Boreal mixedwood understory communities are highly dynamic because they are simultaneously influenced by canopy succession, small-scale environmental variation and disturbances. In such a system, fire initiates succession but canopy gaps, resulting from spruce budworm outbreaks or other disturbances, also affect forest dynamics during succession. In this study, we assessed the short-term response of understory communities along a canopy gap severity gradient in mature and old-growth forest stands. The objective was to determine if resource heterogeneity is driving understory community composition and diversity. We created artificial gaps of varying severities in stands of two late successional forest stages. Four treatments, i.e., total cut, conifer cut, girdling of conifers and a control treatment, were each applied over an area of 100 m2 and replicated three times in both types of forest stands. The cover of each understory species was measured before treatment application and 1, 2 and 11 years following it, in seven 1 m2 plots per replicate. Percent transmission of photosynthetic photon flux density and soil temperature were also measured on different occasions after the beginning of the experiment. Species richness, alpha diversity and total cover all increased following gap creation, and this increase was sustained up to 11 years. Increases were more pronounced in total cut than in other treatments. The increase in species richness and diversity was also more pronounced in old-growth stands, while total cover increase was greater in mature stands. Gap creation resulted in significant changes in abundance patterns that were associated with the resident species assemblages. The magnitude of changes in community composition did not significantly differ between mature and old-growth forests and differences in species composition among stands were maintained following gap creation. This study supports the evidence that resource heterogeneity resulting from small-scale disturbances will contribute to maintaining diversity among understory communities by allowing species more typical of early-successional communities to coexist with species from late-successional ones. Such a process may be essential for maintaining the species pool likely to re-invade the stands following major disturbances.
Bartels, S. and H. Chen. 2010. Is understory plant species diversity driven by resource quantity or resource heterogeneity? Ecology 91:1931–1938.
Chen H. , 'Is understory plant species diversity driven by resource quantity or resource heterogeneity? ' (2010 ) 91 Ecology : 1931 -1938 .
Bergeron, Y. 1991. The influence of island and mainland lakeshore landscapes on boreal forest fire regimes. Ecology 72:1980–1992.
Bergeron Y. , 'The influence of island and mainland lakeshore landscapes on boreal forest fire regimes ' (1991 ) 72 Ecology : 1980 -1992 .
Bergeron, Y. and D. Charron. 1994. Postfire stand dynamics in a southern boreal forest (Québec): a dendroecological approach. Ecoscience 1:173–184.
Charron D. , 'Postfire stand dynamics in a southern boreal forest (Québec): a dendroecological approach ' (1994 ) 1 Ecoscience : 173 -184 .
Bergeron, Y. and M. Dubuc. 1989. Succession in the southern part of the Canadian boreal forest. Vegetatio 79:51–63.
Dubuc M. , 'Succession in the southern part of the Canadian boreal forest ' (1989 ) 79 Vegetatio : 51 -63 .
Bergeron, Y., A. Leduc, H. Morin and C. Joyal. 1995. Balsam fir mortality following the last spruce budworm outbreak in northwestern Quebec. Can. J. Forest Res. 25:1375–1384.
Joyal C. , 'Balsam fir mortality following the last spruce budworm outbreak in northwestern Quebec ' (1995 ) 25 Can. J. Forest Res. : 1375 -1384 .
Bonan, G.B. and H.H. Shugart. 1989. Environmental factors and ecological processes in boreal forests. Ann. Rev. Ecol. Syst. 20:1–28.
Shugart H.H. , 'Environmental factors and ecological processes in boreal forests ' (1989 ) 20 Ann. Rev. Ecol. Syst. : 1 -28 .
Carleton, T.J. and P.F. Maycock. 1981. Understorey-canopy affinities in boreal forest vegetation. Can. J.Bot. 59:1709–1716.
Maycock P.F. , 'Understorey-canopy affinities in boreal forest vegetation ' (1981 ) 59 Can. J.Bot. : 1709 -1716 .
Carleton, T.J. and P.F. Maycock. 1980. Vegetation of the boreal forests south of James Bay: non-centered component analysis of the vascular flora. Ecology 61:1199–1212.
Maycock P.F. , 'Vegetation of the boreal forests south of James Bay: non-centered component analysis of the vascular flora ' (1980 ) 61 Ecology : 1199 -1212 .
Côté, L., S. Brown, D. Paré, J.W. Fyles and J. Bauhus. 2000. Dynamics of carbon and nitrogen mineralization in relation to stand type, stand age, and soil texture in the boreal mixedwood. Soil Biol. Biochem. 32:1079–1090.
Bauhus J. , 'Dynamics of carbon and nitrogen mineralization in relation to stand type, stand age, and soil texture in the boreal mixedwood ' (2000 ) 32 Soil Biol. Biochem. : 1079 -1090 .
Dansereau, P.-R. and Y. Bergeron. 1993. Fire history in the southern boreal forest of northwestern Quebec. Can. J. Forest Res. 23:25–32.
Bergeron Y. , 'Fire history in the southern boreal forest of northwestern Quebec ' (1993 ) 23 Can. J. Forest Res. : 25 -32 .
De Grandpré, L. and Y. Bergeron. 1997. Diversity and stability of understorey communities following disturbance in the southern boreal forest. J. Ecol. 85:777–784.
Bergeron Y. , 'Diversity and stability of understorey communities following disturbance in the southern boreal forest ' (1997 ) 85 J. Ecol. : 777 -784 .
De Grandpé, L., D. Gagnon and Y. Bergeron. 1993. Changes in the understory of Canadian southern boreal forest after fire. J. Veg. Sci. 4:803–810.
Bergeron Y. , 'Changes in the understory of Canadian southern boreal forest after fire ' (1993 ) 4 J. Veg. Sci. : 803 -810 .
Environment Canada. 2005. Canadian Climate Normals or Averages 1971–2000. URL: www.climate.weatheroffice.ec.gc.ca/climate.normals/index_e.html
Eriksson, O. and L. Jerling. 1990. Hierarchical selection and risk spreading in clonal plants. In: J. van Groenendael and H. de Kroon (eds.), Clonal Growth in Plants. SPB Academic Publishing, The Hague. pp. 79–94.
Jerling L. , '', in Clonal Growth in Plants , (1990 ) -.
Frelich, L.E. and P.B. Reich. 1999. Neighborhood effects, disturbance severity, and community stability in forests. Ecosystems 2:151–166.
Reich P.B. , 'Neighborhood effects, disturbance severity, and community stability in forests ' (1999 ) 2 Ecosystems : 151 -166 .
Gilliam, F.S. and M.R. Roberts. 2003. The Herbaceous Layer in Forests of Eastern North America. Oxford Univ. Press, New York.
Roberts M.R. , '', in The Herbaceous Layer in Forests of Eastern North America , (2003 ) -.
Halpern, C.B. 1989. Early successional patterns of forest species: interactions of life history traits and disturbance. Ecology 70:704–720.
Halpern C.B. , 'Early successional patterns of forest species: interactions of life history traits and disturbance ' (1989 ) 70 Ecology : 704 -720 .
Hart, S.A. and H.Y.H. Chen. 2008. Fire, logging, and overstory affect understory abundance, diversity, and composition in boreal forest. Ecol. Monogr. 78:123–140.
Chen H.Y.H. , 'Fire, logging, and overstory affect understory abundance, diversity, and composition in boreal forest ' (2008 ) 78 Ecol. Monogr. : 123 -140 .
Hart, S.A. and H.Y.H. Chen. 2006. Understory vegetation dynamics of North American boreal forests. Crit. Rev. Plant Sci. 25:381–397.
Chen H.Y.H. , 'Understory vegetation dynamics of North American boreal forests ' (2006 ) 25 Crit. Rev. Plant Sci. : 381 -397 .
Hugues, J.W. and T.J. Fahey. 1991. Colonization dynamics of herbs and shrubs in a disturbed northern hardwood forest. J. Ecol. 79:605–616.
Fahey T.J. , 'Colonization dynamics of herbs and shrubs in a disturbed northern hardwood forest ' (1991 ) 79 J. Ecol. : 605 -616 .
Hutchings, M.J. and M. Mogie. 1990. The spatial structure of clonal plants control and consequences. In: J. van Groenendael and H. de Kroon (eds.), Clonal Growth in Plants. SPB Academic Publishing, The Hague. pp. 57–76.
Mogie M. , '', in Clonal Growth in Plants , (1990 ) -.
Jonsson, B.G. and P.-A. Esseen. 1990. Treefall disturbance maintains high bryophyte diversity in a boreal spruce forest. J. Ecol. 78:924–936.
Esseen P.-A. , 'Treefall disturbance maintains high bryophyte diversity in a boreal spruce forest ' (1990 ) 78 J. Ecol. : 924 -936 .
Jost, L. 2007. Partitioning diversity into independent alpha and beta components. Ecology 88:2427–2439.
Jost L. , 'Partitioning diversity into independent alpha and beta components ' (2007 ) 88 Ecology : 2427 -2439 .
Kemball, K.J., G.G. Wang and Q.-L. Dang. 2005. Response of understory plant community of boreal mixedwood stands to fire, logging, and spruce budworm outbreak. Can. J. Bot. 83:1550–1560.
Dang Q.-L. , 'Response of understory plant community of boreal mixedwood stands to fire, logging, and spruce budworm outbreak ' (2005 ) 83 Can. J. Bot. : 1550 -1560 .
Kenward, M. G. and J.H. Roger. 1997. Small sample inference for fixed effects from restricted maximum likelihood. Biometrics 53:983–997.
Roger J.H. , 'Small sample inference for fixed effects from restricted maximum likelihood ' (1997 ) 53 Biometrics : 983 -997 .
Kneeshaw, D.D. and Y. Bergeron. 1998. Canopy gap characteristics and tree replacement in the southeastern boreal forest. Ecology 79:783–794.
Bergeron Y. , 'Canopy gap characteristics and tree replacement in the southeastern boreal forest ' (1998 ) 79 Ecology : 783 -794 .
Kneeshaw, D., Y. Bergeron and L. De Grandpré. 1998. Early response of Abies balsamea seedlings to artificially created openings. J. Veg. Sci. 9:543–550.
Grandpré L. , 'Early response of Abies balsamea seedlings to artificially created openings ' (1998 ) 9 J. Veg. Sci. : 543 -550 .
Légaré, S., Y. Bergeron and D. Paré. 2002. Influence of forest composition on understory cover of boreal mixedwood forests of western Quebec, Canada. Silva Fenn. 36:353–366.
Paré D. , 'Influence of forest composition on understory cover of boreal mixedwood forests of western Quebec, Canada ' (2002 ) 36 Silva Fenn. : 353 -366 .
Legendre, P. and L. Legendre. 1998. Numerical Ecology, Second English Edition. Elsevier, Amsterdam.
Legendre L. , '', in Numerical Ecology, Second English Edition , (1998 ) -.
Morin, H., D. Laprise, and Y. Bergeron. 1993. Chronology of spruce budworm outbreaks near Lake Duparquet, Abitibi region, Quebec. Can. J. Forest Res. 23:1497–1506.
Bergeron Y. , 'Chronology of spruce budworm outbreaks near Lake Duparquet, Abitibi region, Quebec ' (1993 ) 23 Can. J. Forest Res. : 1497 -1506 .
Noble, I.R. and R.O. Slatyer. 1980. The use of vital attributes to predict successional changes in plant communities subject to recurrent disturbances. Vegetatio 43:5–21.
Slatyer R.O. , 'The use of vital attributes to predict successional changes in plant communities subject to recurrent disturbances ' (1980 ) 43 Vegetatio : 5 -21 .
Økland, R.H. 2000. Understorey vegetation development in North Finnish Picea forests after disturbance: re-analysis of Sirén’s data. J. Veg. Sci. 11:533–546.
Økland R.H. , 'Understorey vegetation development in North Finnish Picea forests after disturbance: re-analysis of Sirén’s data ' (2000 ) 11 J. Veg. Sci. : 533 -546 .
Økland, R.H. and O. Eilertsen. 1996. Dynamics of understory vegetation in an old-growth boreal coniferous forest, 1988–1993. J. Veg. Sci. 7:747–762.
Eilertsen O. , 'Dynamics of understory vegetation in an old-growth boreal coniferous forest, 1988–1993 ' (1996 ) 7 J. Veg. Sci. : 747 -762 .
Økland, R.H., K. Rydgren and T. Økland. 2003. Plant species composition of boreal spruce swamp forests: closed doors and windows of opportunity. Ecology 84:1909–1919.
Økland T. , 'Plant species composition of boreal spruce swamp forests: closed doors and windows of opportunity ' (2003 ) 84 Ecology : 1909 -1919 .
Oksanen, J., F. Guillaume Blanchet, R. Kindt, P. Legendre, R.B. O’Hara, G.L. Simpson, P. Solymos, M.H.H. Stevens and H. Wagner 2010. vegan: Community Ecology Package. R package, version 1.17-4. http://CRAN.R-project.org/package=vegan
Paré, D. and Y. Bergeron. 1996. Effect of colonizing tree species on soil nutrient availability in a clay soil of the boreal mixedwood. Can. J. Forest Res. 26:1022–1031.
Bergeron Y. , 'Effect of colonizing tree species on soil nutrient availability in a clay soil of the boreal mixedwood ' (1996 ) 26 Can. J. Forest Res. : 1022 -1031 .
Parent, S. and C. Messier. 1996. A simple and efficient method to estimate microsite light availability under a forest canopy. Can. J. Forest Res. 26:151–154.
Messier C. , 'A simple and efficient method to estimate microsite light availability under a forest canopy ' (1996 ) 26 Can. J. Forest Res. : 151 -154 .
Rees, D.C. and G.P. Juday. 2002. Plant species diversity on logged versus burned sites in central Alaska. Forest Ecol. Manage. 155:291–302.
Juday G.P. , 'Plant species diversity on logged versus burned sites in central Alaska ' (2002 ) 155 Forest Ecol. Manage. : 291 -302 .
Reyes, G.P., D. Kneeshaw, L. De Grandpré and A. Leduc. 2010. Changes in woody vegetation abundance and diversity after natural disturbances causing different levels of mortality. J. Veg. Sci. 21:406–417.
Leduc A. , 'Changes in woody vegetation abundance and diversity after natural disturbances causing different levels of mortality ' (2010 ) 21 J. Veg. Sci. : 406 -417 .
Rich, R.L., L.E. Frelich and P.B. Reich. 2007. Wind-throw mortality in the southern boreal forest: effects of species, diameter and stand age. J. Ecol. 95:1261–1273.
Reich P.B. , 'Wind-throw mortality in the southern boreal forest: effects of species, diameter and stand age ' (2007 ) 95 J. Ecol. : 1261 -1273 .
Roberts, M.R. 2004. Response of the herbaceous layer to natural disturbance in North American forests. Can. J. Bot. 82:1273–1283.
Roberts M.R. , 'Response of the herbaceous layer to natural disturbance in North American forests ' (2004 ) 82 Can. J. Bot. : 1273 -1283 .
Rowe, J.S. 1983. Concepts of fire effects on plant individuals and species. In: R.W. Wein and D.A. MacLean (eds.), The Role of Fire in Northern Circumpolar Ecosystems. Scope, Vol. 18. Wiley, New York. pp. 135–144.
Rowe J.S. , '', in The Role of Fire in Northern Circumpolar Ecosystems. Scope, Vol. 18 , (1983 ) -.
Rydgren, K., G. Hestmark and R.H. Økland. 1998. Revegetation following experimental disturbance in a boreal old-growth Picea abies forest. J. Veg. Sci. 9:763–776.
Økland R.H. , 'Revegetation following experimental disturbance in a boreal old-growth Picea abies forest ' (1998 ) 9 J. Veg. Sci. : 763 -776 .
Rydgren, K., R.H. Økland and G. Hestmark. 2004. Disturbance severity and community resilience in a boreal forest. Ecology 85:1906–1915.
Hestmark G. , 'Disturbance severity and community resilience in a boreal forest ' (2004 ) 85 Ecology : 1906 -1915 .
SAS Institute Inc. 2002. SAS/STAT® User’s Guide, Version 9.1. SAS Institute Inc., Cary, NC, USA.
'', in SAS/STAT® User’s Guide, Version 9.1 , (2002 ) -.
Saucier, J.-P., J.-F. Bergeron, P. Grondin and A. Robitaille. 1998. Les régions écologiques du Québec méridional (3e version): un des éléments du système hiérarchique de classification écologique du territoire mis au point par le ministère des Ressources naturelles du Québec. L’Aubelle 124:S1–S12.
Robitaille A. , 'Les régions écologiques du Québec méridional (3e version): un des éléments du système hiérarchique de classification écologique du territoire mis au point par le ministère des Ressources naturelles du Québec ' (1998 ) 124 L’Aubelle : S1 -S12 .
Schwarz, P.A., T.J. Fahey and C.E. McCulloch. 2003. Factors controlling spatial variation of tree species abundance in a forested landscape. Ecology 84:1862–1878.
McCulloch C.E. , 'Factors controlling spatial variation of tree species abundance in a forested landscape ' (2003 ) 84 Ecology : 1862 -1878 .
Takafumi, H. and T. Hiura. 2009. Effects of disturbance history and environmental factors on the diversity and productivity of understory vegetation in a cool-temperate forest in Japan. Forest Ecol. Manage. 257:843–857.
Hiura T. , 'Effects of disturbance history and environmental factors on the diversity and productivity of understory vegetation in a cool-temperate forest in Japan ' (2009 ) 257 Forest Ecol. Manage. : 843 -857 .
Taylor, S.J., T.J. Carleton and P. Adams. 1987. Understorey vegetation change in a Picea mariana chronosequence. Vegetatio 73:63–72.
Adams P. , 'Understorey vegetation change in a Picea mariana chronosequence ' (1987 ) 73 Vegetatio : 63 -72 .
Tessier, J.T. 2007. Re-establishment of three dominant herbaceous understory species following fine-scale disturbance in a Catskill northern hardwood forest. J. Torrey Bot. Soc. 134:34–44.
Tessier J.T. , 'Re-establishment of three dominant herbaceous understory species following fine-scale disturbance in a Catskill northern hardwood forest ' (2007 ) 134 J. Torrey Bot. Soc. : 34 -44 .
Thomas, S.C., C.B. Halpern, D.A. Falk, D.A. Liguori and K.A. Austin. 1999. Plant diversity in managed forests: understory responses to thinning and fertilization. Ecol. Appl. 9:864–879.
Austin K.A. , 'Plant diversity in managed forests: understory responses to thinning and fertilization ' (1999 ) 9 Ecol. Appl. : 864 -879 .
Veillette, J.J. 1994. Evolution and paleohydrology of glacial Lakes Barlow and Ojibway. Quaternary Sci. Rev. 13: 945–971.
Veillette J.J. , 'Evolution and paleohydrology of glacial Lakes Barlow and Ojibway ' (1994 ) 13 Quaternary Sci. Rev. : 945 -971 .