics in boreal forests. For Ecol Manag 328:193–201CrossRefGoogle Scholar
  56. Heikkala O, Martikainen P, Kouki J (2016a) Decadal effects of emulating natural disturbances in forest management on saproxylic beetle assemblages. Biol Conserv 194:39–47CrossRefGoogle Scholar
  57. Heikkala O, Seibold S, Koivula M et al (2016b) Retention forestry and prescribed burning result in functionally different saproxylic beetle assemblages than clear-cutting. For Ecol Manag 359:51–58CrossRefGoogle Scholar
  58. Heikkala O, Kouki J, Martikainen P (2017) Prescribed burning is an effective and quick method to conserve rare pyrophilous forest-dwelling flat bugs. Insect Conserv Diver 10:32–41CrossRefGoogle Scholar
  59. Hickey JE, Neyland MG, Grove SJ (2006) From little things big things grow: the Warra Silvicultural Systems Trial in Tasmanian wet Eucalyptus obliqua forest. Allg Forst- u J-Ztg 177(Jg, 6/7):113–119Google Scholar
  60. Hjältén J, Stenbacka F, Andersson J (2010) Saproxylic beetle assemblages on low stumps, high stumps and logs: implications for environmental effects of stump harvesting. For Ecol Manag 260:1149–1155CrossRefGoogle Scholar
  61. Hjältén J, Stenbacka F, Pettersson RB et al (2012) Micro and macro-habitat associations in saproxylic beetles: implications for biodiversity management. PLoS One 7(7):e41100. Scholar
  62. Hofgaard A (1993) Structure and regeneration patterns in a virgin Picea abies forest in northern Sweden. J Veg Sci 4:601–608CrossRefGoogle Scholar
  63. Hunter ML (1993) Natural fire regimes as spatial models for managing boreal forests. Biol Conserv 65:115–120CrossRefGoogle Scholar
  64. Hunter ML (1999) Maintaining biodiversity in forest ecosystems. Cambridge University Press, CambridgeCrossRefGoogle Scholar
  65. Hyvärinen E, Kouki J, Martikainen P et al (2005) Short-term effects of controlled burning and green-tree retention on beetle (Coleoptera) assemblages in managed boreal forests. For Ecol Manag 212:315–332CrossRefGoogle Scholar
  66. Hyvärinen E, Kouki J, Martikainen P (2006) Fire and green-tree retention in conservation of red-listed and rare deadwood-dependent beetles in Finnish boreal forests. Conserv Biol 20:1711–1719PubMedCrossRefPubMedCentralGoogle Scholar
  67. Hyvärinen E, Kouki J, Martikainen P (2009) Prescribed fires and retention trees help to conserve beetle diversity in managed boreal forests despite their transient negative effects on some beetle groups. Insect Conserv Diver 2:93–105CrossRefGoogle Scholar
  68. Jacobs JM, Spence JR, Langor DW (2007a) Influence of boreal forest succession and dead wood qualities on saproxylic beetles. Agr Forest Entomol 9:3–16CrossRefGoogle Scholar
  69. Jacobs JM, Spence JR, Langor DW (2007b) Variable retention harvesting of white spruce stands and saproxylic beetle assemblages. Can J For Res 37:1631–1642CrossRefGoogle Scholar
  70. Johansson V, Felton A, Ranius T (2016) Long-term landscape scale effects of bioenergy extraction on dead wood-dependent species. For Ecol Manag 371:103–113CrossRefGoogle Scholar
  71. Jonsell M, Hansson J, Wedmo L (2007) Diversity of saproxylic beetle species in logging residues in Sweden – comparisons between tree species and diameters. Biol Conserv 138:89–99CrossRefGoogle Scholar
  72. Jonsson BG (2000) Availability of coarse woody debris in a boreal old-growth Picea abies forest. J Veg Sci 11:51–56CrossRefGoogle Scholar
  73. Jonsson BG, Siitonen J (2012) Dead wood and sustainable forest management. In: Stokland JN, Siitonen J, Jonsson BG (eds) Biodiversity in dead wood. Cambridge University Press, New York, pp 302–337CrossRefGoogle Scholar
  74. Jonsson BG, Kruys N, Ranius T (2005) Ecology of species living on dead wood – lessons for dead wood management. Silva Fenn 39:289–309CrossRefGoogle Scholar
  75. Jönsson MT, Fraver S, Jonsson BG et al (2007) Eighteen years of tree mortality and structural change in an experimentally fragmented Norway spruce forest. For Ecol Manag 242:306–313CrossRefGoogle Scholar
  76. Keller M, Palace M, Asner GP et al (2004) Coarse woody debris in undisturbed and logged forests in the eastern Brazilian Amazon. Glob Chang Biol 10:784–795CrossRefGoogle Scholar
  77. Klimaszewski J, Langor DW, Savard K et al (2007) Rove beetles (Coleoptera: Staphylinidae) in yellow birch-dominated stands of southeastern Quebec, Canada: diversity, abundance, and description of a new species. Can Entomol 139:793–833CrossRefGoogle Scholar
  78. Kouki J, Löfman S, Martikainen P et al (2001) Forest fragmentation in Fennoscandia: linking habitat requirements of wood-associated threatened species to landscape and habitat changes. Scand J For Res Suppl 3:27–37CrossRefGoogle Scholar
  79. Kunttu P, Junninen K, Kouki J (2015) Dead wood as an indicator of forest naturalness: a comparison of methods. For Ecol Manag 353:30–40CrossRefGoogle Scholar
  80. Lachat T, Müller J (2018) Importance of primary forests for the conservation of saproxylic insects. In: Ulyshen MD (ed) Saproxylic insects: diversity, ecology and conservation. Springer, Heidelberg, pp 581–605CrossRefGoogle Scholar
  81. Langor DW, Hammond HEJ, Spence J et al (2008) Saproxylic insect assemblages in Canadian forests: diversity, ecology, and conservation. Can Entomol 140:453–474CrossRefGoogle Scholar
  82. Laurance WF (2000) Do edge effects occur over large spatial scales? Trends Ecol Evol 15:134–135PubMedCrossRefPubMedCentralGoogle Scholar
  83. Lee S-I, Spence JR, Langor DW (2014) Succession of saproxylic beetles associated with decomposition of boreal white spruce logs. Agric For Entomol 16:391–405CrossRefGoogle Scholar
  84. Lee S-I, Spence JR, Langor DW et al (2015) Retention patch size and conservation of saproxylic beetles in boreal white spruce stands. For Ecol Manag 358:98–107CrossRefGoogle Scholar
  85. Lee S-I, Spence JR, Langor DW (2017) Combinations of aggregated and dispersed retention improve conservation of saproxylic beetles in boreal white spruce stands. For Ecol Manag 385:116–126CrossRefGoogle Scholar
  86. Lencinas MV, Martínez Pastur G, Gallo E et al (2009) Alternative silvicultural practices with variable retention improve bird conservation in managed South Patagonian forests. For Ecol Manag 258:472–480CrossRefGoogle Scholar
  87. Lencinas MV, Martínez Pastur G, Gallo E et al (2011) Alternative silvicultural practices with variable retention to improve understory plant diversity conservation in southern Patagonian forests. For Ecol Manag 262:1236–1250CrossRefGoogle Scholar
  88. Lindenmayer DB, Franklin JF (2002) Conserving forest biodiversity: a comprehensive multi-scaled approach. Island Press, Washington, DCGoogle Scholar
  89. Lindenmayer DB, Burton PJ, Franklin JF (2008) Salvage logging and its ecological consequences. Island Press, Washington, DCGoogle Scholar
  90. Lindenmayer DB, Franklin JF, Lõhmus A et al (2012) A major shift to the retention approach for forestry can help resolve some global forest sustainability issues. Conserv Lett 5:421–431CrossRefGoogle Scholar
  91. Martikainen P, Kouki J (2003) Sampling the rarest: threatened beetles in boreal forest biodiversity inventories. Biodivers Conserv 12:1815–1831CrossRefGoogle Scholar
  92. Martikainen P, Kouki J, Heikkala O (2006a) The effects of green tree retention and subsequent prescribed burning on ground beetles (Coleoptera: Carabidae) in boreal pine-dominated forests. Ecography 29:659–670CrossRefGoogle Scholar
  93. Martikainen P, Kouki J, Heikkala O et al (2006b) Effects of green tree retention and prescribed burning on the crown damage caused by the pine shoot beetles (Tomicus spp.) in pine-dominated timber harvest areas. J Appl Entomol 130:37–44CrossRefGoogle Scholar
  94. Maser C, Anderson RG, Comack K Jr et al (1979) Dead and down woody material. In: Thomas JW (ed) Wildlife habitats in managed forests: The Blue Mountains of Oregon and Washington, USDA Forest Service Agriculture Handbook, vol 877. Wildlife Management Institute, Washington, DC, pp 78–95Google Scholar
  95. Matveinen-Huju K, Niemelä J, Rita H et al (2006) Retention-tree groups in clear-cuts: do they constitute ‘life-boats’ for spiders and carabids? For Ecol Manag 230:119–135CrossRefGoogle Scholar
  96. McCullough HA (1948) Plant succession on fallen logs in a virgin spruce-fir forest. Ecology 29:508–513CrossRefGoogle Scholar
  97. Mitchell SJ, Beese WJ (2002) The retention system: reconciling variable retention with the principles of silvicultural systems. For Chron 78:397–403CrossRefGoogle Scholar
  98. Næsset E (1999) Relationship between relative wood density of Picea abies logs and simple classification systems of decayed coarse woody debris. Scand J For Res 14:454–461CrossRefGoogle Scholar
  99. Park J, Reid ML (2007) Distribution of a bark beetle, Trypodendron lineatum, in a harvested landscape. For Ecol Manag 242:236–242CrossRefGoogle Scholar
  100. Pearce JL, Venier LA, Eccles G et al (2005) Habitat islands, forest edge and spring-active invertebrate assemblages. Biodivers Conserv 14:2949–2969CrossRefGoogle Scholar
  101. Pinzon J, Spence JR, Langor DW (2012) Responses of ground-dwelling spiders (Araneae) to variable retention harvesting practices in the boreal forest. For Ecol Manag 266:42–53CrossRefGoogle Scholar
  102. Pinzon J, Spence JR, Langor DW et al (2016) Ten-year responses of ground-dwelling spiders to retention harvest in the boreal forest. Ecol Appl 26:2581–2599CrossRefGoogle Scholar
  103. Pitkänen A, Kouki J, Viiri H et al (2008) Effects of controlled forest burning and intensity of timber harvesting on the occurrence of pine weevils, Hylobius spp., in regeneration areas. For Ecol Manag 255:522–529CrossRefGoogle Scholar
  104. Pyper MP (2009) Retention patch characteristics and ground dwelling beetle diversity: implications for natural disturbance-based management. MSc thesis, University of AlbertaGoogle Scholar
  105. Rassi P, Hyvärinen E, Juslén A, Mannerkoski I (eds) (2010) The 2010 red list of Finnish Species. Ympäristöministeriö & Suomen ympäristökeskus, Helsinki, p 685Google Scholar
  106. Sahlin E, Ranius T (2009) Habitat availability in forests and clearcuts for saproxylic beetles associated with aspen. Biodivers Conserv 18:621–638CrossRefGoogle Scholar
  107. Schmitt CB, Burgess ND, Coad L et al (2009) Global analysis of the protection status of the world’s forests. Biol Conserv 142:2122–2130CrossRefGoogle Scholar
  108. Scott RE, Mitchell SJ (2005) Empirical modelling of windthrow risk in partially harvested stands using tree, neighbourhood, and stand attributes. For Ecol Manag 218:193–209CrossRefGoogle Scholar
  109. Serrouya R, D’Eon R (2004) Variable retention forest harvesting: research synthesis and implementation guidelines. Knowledge Exchange and Technology Extension Program (KETE). Sustainable Forest Management Network, EdmontonGoogle Scholar
  110. Siitonen J (2001) Forest management, coarse woody debris and saproxylic organisms: Fennoscandian boreal forests as an example. Ecol Bull 49:11–41Google Scholar
  111. Siitonen J (2012) Threatened saproxylic species. In: Stokland JN, Siitonen J, Jonsson BG (eds) Biodiversity in dead wood. Cambridge University Press, New York, pp 356–379CrossRefGoogle Scholar
  112. Siitonen J, Martikainen P (1994) Occurrence of rare and threatened insects living on decaying Populus tremula: a comparison between Finnish and Russian Karelia. Scand J For Res 9:185–191CrossRefGoogle Scholar
  113. Siitonen J, Martikainen P, Punttila P et al (2000) Coarse woody debris and stand characteristics in mature managed and old-growth boreal mesic forests in southern Finland. For Ecol Manag 128:211–225CrossRefGoogle Scholar
  114. Similä M, Kouki J, Martikainen P et al (2002) Conservation of beetles in boreal pine forests: the effects of forest age and naturalness on species assemblages. Biol Conserv 106:19–27CrossRefGoogle Scholar
  115. Similä M, Kouki J, Martikainen P (2003) Saproxylic beetles in managed and seminatural Scots pine forests: quality of dead wood matters. For Ecol Manag 174:365–381CrossRefGoogle Scholar
  116. Simonsson P, Gustafsson L, Östlund L (2015) Retention forestry in Sweden: driving forces, debate and implementation 1968–2003. Scand J For Res 30:154–173CrossRefGoogle Scholar
  117. Solarik KA, Volney WJA, Lieffers VJ et al (2012) Factors affecting white spruce and aspen survival after partial harvest. J Appl Ecol 49:145–154CrossRefGoogle Scholar
  118. Sollins P (1982) Input and decay of coarse woody debris in coniferous stands in western Oregon and Washington. Can J For Res 12:18–28CrossRefGoogle Scholar
  119. Speight MCD (1989) Saproxylic invertebrates and their conservation. Council of Europe, StrasbourgGoogle Scholar
  120. Spence JR (2001) The new boreal forestry: adjusting timber management to accommodate biodiversity. Trends Ecol Evol 16:591–593CrossRefGoogle Scholar
  121. Spence JR, Volney WJA, Lieffers VJ et al (1999) The Alberta EMEND project: recipe and cook’s argument. In: Veeman TS, Smith DW, Purdy BG et al (eds) Proceedings of the 1999 sustainable forest management network conference-science and practice. Sustaining the boreal forest. SFM Network, Edmonton, pp 583–590Google Scholar
  122. Stevens V (1997) The ecological role of coarse woody debris: an overview of the ecological importance of CWD in B.C. forests. Research Branch, B.C. Ministry of Forests, Victoria, Working Paper 30Google Scholar
  123. Stokland JN, Siitonen J, Jonsson BG (2012) Biodiversity in dead wood. Cambridge University Press, New YorkCrossRefGoogle Scholar
  124. Swanson ME, Franklin JF, Beschta RL et al (2011) The forgotten stage of forest succession: early-successional ecosystems on forest sites. Front Ecol Environ 9:117–125CrossRefGoogle Scholar
  125. Tikkanen O-P, Martikainen P, Hyvärinen E et al (2006) Red-listed boreal forest species of Finland: associations with forest structure, tree species, and decaying wood. Ann Zool Fenn 43:373–383Google Scholar
  126. Tikkanen O-P, Heinonen T, Kouki J et al (2007) Habitat suitability models of saproxylic red-listed boreal forest species in long-term matrix management: cost-effective measures for multi-species conservation. Biol Conserv 140:359–372CrossRefGoogle Scholar
  127. Ulyshen MD, Hanula JL (2009) Habitat associations of saproxylic beetles in the southeastern United States: a comparison of forest types, tree species and wood postures. For Ecol Manag 257:653–664CrossRefGoogle Scholar
  128. Ulyshen MD, Hanula JL (2010) Patterns of saproxylic beetle succession in loblolly pine. Agr For Entomol 12:187–194CrossRefGoogle Scholar
  129. Ulyshen MD, Pawson S, Branco M, Horn S, Hoebeke ER, Gossner MM (2018) Utilization of non-native wood by saproxylic insects. In: Ulyshen MD (ed) Saproxylic insects: diversity, ecology and conservation. Springer, Heidelberg, pp 797–834CrossRefGoogle Scholar
  130. Urgenson LS, Halpern CB, Anderson PD et al (2013) Level and pattern of overstory retention influence rates and forms of tree mortality in mature, coniferous forests of the Pacific Northwest, USA. For Ecol Manag 308:116–127CrossRefGoogle Scholar
  131. Vanderwel MC, Malcolm JR, Smith SM et al (2006) Insect community composition and trophic guild structure in decaying logs from eastern Canadian pine-dominated forests. For Ecol Manag 225:190–199CrossRefGoogle Scholar
  132. Wardlaw T, Grove S, Hopkins A et al (2009) The uniqueness of habitats in old eucalypts: contrasting wood-decay fungi and saproxylic beetles of young and old eucalypts. Tasforests 18:17–32Google Scholar
  133. Wesley J (2002) The impacts of variable retention harvesting on phytophagous insects and their parasitoids in the boreal forest. MSc thesis, University of AlbertaGoogle Scholar
  134. Wood CM (2012) Saproxylic beetles (Coleoptera) associated with aspen deadwood in broad-leaved boreal mixedwood stands. MSc thesis, University of AlbertaGoogle Scholar
  135. Work TT, Spence JR, Volney WJA et al (2003) Integrating biodiversity and forestry practices in western Canada. For Chron 79:906–916CrossRefGoogle Scholar
  136. Work TT, Jacobs JM, Spence JR et al (2010) High levels of green-tree retention are required to preserve ground beetle biodiversity in boreal mixedwood forests. Ecol Appl 20:741–751PubMedCrossRefPubMedCentralGoogle Scholar
  137. Zielonka T (2006) When does dead wood turn into a substrate for spruce replacement? J Veg Sci 17:739–746CrossRefGoogle Scholar

Copyright information

© This is a U.S. government work and its text is not subject to copyright protection in the United States; however, its text may be subject to foreign copyright protection.  2018

Authors and Affiliations

  • Seung-Il Lee
    • 1
    Email author
  • John R. Spence
    • 1
  • David W. Langor
    • 2
  1. 1.Department of Renewable ResourcesUniversity of AlbertaEdmontonCanada
  2. 2.Natural Resources Canada, Canadian Forest Service, Northern Forestry CentreEdmontonCanada

Personalised recommendations