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Long-term responses of ground beetles (Coleoptera: Carabidae) to clear-cutting and wildfire in lodgepole pine stands of western Alberta, Canada

Published online by Cambridge University Press:  15 August 2022

Vincent Belluz
Affiliation:
Department of Renewable Resources, 751 General Services Building, University of Alberta, Edmonton, Alberta, T6G 2H1, Canada
David W. Langor*
Affiliation:
Natural Resources Canada, Canadian Forest Service, 5320-122 Street NW, Edmonton, Alberta, T6H 3S5, Canada
Jari K. Niemelä
Affiliation:
University of Helsinki, P.O. Box 3, FI-00014, Helsinki, Finland
Fangliang He
Affiliation:
Department of Renewable Resources, 751 General Services Building, University of Alberta, Edmonton, Alberta, T6G 2H1, Canada
John R. Spence
Affiliation:
Department of Renewable Resources, 751 General Services Building, University of Alberta, Edmonton, Alberta, T6G 2H1, Canada
*
*Corresponding author. Email: david.langor@nrcan-rncan.ca

Abstract

We studied responses of carabid beetles (Coleoptera: Carabidae) to clear-cutting and wildfire in lodgepole pine forests in the foothills of Alberta, Canada during 2013–2014 and compared the results with those from a similar study conducted in the same area during 1989–1990. Assemblages from stands regenerating 12–53 years after harvest gradually recovered towards their presumed preharvest condition represented by old pyrogenic stands. Assemblage structure in postharvest stands of similar age had also largely converged with that in stands that had burned in 1957 and 1997. Composition of ground vegetation, mineral soil cover, and basal area of trees and shrubs were significantly correlated with carabid assemblage structure, suggesting that plant successional gradients and patterns in carabid assemblages are driven by similar factors. We found that no carabid species was strictly associated with old pyrogenic stands, although assemblages in pyrogenic stands were distinctive. We predict that composition of carabid assemblages in harvested stands will recover and roughly match the variable structure of assemblages remaining in old, never-cut pyrogenic stands, given sufficient time (≥ 50 years). Nonetheless, the carabid fauna of the eastern slopes of Alberta’s Rocky Mountains appears to be changing in response to factors other than forestry. Warming climate is an explanation consistent with the changes observed.

Type
Research Paper
Copyright
© The Author(s) and Her Majesty, the Queen, in right of Canada, 2022. Published by Cambridge University Press on behalf of the Entomological Society of Canada

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Footnotes

deceased

Subject editor: Kevin Floate

References

Alberta Climate Information Service. 2018. Historical weather data, Alberta Climate Information Service [online]. Alberta Agriculture and Forestry, Edmonton, Alberta, Canada. Available from https://agriculture.alberta.ca/acis [accessed 1 March 2018].Google Scholar
Beckingham, J.D., Corns, I.G.W., and Archibald, J.H. 1996. Field guide to ecosites of west–central Alberta. Special report 9. Natural Resources Canada, Canadian Forest Service, Northwest Region, Northern Forestry Centre, Edmonton, Alberta, Canada.Google Scholar
Bedford, S.E. and Usher, M.B. 1994. Distribution of arthropod species across the margins of farm woodlands. Agriculture, Ecosystems and Environment, 48: 295305.CrossRefGoogle Scholar
Bell, A.J., Phillips, I.D., Floate, K.D., Hoemsen, B.M., and Phillips, C.E. 2014. Effects of pitfall lid transparency and habitat structure on the catches of carabid beetles (Coleoptera: Carabidae) in tame pasture. Environmental Entomology, 43: 139145.CrossRefGoogle Scholar
Bergeron, J.A.C., Spence, J.R., Volney, W.J.A., Pinzon, J., and Hartley, D.J. 2013. Effect of habitat type and pitfall trap installation on captures of epigaeic arthropod assemblages in the boreal forest. The Canadian Entomologist, 145: 547565. https://doi.org/10.4039/tce.2013.38.CrossRefGoogle Scholar
Blanchet, F.G., Bergeron, J.A.C., Spence, J.R., and He, F. 2013. Landscape effects of disturbance, habitat heterogeneity and spatial autocorrelation for a ground beetle (Carabidae) assemblage in mature boreal forest. Ecography, 36: 636647.CrossRefGoogle Scholar
Bott, R., Murphy, P., and Udell, R. 2003. Learning from the forest: a fifty-year journey in sustainable forest management. Fifth House Ltd., Calgary, Alberta, Canada. 242 pp.Google Scholar
Boulinier, T., Nichols, J.D., Sauer, J.R., Hines, J.E., and Pollock, K.H. 1998. Estimating species richness: the importance of heterogeneity in species detectability. Ecology, 79: 10181028.CrossRefGoogle Scholar
Bousquet, Y., Bouchard, P., Davies, A.E., and Sikes, D.S. 2013. Checklist of beetles (Coleoptera) of Canada and Alaska. Second edition. Pensoft, Sofia, Bulgaria–Moscow, Russia.Google ScholarPubMed
Brandmayr, P. and Pizzolotto, R. 2016. Climate change and its impact on epigean and hypogean carabid beetles. Periodicum Biologorum, 118: 147162 CrossRefGoogle Scholar
Bruce, D. 1955. A new way to look at trees. Journal of Forestry, 53: 163167.Google Scholar
Buddle, C.M., Beguin, J., Bolduc, E., Mercado, A., Sackett, T.E., Selby, R.D., et al. 2005. The importance and use of taxon sampling curves for comparative biodiversity research with forest arthropod assemblages. The Canadian Entomologist, 137: 120127. https://doi.org/10.4039/n04-040.CrossRefGoogle Scholar
Buddle, C.M., Langor, D.W., Pohl, G.R., and Spence, J.R. 2006. Arthropod responses to harvesting and wildfire: implications for emulation of natural disturbance in forest management. Biological Conservation, 128: 346357.CrossRefGoogle Scholar
Buddle, C.M., Spence, J.R., and Langor, D.W. 2000. Succession of boreal forest spider assemblages following wildfire and harvesting. Ecography, 23: 424436.CrossRefGoogle Scholar
Burton, P.J., Messier, C., Smith, D.W., and Adamowicz, W.L. (editors). 2003. Towards sustainable management of the boreal forest. NRC Research Press, Ottawa, Ontario, Canada. 1039 pp.Google Scholar
Byrne, J.M., Fagre, D.B., MacDonald, R., and Muhlfeld, C.C. 2014. Climate change and the Rocky Mountains. Chapter 20. In Impact of global changes on mountains: responses and adaptation. Edited by V. Grover, A. Borsdorf, J. Breuste, P. Tiwari, and F. Frangetto. CRC Press, Boca Raton, Louisiana, United States of America. Pp. 432–463.Google Scholar
Chao, A. and Jost, L. 2012. Coverage-based rarefaction and extrapolation: standardizing samples by completeness rather than size. Ecology, 93: 25332547.CrossRefGoogle ScholarPubMed
Cobb, T.P., Langor, D.W., and Spence, J.R. 2007. Biodiversity and multiple disturbances: boreal forest ground beetle (Coleoptera: Carabidae) responses to wildfire, harvesting, and herbicide. Canadian Journal of Forest Research, 37: 13101323.CrossRefGoogle Scholar
Coogan, S.C.P., Robinne, F.N., Jain, P., and Flannigan, M.D. 2019. Scientists’ warning on wildfire: a Canadian perspective. Canadian Journal of Forest Research, 49: 10151023.CrossRefGoogle Scholar
Corns, I.G. and La Roi, G.H.L. 1976. A comparison of mature with recently clear-cut and scarified lodgepole pine forests in the lower foothills of Alberta. Canadian Journal of Forest Research, 6: 2032.CrossRefGoogle Scholar
Digweed, S., Currie, C.R., Carcamo, H., and Spence, J.R. 1995. Digging out the “digging-in effect” of pitfall traps: confluences of depletion and disturbance on catches of ground beetles (Coleoptera: Carabidae). Pedobiologia, 39: 561576.Google Scholar
Everitt, B.S. 1992. The analysis of contingency tables. Second edition. Chapman and Hall, London, United Kingdom.CrossRefGoogle Scholar
Fahy, O. and Gormally, M. 1998. A comparison of plant and carabid communities in an Irish oak woodland with a nearby conifer plantation and clearfelled site. Forest Ecology and Management, 110: 263273.CrossRefGoogle Scholar
Flannigan, M.D., Krawchuk, M.A., Groot, W.J.D., Wotton, B.M., and Gowman, L.M. 2009. Implications of changing climate for global wildland fire. International Journal of Wildland Fire, 18: 483507.CrossRefGoogle Scholar
Fuller, R.J., Oliver, T.H., and Leather, S.R. 2008. Forest management effects on carabid beetle communities in coniferous and broadleaved forests: implications for conservation. Insect Conservation and Diversity, 1: 242252.CrossRefGoogle Scholar
Gandhi, K.J., Spence, J.R., Langor, D.W., and Morgantini, L.E. 2001. Fire residuals as habitat reserves for epigaeic beetles (Coleoptera: Carabidae and Staphylinidae). Biological Conservation, 102: 131141.CrossRefGoogle Scholar
Gandhi, K.J., Spence, J.R., Langor, D.W., Morgantini, L.E., and Cryer, K.J. 2004. Harvest retention patches are insufficient as stand analogues of fire residuals for litter-dwelling beetles in northern coniferous forests. Canadian Journal of Forest Research, 34: 13191331.CrossRefGoogle Scholar
Gobbi, M., Bernardi, F.D., Pelfini, M., Rossaro, B., and Brandmayr, P. 2006. Epigean arthropod succession along a 154-year glacier foreland chronosequence in the Forni Valley (Central Italian Alps). Arctic, Antarctic, and Alpine Research, 38: 357362.CrossRefGoogle Scholar
Gough, L., Shaver, G.R., Carroll, J., Royer, D.L., and Laundre, J.A. 2000. Vascular plant species richness in Alaskan Arctic tundra: the importance of soil pH. Journal of Ecology, 88: 5466.CrossRefGoogle Scholar
Hansen, A.J. and Phillips, L.B. 2015. Which tree species and biome types are most vulnerable to climate change in the US Northern Rocky Mountains? Forest Ecology and Management, 338: 6883.CrossRefGoogle Scholar
Heliölä, J., Koivula, M., and Niemelä, J. 2001. Distribution of carabid beetles (Coleoptera, Carabidae) across a boreal forest–clearcut ecotone. Conservation Biology, 15: 370377.CrossRefGoogle Scholar
Hillebrand, H., Blasius, B., Borer, E.T., Chase, J.M., Downing, J.A., Eriksson, B.K., et al. 2018. Biodiversity change is uncoupled from species richness trends: consequences for conservation and monitoring. Journal of Applied Ecology, 55: 169184.CrossRefGoogle Scholar
Holliday, N.J. 1991. Species responses of carabid beetles (Coleoptera: Carabidae) during post-fire regeneration of boreal forest. The Canadian Entomologist, 123: 13691389. https://doi.org/10.4039/Ent1231369-6.CrossRefGoogle Scholar
Holliday, N.J., Floate, K.D., Cárcamo, H., Pollock, D.A., Stjernberg, A., and Roughley, R.E. 2014. Ground beetles (Coleoptera: Carabidae) of the prairie grasslands of Canada. In Arthropods of Canadian grasslands. Volume 4: Biodiversity and Systematics. Part 2. Edited by D.J. Giberson and H.A. Cárcamo. Biological Survey of Canada, Canada. Pp. 1–85.Google Scholar
Honek, A. 1997. The effect of temperature on the activity of Carabidae (Coleoptera) in a fallow field. European Journal of Entomology, 94: 97104.Google Scholar
Hsieh, T.C., Ma, K.H., and Chao, A. 2016. iNEXT: an R package for rarefaction and extrapolation of species diversity (Hill numbers). Methods in Ecology and Evolution, 7: 14511456.CrossRefGoogle Scholar
Hunter, M.L. Jr. 1993. Natural disturbance regimes as spatial models for managing boreal forests. Biological Conservation, 65: 115120.CrossRefGoogle Scholar
Johansson, T., Hjältén, J., Olsson, J., Dynesius, M., and Roberge, J.M. 2016. Long-term effects of clear-cutting on epigaeic beetle assemblages in boreal forests. Forest Ecology and Management, 359: 6573.CrossRefGoogle Scholar
Jonsson, B.G. and Esseen, P.A. 1990. Treefall disturbance maintains high bryophyte diversity in a boreal spruce forest. The Journal of Ecology, 78: 924936.CrossRefGoogle Scholar
Jouveau, S., Toïgo, M., Giffard, B., Castagneyrol, B., Van Halder, I., Vétillard, F. and Jactel, H. 2020. Carabid activity-density increases with forest vegetation diversity at different spatial scales. Insect Conservation and Diversity, 13: 3646.CrossRefGoogle Scholar
Karen, M., O’Halloran, J., Breen, J., Giller, P., Pithon, J., and Kelly, T. 2008. Distribution and composition of carabid beetle (Coleoptera, Carabidae) communities across the plantation forest cycle: implications for management. Forest Ecology and Management, 256: 624632.CrossRefGoogle Scholar
Koivula, M., Cobb, T., Déchêne, A.D., Jacobs, J., and Spence, J.R. 2006. Responses of two Sericoda Kirby, 1837 (Coleoptera: Carabidae) species to forest harvesting, wildfire, and burn severity. Entomologica Fennica, 17: 315324.CrossRefGoogle Scholar
Koivula, M., Kukkonen, J., and Niemelä, J. 2002. Boreal carabid-beetle (Coleoptera, Carabidae) assemblages along the clear-cut originated succession gradient. Biodiversity and Conservation, 11: 12691288.CrossRefGoogle Scholar
Langor, D.W., Hammond, H.E.J., Spence, J.R., Jacobs, J.M., and Cobb, T.P. 2008. Saproxylic beetle assemblages in Canadian forests: diversity, ecology, and conservation. The Canadian Entomologist, 140: 453474. https://doi.org/10.4039/n07-LS02.CrossRefGoogle Scholar
Langor, D.W., Pohl, G.R., and Hammond, J.E.H. 2006. A coarse-filter approach to conserving arthropod biodiversity in Canadian forests. Arthropods of Canadian Forests Newsletter, 2: 913. Available from https://biologicalsurvey.ca/acf/forestarthropodsno2.pdf [accessed 10 March 2022].Google Scholar
Lee, T.D. and La Roi, G.H. 1979. Gradient analysis of bryophytes in Jasper National Park, Alberta. Canadian Journal of Botany, 57: 914925.CrossRefGoogle Scholar
Legendre, P. and Legendre, L. 2012. Numerical ecology. Third English edition. Elsevier, Amsterdam, The Netherlands. 990 pp.Google Scholar
Lemieux, J.P. and Lindgren, B.S. 2004. Ground beetle responses to patch retention harvesting in high-elevation forests of British Columbia. Ecography, 27: 557566.CrossRefGoogle Scholar
Lindroth, C.H. 1961. The ground beetles (Carabidae, excl. Cicindelinae) of Canada and Alaska. Part 2. Opuscula Entomologica Supplementum, 20: 1200.Google Scholar
Lindroth, C.H. 1963. The ground beetles (Carabidae, excl. Cicindelinae) of Canada and Alaska. Part 3. Opuscula Entomologica Supplementum, 24: 201408.Google Scholar
Lindroth, C.H. 1966. The ground beetles (Carabidae, excl. Cicindelinae) of Canada and Alaska. Part 4. Opuscula Entomologica Supplementum, 29: 408648.Google Scholar
Lindroth, C.H. 1968. The ground beetles (Carabidae, excl. Cicindelinae) of Canada and Alaska. Part 5. Opuscula Entomologica Supplementum, 33: 649944.Google Scholar
Lindroth, C.H. 1969a. The ground beetles (Carabidae, excl. Cicindelinae) of Canada and Alaska. Part 1. Opuscula Entomologica Supplementum, 35: IXLVIII.Google Scholar
Lindroth, C.H. 1969b. The ground beetles (Carabidae, excl. Cicindelinae) of Canada and Alaska. Part 6. Opuscula Entomologica Supplementum, 34: 9451192.Google Scholar
Magura, T., Ködöböcz, V., and Tóthmérész, B. 2001b. Effects of habitat fragmentation on carabids in forest patches. Journal of Biogeography, 28: 129138.CrossRefGoogle Scholar
Magura, T., Tóthmérész, B., and Molnar, T. 2001a. Edge effects on carabid assemblages along forest-grass transects. Web Ecology, 2: 713 CrossRefGoogle Scholar
Niemelä, J., Haila, Y., Halme, E., Pajunen, T., and Punttila, P. 1992. Small-scale heterogeneity in the spatial distribution of carabid beetles in the southern Finnish taiga. Journal of Biogeography, 19: 173181.CrossRefGoogle Scholar
Niemelä, J., Haila, Y., and Punttila, P. 1996. The importance of small-scale heterogeneity in boreal forests: variation in diversity in forest-floor invertebrates across the succession gradient. Ecography, 19: 352368.CrossRefGoogle Scholar
Niemelä, J., Koivula, M., and Kotze, D.J. 2007. The effects of forestry on carabid beetles (Coleoptera: Carabidae) in boreal forests. Journal of Insect Conservation, 11: 518.CrossRefGoogle Scholar
Niemelä, J., Langor, D., and Spence, J.R. 1993. Effects of clear-cut harvesting on boreal ground-beetle assemblages (Coleoptera: Carabidae) in western Canada. Conservation Biology, 7: 551561.CrossRefGoogle Scholar
Niemelä, J. and Spence, J.R. 1994. Distribution of forest dwelling carabids (Coleoptera): spatial scale and the concept of communities. Ecography, 17: 166175.CrossRefGoogle Scholar
Oksanen, J., Blanchet, F., Kindt, R., Legendre, P., Minchin, P.R., O’Hara, R.B., et al. 2013. Package ‘vegan’. R package version 2.5–2 [online]. Available from http://CRAN.R-project.org/package=vegan [accessed 1 August 2018].Google Scholar
Osawa, N., Terai, A., Hirata, K., Nakanishi, A., Makino, A., Sakai, S., and Sibata, S. 2004. Logging impacts on forest carabid assemblages in Japan. Canadian Journal of Forest Research, 35: 26982708.CrossRefGoogle Scholar
Pearce, J.L. and Venier, L.A. 2006. The use of ground beetles (Coleoptera: Carabidae) and spiders (Araneae) as bioindicators of sustainable forest management: a review. Ecological Indicators, 6: 780793.CrossRefGoogle Scholar
Pechony, O. and Shindell, D.T. 2010. Driving forces of global wildfires over the past millennium and the forthcoming century. Proceedings of the National Academy of Science, 107: 1916719170.CrossRefGoogle ScholarPubMed
Phillips, I.D., Cobb, T.P., Spence, J.R., and Brigham, R.M. 2006. Salvage logging, edge effects, and carabid beetles: connections to conservation and sustainable forest management. Environmental Entomology, 35: 950957.CrossRefGoogle Scholar
Pinzon, J., Wu, L., He, F., and Spence, J.R. 2018. Fine-scale forest variability and biodiversity in the boreal mixedwood forest. Ecography, 41: 753769.CrossRefGoogle Scholar
Pohl, G.R., Langor, D.W., and Spence, J.R. 2007. Rove beetles and ground beetles (Coleoptera: Staphylinidae, Carabidae) as indicators of harvest and regeneration practices in western Canadian foothills forests. Biological Conservation, 137: 294307.CrossRefGoogle Scholar
Pozsgai, G., Baird, J., Littlewood, N.A., Pakeman, R.J., and Young, M.R. 2018. Phenological changes of the most commonly sampled ground beetle (Coleoptera: Carabidae) species in the UK environmental change network. International Journal of Biometeorology, 62: 10631074.CrossRefGoogle ScholarPubMed
R Core Team. 2016. R: a language and environment for statistical computing [online.] R Foundation for Statistical Computing, Vienna, Austria. Available from https://www.R-project.org/ [accessed 1 September 2018].Google Scholar
Rainio, J. and Niemelä, J. 2003. Ground beetles (Coleoptera: Carabidae) as bioindicators. Biodiversity and Conservation, 12: 487506.CrossRefGoogle Scholar
Royer, F. and Dickinson, R. 2007. Plants of Alberta: trees, shrubs, wildflowers, ferns, aquatic plants and grasses. Lone Pine Publishing, Edmonton, Alberta, Canada. 392 pp.Google Scholar
Saint-Germain, M., Larrivée, M., Drapeau, P., Fahrig, L., and Buddle, C.M. 2005. Short-term response of ground beetles (Coleoptera: Carabidae) to fire and logging in a spruce-dominated boreal landscape. Forest Ecology and Management, 212: 118126.CrossRefGoogle Scholar
Saska, P. and Honek, A. 2003. Temperature and development of central European species of Amara (Coleoptera: Carabidae). European Journal of Entomology, 100: 509516.CrossRefGoogle Scholar
Spence, J.R. 2001. The new boreal forestry: adjusting timber management to accommodate biodiversity. Trends in Ecology and Evolution, 16: 591593.CrossRefGoogle Scholar
Spence, J.R., Langor, D.W., Niemelä, J., Carcamo, H.A., and Currie, C.R. 1996. Northern forestry and carabids: the case for concern about old-growth species. Annales Zoologici Fennici, 33: 173184.Google Scholar
Spence, J.R. and Niemelä, J.K. 1994. Sampling carabid assemblages with pitfall traps: the madness and the method. The Canadian Entomologist, 126: 881894. https://doi.org/10.4039/Ent126881-3.CrossRefGoogle Scholar
Strickler, G.S. 1959. Use of the densiometer to estimate density of forest canopy on permanent sample plots. PNW Old Series Research Notes, 180: 15.Google Scholar
Tews, J., Brose, U., Grimm, V., Tielbörger, K., Wichmann, M.C., Schwager, M., and Jeltsch, F. 2004. Animal species diversity driven by habitat heterogeneity/diversity: the importance of keystone structures. Journal of Biogeography, 31: 7992.CrossRefGoogle Scholar
Tseng, M., Kaur, K.M., Soleimani Pari, S., Sarai, K., Chan, D., Yao, C.H., et al. 2018. Decreases in beetle body size linked to climate change and warming temperatures. Journal of Animal Ecology, 87: 647659.CrossRefGoogle ScholarPubMed
Turin, H. and den Boer, P.J. 1988. Changes in the distribution of carabid beetles in the Netherlands since 1880. II. Isolation of habitats and long-term time trends in the occurrence of carabid species with different powers of dispersal (Coleoptera, Carabidae). Biological Conservation, 44: 179200.CrossRefGoogle Scholar
Turin, H., Kotze, D.J., Müller-Kroehling, S., Saska, P., Spence, J.R., and Heijerman, T. 2022. Ecology and conservation of the Dutch ground beetle fauna: lessons from 66 years of pitfall trapping. Wageningen Academic Publishers, Wageningen, The Netherlands. 452 pp.CrossRefGoogle Scholar
Udell, R., Murphy, P.J., Bott, R., and Stevenson, R.E. 2002. The Hinton Forest 1955–2000: a case study in adaptive forest management. Foothills Model Forest History Series Report [online]. Available from https://friresearch.ca/sites/default/files/FHP_2014_05_Book_HintonForestOpt.pdf [accessed 1 September 2018].Google Scholar
Van Wagner, C.E., Finney, M.A., and Heathcott, M. 2006. Historical fire cycles in the Canadian Rocky Mountain parks. Forest Science, 52: 704717.Google Scholar
Venier, L.A., Work, T.T., Klimaszewski, J., Morris, D.M., Bowden, J.J., Kwiaton, M.M., et al. 2017. Ground-dwelling arthropod response to fire and clearcutting in jack pine: implications for ecosystem management. Canadian Journal of Forest Research, 47: 16141631.CrossRefGoogle Scholar
Venn, S. 2016. To fly or not to fly. European Journal of Entomology, 113: 587600.CrossRefGoogle Scholar
Wikars, L.O. and Schimmel, J. 2001. Immediate effects of fire severity on soil invertebrates in cut and uncut pine forests. Forest Ecology and Management, 141: 189200.CrossRefGoogle Scholar
Work, T.T., Jacobs, J.M., Spence, J.R., and Volney, W.J. 2010. High levels of green-tree retention are required to preserve ground beetle biodiversity in boreal mixedwood forests. Ecological Applications, 20: 741751.CrossRefGoogle ScholarPubMed
Work, T., Klimaszewski, J., Thiffault, E., Bourdon, C., Pare, D., Bousquet, Y., et al. 2013. Initial responses of rove and ground beetles (Coleoptera: Staphylinidae, Carabidae) to removal of logging residues following clearcut harvesting in the boreal forest of Quebec, Canada. ZooKeys, 258: 3152.Google Scholar
Work, T.T., Koivula, M., Klimaszewski, J., Langor, D., Spence, J., Sweeney, J., and Hébert, C. 2008. Evaluation of carabid beetles as indicators of forest change in Canada. The Canadian Entomologist, 140: 393414. https://doi.org/10.4039/n07-LS07.CrossRefGoogle Scholar
Work, T.T., Shorthouse, D.P., Spence, J.R., Volney, W.J.A., and Langor, D. 2004. Stand composition and structure of the boreal mixedwood and epigaeic arthropods of the Ecosystem Management Emulating Natural Disturbance (EMEND) landbase in northwestern Alberta. Canadian Journal of Forest Research, 34: 417430.CrossRefGoogle Scholar
Zou, Y., Sang, W., Wang, S., Warren-Thomas, E., Liu, Y., Yu, Z., et al. 2015. Diversity patterns of ground beetles and understorey vegetation in mature, secondary, and plantation forest regions of temperate northern China. Ecology and Evolution, 5: 531542.CrossRefGoogle Scholar
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