Open lichen woodlands (LWs) are degraded stands that lack the ability to regenerate naturally due to a succession of natural and/or anthropogenic disturbances. As they represent both interesting forest restoration and carbon sequestration opportunities, we tested disc scarification and planting of two sizes of containerized black spruce ( Picea mariana Mill. (BSP)) seedlings for their afforestation. We compared treatment of unproductive LWs to reforestation of harvested, closed-crown black spruce-feathermoss (BSFM) stands. After one year, seedling survival and nutritional status were equivalent among stand types but despite higher root elongation index ( REI), planted seedlings in LWs had lower relative growth rate, smaller total biomass and stem diameter than those in BSFM stands. Soil fertility variables, soil temperature, nor seedling water potential, helped at explaining this early growth response. Disc scarification significantly improved seedling first-year survival, biomass and foliar nutrient concentrations of P, Ca, and Mg. Smaller planting stock showed higher REI, higher shoot water potential, and higher foliar nutrient concentration of all but one of the measured nutrients (N, P, K and Mg). Hence, preliminary results suggest that planting of smaller containerized black spruce stock, combined with disc scarification, shows potential for afforestation of unproductive LWs. The impact of the lichen mat and other potential growth limiting factors on afforestation of these sites requires further investigation.
References
[1]
Coulombe, G.; Huot, J.; Arsenault, J.; Bauce, é.; Bernard, J.T.; Bouchard, A.; Liboiron, M.A.; Szaraz, G. Commission D’étude sur la Gestion de la Forêt Publique Québécoise; Bibliothèque Nationale du Québec: Québec, Canada, 2004; p. 307.
[2]
Arseneault, D.; Payette, S. A postfire shift from lichen-spruce to lichen-tundra vegetation at tree line. Ecology 1992, 73, 1067–1081, doi:10.2307/1940181.
[3]
Payette, S.; Bhiry, N.; Delwaide, A.; Simard, M. Origin of the lichen woodland at its southern range limit in eastern Canada: The catastrophic impact of insect defoliators and fire on the spruce-moss forest. Can. J. For. Res. 2000, 30, 288–305, doi:10.1139/x99-207.
[4]
Gagnon, R.; Morin, H. Les forêts d’épinette noire du Québec: Dynamique, perturbations et biodiversité. Nat. Can. 2001, 125, 26–35.
[5]
Girard, F.; Payette, S.; Gagnon, R. Rapid expansion of lichen woodlands within the closed-crown boreal forest zone over the last 50 years caused by stand disturbances in eastern Canada. Can. J. Biogeogr. 2008, 35, 529–537, doi:10.1111/j.1365-2699.2007.01816.x.
[6]
Jasinski, J.P.P.; Payette, S. The creation of alternative stable states in the southern boreal forest, Québec, Canada. Ecol. Monogr. 2005, 75, 561–583, doi:10.1890/04-1621.
[7]
Hébert, F.; Boucher, J.F.; Bernier, P.Y.; Lord, D. Growth response and water relations of 3-year-old planted black spruce and jack pine seedlings in site prepared lichen woodlands. For. Ecol. Manag. 2006, 223, 226–236, doi:10.1016/j.foreco.2005.11.005.
[8]
C？té, D.; Girard, F.; Hébert, F.; Bouchard, S.; Gagnon, R.; Lord, D. Is the closed-crown boreal forest resilient after successive stand disturbances? A quantitative demonstration from a case study. J. Veg. Sci. 2013, 24, 664–674, doi:10.1111/j.1654-1103.2012.01488.x.
[9]
Gaboury, S.; Boucher, J.F.; Villeneuve, C.; Lord, D.; Gagnon, R. Estimating the net carbon balance of boreal open woodland afforestation: A case study in Québec’s closed-crown boreal forest. For. Ecol. Manag. 2009, 257, 483–494, doi:10.1016/j.foreco.2008.09.037.
[10]
Mallik, A.U. Ecology of a forest weed of Newfoundland: Vegetative regeneration strategy of Kalmia angustifolia. Can. J. Bot. 1993, 71, 161–166, doi:10.1139/b93-018.
[11]
Mallik, A.U. Growth and physiological responses of black spruce (Picea mariana) to sites dominated by Ledum groenlandicum. J. Chem. Ecol. 1996, 22, 575–585, doi:10.1007/BF02033656.
[12]
Yamasaki, S.H.; Fyles, J.W.; Egger, K.N.; Titus, B.D. The effect of Kalmia angustifolia on the growth, nutrition, and ectomycorrhizal symbiont community of black spruce. For. Ecol. Manag. 1998, 105, 197–207, doi:10.1016/S0378-1127(97)00285-5.
[13]
Thiffault, N.; Cyr, G.; Prégent, G.; Jobidon, R.; Charrette, L. Régénération artificielle des pessières noires à éricacées: Effets du scarifiage, de la fertilisation et du type de plants après 10 ans. For. Chron. 2004, 80, 141–149.
[14]
Thiffault, N.; Jobidon, R. How to shift unproductive Kalmia angustifolia–Rhododendron groenlandicum heath to productive conifer plantation. Can. J. For. Res. 2006, 36, 2364–2376, doi:10.1139/x06-090.
[15]
LeBel, P.; Thiffault, N.; Bradley, R. Kalmia removal increases nutrient supply and growth of black spruce seedlings: An effect fertilizer cannot emulate. For. Ecol. Manag. 2008, 256, 1780–1784, doi:10.1016/j.foreco.2008.02.050.
[16]
Moroni, M.T.; Thiffault, N.; Titus, B.D.; Mante, C.; Makeschin, F. Controlling Kalmia and reestablishing conifer dominance enhances soil fertility indicators in central Newfoundland, Canada. Can. J. For. Res. 2009, 39, 1270–1279, doi:10.1139/X09-061.
[17]
Steijlen, I.; Nilsson, M.C.; Zackrisson, O. Seed regeneration of scots pine in boreal forest stands dominated by lichen and feather moss. Can. J. For. Res. 1995, 25, 713–723, doi:10.1139/x95-079.
[18]
Fisher, R.F. Possible allelopathic effects of reindeer-moss (Cladonia) on jack pine and white spruce. For. Sci. 1979, 25, 256–260.
[19]
Houle, G.; Filion, L. The effects of lichens on white spruce seedling establishment and juvenile growth in a spruce-lichen woodland of subarctic Quebec. Ecoscience 2003, 10, 80–84.
[20]
Burdett, A.N. Physiological processes in plantation establishment and the development of specifications for forest planting stock. Can. J. For. Res. 1990, 20, 415–427, doi:10.1139/x90-059.
[21]
Bernier, P.Y. Comparing natural and planted black spruce seedlings 1. Water relations and growth. Can. J. For. Res. 1993, 23, 2427–2434, doi:10.1139/x93-300.
[22]
Girard, F. Remise en Production des Pessières à Lichens de la Forêt Boréale Commerciale: Nutrition et Croissance de Plants D’épinette Noire Trois Ans Après Traitements de Préparation de Terrain; Université du Québec à Chicoutimi: Chicoutimi, Canada, 2004.
[23]
Jobidon, R. Light threshold for optimal black spruce (Picea mariana) seedling growth and development under brush competition. Can. J. For. Res. 1994, 24, 1629–1635, doi:10.1139/x94-211.
[24]
Lamhamedi, M.; Bernier, P.Y.; Hébert, C. Effect of shoot size on the gas exchange and growth of containerized Picea mariana seedlings under different watering regimes. New For. 1997, 13, 209–223, doi:10.1023/A:1006586325524.
[25]
Nabuurs, G.J.; Masera, O.; Andrasko, K.; Benitez-Ponce, P.; Boer, R.; Dutschke, M.; Elsiddig, E.; Ford-Robertson, J.; Frumhoff, P.; Karjalainen, T.; et al. Forestry. In Climate Change 2007: Mitigation. Contribution of Working Group III to the Fourth Assessment Report of the Intergovernmental Panel on Climate Change; Cambridge University Press: Cambridge, UK, 2007; pp. 541–584.
[26]
Peters-Stanley, M.; Hamilton, K.; Marcello, T.; Sjardin, M. Back to the Future: State of the Voluntary Carbon Markets 2011; Ecosystem Marketplace: Washington, DC, USA, 2011; p. 93.
[27]
Saucier, J.P.; Bergeron, J.F.; Grondin, P.; Robitaille, A. Les Régions écologiques du Québec Méridional; Ministère des Ressources Naturelles du Québec: Ste-Foy, Canada, 2000.
[28]
Environment Canada National Climate Data and Information Archive. Available online: http://climate.weatheroffice.gc.ca/climateData/canada_e.html (accessed on 1 April 2013).
[29]
Scholander, P.F.; Hammel, H.T.; Bradstreet, E.D.; Hemmingsen, E.A. Sap pressure in vascular plants, negative hydrostatic pressure can be measured in plants. Science 1965, 148, 339–346.
[30]
Hoffman, W.A.; Poorter, H. Avoiding bias in calculation of Relative Growth Rate. Ann. Bot. Lond. 2002, 80, 37–42, doi:10.1093/aob/mcf140.
[31]
Steel, R.G.D.; Torrie, J.H. Principles and Procedures of Statistics: A Biometrical Approach; McGraw-Hill Publishing Company: New York, NY, USA, 1980; p. 633.
[32]
Zar, J.H. Biostatistical Analysis, 4th ed. ed.; Prentice Hall: Upper Saddle River, NJ, USA, 1999; p. 266.
[33]
Devore, J.; Peck, R. Introductory Statistics, 2nd ed. ed.; West Publishing Company: St. Paul, MN, USA, 1994; p. 640.
[34]
Sedia, E.G.; Ehrenfeld, J.G. Lichens and mosses promote alternate stable plant communities in the New Jersey Pinelands. Oikos 2003, 100, 447–458, doi:10.1034/j.1600-0706.2003.12058.x.
[35]
Trofymow, J.A.; van den Driessche, R. Mycorizha. In Mineral Nutrition of Conifer Seedlings; CRC Press: Boston, MA, USA, 1991; pp. 183–229.
[36]
Johansson, K.; Nilson, U.; Lee, A.H. Interaction between soil scarification and Norway spruce seedling types. New For. 2007, 33, 13–27, doi:10.1007/s11056-006-9010-y.
[37]
Rudinger, M.; Hallgren, S.W.; Steudle, E.; Schulze, E.D. Hydraulic and osmotic properties of spruce roots. J. Exp. Bot. 1994, 45, 1413–1425, doi:10.1093/jxb/45.10.1413.
[38]
Kramer, P.J. Water Deficits and Plant Growth. In Water Relations of Plants; Academic Press: New York, NY, USA, 1983; pp. 342–389.
[39]
Grossnickle, S.C. Planting stress in newly planted jack pine and white spruce 1. Factors influencing water uptake. Tree Physiol. 1988, 4, 71–83, doi:10.1093/treephys/4.1.71.
[40]
Kozlowski, T.T.; Davies, W.J. Control of water balance in transplanted trees. J. Arboric. 1975, 1, 1–10.
[41]
Kramer, P.J.; Kozlowski, T.T. Physiology of Woody Plants; Academic Press Inc.: New York, NY, USA, 1979; pp. 58–112.
[42]
Folk, R.S.; Grossnickle, S.C. Stock-type patterns of phosphorus uptake, retranslocation, net photosynthesis and morphological development in interior spruce seedlings. New For. 2000, 19, 27–49, doi:10.1023/A:1006618312161.
[43]
Colombo, S.J. Second year shoot development in black spruce Picea mariana (Mill) BSP container seedlings. Can. J. For. Res. 1986, 16, 68–73, doi:10.1139/x86-011.
[44]
Bigras, F.J.; D’Aoust, A.L. Hardening and dehardening of shoot and roots of containerized black spruce and white spruce seedlings under short and long days. Can. J. For. Res. 1992, 22, 388–396, doi:10.1139/x92-051.
[45]
Malik, V.; Timmer, V.R. Growth, nutrient dynamics, and interspecific competition of nutrient-loaded black spruce seedlings on a boreal mixedwood site. Can. J. For. Res. 1996, 26, 1651–1659, doi:10.1139/x26-186.
[46]
Timmer, V.R.; Morrow, L.D. Predicting Fertilizer Growth Response and Nutrient Status of Jack Pine by Foliar Diagnosis. In Proceedings of the Forest Soils and Treatment Impacts, Sixth North American Forest Soils Conference, University of Tennessee, Knoxville, TN, USA, 1983; pp. 335–351.
[47]
Givnish, T.J. Adaptation to sun and shade: A whole-plant perspective. Funct. Plant Biol. 1988, 15, 63–92.
[48]
Canham, C.D. Different responses to gaps among shade-tolerant tree species. Ecology 1989, 70, 548–550, doi:10.2307/1940200.
[49]
Bazzaz, F.A.; Wayne, P.M. Coping with Environmental Heterogeneity: The Physiological Ecology of Tree Seedling Regeneration across the Gap-Understory Continuum. In Exploitation of Environmental Heterogeneity by Plants: Ecophysiological Processes Above- and Belowground; Caldwell, M.M., Pearcy, R.W., Eds.; Academic Press Inc.: San Diego, CA, USA, 1994; pp. 349–390.
[50]
Walters, M.B.; Reich, P.B. Trade-offs in low-light CO2 exchange: A component of variation in shade tolerance among cold temperate tree seedlings. Funct. Ecol. 2000, 14, 155–165, doi:10.1046/j.1365-2435.2000.00415.x.
[51]
Tan, W.X.; Blake, T.J.; Boyle, T.J.B. Drought tolerance in faster-growing and slower-growing black spruce (Picea mariana) Progenies 1. Stomatal and gas exchange responses to osmotic stress. Physiol. Plant. 1992, 85, 639–644, doi:10.1111/j.1399-3054.1992.tb04766.x.
[52]
Bassman, J.H. Influence of two site preparation treatments on ecophysiology of planted Picea engelmannii × glauca seedlings. Can. J. For. Res. 1989, 19, 1359–1370, doi:10.1139/x89-210.
[53]
Grossnickle, S.C.; Heikurinen, J. Site preparation: Water relations and growth of newly planted jack pine and white spruce. New For. 1989, 3, 99–123, doi:10.1007/BF00021576.
[54]
？rlander, G.; Gemmel, P.; Hunt, J. Site Preparation: A Swedish Overview; British Columbia Ministry of Forests, Canada/BC Economic & Regional Development Agreement: Victoria, BC, Canada, 1990; p. 61.
[55]
Prévost, M. Effet du scarifiage sur les propriétés du sol et de l’ensemencement naturel dans une pessière noire à mousses de la forêt boréale québécoise. Can. J. For. Res. 1996, 26, 72–86, doi:10.1139/x26-008.
[56]
Boucher, J.F.; Wetzel, S.; Munson, A.D. Leaf level response of planted eastern white pine (Pinus strobus L.) seven years after intensive silvicultural treatments. For. Ecol. Manag. 1998, 107, 291–307, doi:10.1016/S0378-1127(97)00350-2.
[57]
Bedford, L.; Sutton, R.F. Site preparation for establishing lodgepole pine in the sub-boreal spruce zone of interior British Columbia: The Bednesti trial, 10-year results. For. Ecol. Manag. 2000, 126, 227–238, doi:10.1016/S0378-1127(99)00090-0.
[58]
Thiffault, N.; Jobidon, R.; Munson, A.D. Performance and physiology of large containerized and bare-root spruce seedlings in relation to scarification and competition in Quebec (Canada). Ann. For. Sci. 2003, 60, 645–655, doi:10.1051/forest:2003057.
[59]
Thiffault, N.; Hébert, F.; Jobidon, R. Planted Picea mariana growth and nutrition as influenced by silviculture × nursery interaction on an ericaceous-dominated site. Silva Fenn. 2012, 46, 667–682.
[60]
Bowen, G.D. Soil Temperature, Root Growth, and Plant Function. In Plant Roots: The Hidden Half; Waisel, Y., Eshel, A., KapFkafi, U., Eds.; Marcel Dekker Inc.: New York, NY, USA, 1991; pp. 309–330.
[61]
Dodd, I.C.; He, J.; Turnbull, C.G.N.; Lee, S.K.; Critchley, C. The influence of supra-optimal root-zone temperature on growth and stomatal conductance in Capsicum annuum L. J. Exp. Bot. 2000, 51, 239–248, doi:10.1093/jexbot/51.343.239.
[62]
Nobel, P.S. Physicochemical & Environmental Plant Physiology; Academic Press: San Diego, CA, USA, 1999; p. 474.
[63]
Brais, S. Persistence of soil compaction and effects on seedling growth in Northwestern Quebec. Soil Sci. Soc. Am. J. 2001, 65, 1263–1271, doi:10.2136/sssaj2001.6541263x.
[64]
Boivin, R.; Miller, B.D.; Timmer, V.R. Late-season fertilisation of Picea mariana seedlings under greenhouse culture: Biomass and nutrient dynamics. Ann. For. Sci. 2002, 59, 255–264, doi:10.1051/forest:2002021.
[65]
Mansuy, N.; Gauthier, S.; Bergeron, Y. Afforestation opportunities when stand productivity is driven by a high risk of natural disturbance: A review of the open lichen woodland in the eastern boreal forest of Canada. Mitig. Adapt. Strateg. Glob. Change 2012, 18, 245–264, doi:10.1007/s11027-012-9362-x.
[66]
Boucher, J.-F.; Tremblay, P.; Gaboury, S.; Villeneuve, C. Can boreal afforestation help offset incompressible GHG emissions from Canadian industries? Process Saf. Environ. 2012, 90, 459–466, doi:10.1016/j.psep.2012.10.011.
