We used multivariate analysis to model boreal forest stand structure and dynamics at Riding Mountain National Park, Manitoba based on data from 202 sampled stands. Eight forest stand-types were recognized based on canopy composition: black spruce on peat substrates, jack pine — black spruce, bur oak, eastern deciduous (green ash — American elm — Manitoba maple), balsam fir, trembling aspen — paper birch — mountain maple, trembling aspen — balsam poplar, and white spruce. The first four stand-types occur in edaphically distinct environments, while the four remaining boreal mixedwood stand-types occur in edaphically similar environments. We found that the composition and abundance of advance regeneration were best predicted by current canopy composition (redundancy = 54.4%); this reflects both the limited dispersal of conifer seeds and the strong vegetative reproductive capacity of hardwoods. Biotically-controlled site factors such as bareground, herb and shrub cover, ungulate browsing intensity, and stand age were also reasonably good predictors of advance regeneration (redundancy = 31.7%). Edaphic variables such as soil pH, conductivity, particle size, organic horizon depth and slope proved to be poor predictors of advance regeneration, however (redundancy = 18.1%). Size-class ordination indicated that many stand-types have relatively short successional trajectories, suggesting limited change in forest canopy composition over time. There are two exceptions: in the jack pine — black spruce stand-type, black spruce will increase over time, and in the trembling aspen — paper birch — mountain maple stand-type, eastern deciduous species (green ash, American elm, Manitoba maple, and bur oak) are forecast to become increasingly dominant. We also describe a synoptic model of mixedwood boreal forest stand dynamics for the Riding Mountain area. The model includes a number of factors that we consider to be critical determinants of forest dynamics, such as seed source availability, small and large-scale disturbances, species life-history characteristics, and environmental gradients. Our succession model is more similar those described for eastern than western Canada, which may reflect the lower frequency of catastrophic fires in the Riding Mountain area compared to boreal forests further west. Our model emphasizes that successional trajectories do not converge towards a single self-perpetuating “climax”. Instead, successional vectors may diverge, converge or remain cyclical, and multiple potential pathways are possible for each stand-type. Our results also illustrate that species assemblages, and the propensity for canopy change in the absence of fire, are governed by the cumulative and synergistic effects of climate, topography, disturbance frequency, size and intensity, edaphic conditions, and the proximity of parental seed sources. Fire suppression in the southern boreal forest has resulted in a paradigm shift in disturbance regime, from large, synchronous catastrophic fires to small-scale, asynchronous gap formation. A major challenge for boreal forest ecologists is to determine the long-term consequences of this paradigm shift on the composition, structure and health of boreal forest stands and landscapes.
Riding Mountain National Park
Canonical correspondence analysis
Achuff, P.L. and G.H. La Roi. 1977. Picea-Abies forests in the highlands of northern Alberta. Vegetatio 33: 127–146.
Agriculture Canada Expert Committee on Soil Survey. 1987. The Canadian System of Soil Classification. 2nd ed. Agric. Can. Publ. 1646, Ottawa.
Bailey, R.H. 1968. Notes on the vegetation of Riding Mountain National Park, Manitoba. National Parks Forest Survey No. 2. Dept. Forestry Rural Develop., Forest Management Inst., Ottawa.
Bergeron, Y. and M. Dubuc. 1989. Succession in the southern part of the Canadian boreal forest. Vegetatio 79: 51–63.
Bergeron, Y. and P. Dansereau. 1993. Predicting the composition of Canadian southern boreal forest in different fire cycles. J. Veg. Sci. 4: 827–832.
Bergeron, Y. 2000. Species and stand dynamics in the mixed woods of Québec’s southern boreal forest. Ecology 81:1500–1516.
Bird, R. D. 1961. Ecology of the Aspen Parkland of Western Canada. Research Branch, Canada Dept. Agriculture, Publ. 1066, Ottawa.
Borcard, D. and P. Legendre. 1994. Environmental control and spatial structure in ecological communities: an example using oribatid mites (Acari, Oribates). Environ. Ecol. Stat. 1: 37–61.
Brandt, J. P. 1993. Forest Insect and Disease Conditions in West-central Canada in 1993 and Predictions for 1994. Inf. Rep. NOR-X-335. Nat. Res. Can., Can. For. Serv., Northwest Reg., North. For. Cent., Edmonton.
Caners, R. 2001. Landscape-level vegetation dynamics in Riding Mountain National Park, Manitoba, Canada. MSc. Thesis, Univ. Manitoba, Winnipeg. 270 pages.
Carleton, T.J. and P.F. Maycock. 1978. Dynamics of the boreal forest south of James Bay. Can. J. Bot. 56: 1157–1173.
Chen, H.Y.H. and R.V. Popadiouk. 2002. Dynamics of North American boreal mixedwoods. Environ. Rev. 10: 137–166.
Cody, W.J. 1988. Plants of Riding Mountain National Park, Manitoba. Publ. 1818/E, Res. Branch, Agric. Canada, Ottawa.
Cogbill, C.V. 1985. Dynamics of the boreal forests of the Laurentian Highlands, Canada. Can. J. For. Res. 15: 252–261
Connell, J.H. and R.O. Slatyer. 1977. Mechanisms of succession in natural communities and their role in community stability and organization. Am. Nat. 111: 1119–1144.
Cornett, M.W., K.J. Puettmann and P.B. Reich. 1998. Canopy type, forest floor, predation, and competition influence conifer seedling emergence and early survival in two Minnesota conifer-deciduous forests. Can. J. For. Res. 28: 196–205.
Dickson, J.R. 1909. The Riding Mountain Forest Reserve. Bulletin No. 6, Dept. Interior, Forestry Branch, Ottawa.
Dix, R.L. and J. M. A. Swan. 1971. The roles of disturbance and succession in upland forest at Candle Lake, Saskatchewan. Can. J. Bot. 49: 657–676.
Drobyshev, I. and B. Nihlgård. 2000. Growth response of spruce saplings in relation to climatic conditions along a gradient of gap size. Can. J. For. Res. 30: 930–938.
Egler, F.E. 1954. Vegetation science concepts I. Initial floristic composition, a factor in old-field vegetation development. Vegetatio 4:412–417.
Environment Canada. 1993. Canadian climate normals 1961–1991. Vol. 2. Atmospheric Environ. Serv., Ottawa.
Evans, D.L.S. 1923. Report of land classification surveys in part of Riding Mountain Forest Reserve. Topographical Survey of Canada, Dept. Interior, Ottawa.
Fastie, C.L. 1995. Causes and ecosystem consequences of multiple successional pathways of primary succession at Glacier Bay, Alaska. Ecology 76: 1899–1916.
Foré, S.A, J.L. Vankat and R.L. Schaefer. 1997. Temporal variation in the woody understory of an old-growth Fagus-Acer forest and implications for overstory recruitment. J. Veg. Sci. 8: 607–614.
Foster, B.L. and D. Tilman. 2000. Dynamic and static views of succession: testing the descriptive power of chronosequence approach. Plant Ecol. 146: 1–10.
Freiich, L.E. and P.B. Reich. 1995. Spatial patterns and succession in a Minnesota southern-boreal forest. Ecol. Monogr. 65: 325–346.
Galipeau, C., D. Kneeshaw and Y. Bergeron. 1997. White spruce and balsam fir colonization in a site in the southeastern boreal forest as observed 68 years after fire. Can. J. For. Res. 27: 139–147.
Greene, D.F. and E.A. Johnson. 1995. Long-distance wind dispersal of tree seeds. Can. J. Bot. 73: 1036–1045.
Greene, D.F. and E.A. Johnson. 1996. Wind dispersal of seeds from a forest into a clearing. Ecology 77: 595–609.
Greene, D.F. and E.A. Johnson. 1997. Secondary dispersal of tree seeds on snow. J. Ecol. 85: 329–340.
Greene, D.F., J.C. Zasada, L. Sirois, D. Kneeshaw, H. Morin, I. Charron and M.-J. Simard. 1999. A review of the regeneration dynamics of North American boreal forest tree species. Can. J. For. Res. 29: 824–839.
Grigal, D.F. and L.F. Ohmann. 1975. Classification, description, and dynamics of upland plant communities within a Minnesota wilderness area. Ecol. Monogr. 45: 389–407.
Outsell, S.L. and E.A. Johnson. 2001. Accurately ageing trees and examining their height-growth rates: implications for interpreting forest dynamics. J. Ecol. 90: 153–166.
Halliday, W.E.D. 1932. Vegetation and site studies – Clear Lake, Riding Mountain National Park, Manitoba.
Heinselman, M.L. 1973. Fire in the virgin forests of the Boundary Waters Canoe Area, Minnesota. Quaternary Res. 3: 329–362.
Heinselman, M.L. 1978. Fire intensity and frequency as factors in the distribution and structure of northern ecosystems. In: Mooney, H.A., T.M. Bonnicksen and N.L. Christensen (eds.). 1978. Conference Proceedings, Fire Regimes and Ecosystem Properties, 11–15 Dec. 1978. Honolulu, Hawaii. USDA For. Serv. Gen. Tech. Rep. WO-26, Washington D.C.
Heinselman, M.L. 1981. Fire and succession in the conifer forests of northern North America. In: West, D.C., H.H. Shugart and D.B. Botkin (eds.). Forest Succession: Concepts and Application. Springer-Verlag, New York. Pages 374–406.
Hill, M.O. 1973. Diversity and evenness: a unifying notation and its consequences. Ecology 54: 427–432.
Johnson, E.A. 1992. Fire and Vegetation Dynamics: Studies from the North American Boreal Forest. Cambridge Univ. Press, Cambridge.
Johnson, E.A., K. Miyanishi and J.M.H. Weir. 1995. Old-growth, disturbance, and ecosystem management. Can. J. Bot. 73: 918–926.
Johnson, E.A., K. Miyanishi and J.M.H. Weir. 1998. Wildfires in the western Canadian boreal forest: Landscape patterns and ecosystem management. J. Veg. Sci. 9: 603–610.
Johnson, E.A., K. Miyanishi and H. Kleb. 1994. The hazards of interpretation of static age structures as shown by stand reconstructions in a Pinus conforta – Picea engelmanii forest. J. Ecol. 82: 923–931.
Kalra, Y.P. and D.G. Maynard. 1991. Methods manual for forest soil and plant analysis. Inf. Rep. NOR-X-319. For. Can., Northwest Reg., North. For. Cent. Edmonton.
Kenkel, N.C. 1986. Structure and dynamics of jack pine stands near Elk Lake, Ontario: amultivariate approach. Can. J. Bot. 64: 486–497.
Kenkel, N.C., D.J. Walker, P.R. Watson, R.T. Caners and R.A. Lastra. 1997. Vegetation dynamics in boreal forest ecosystems. Coenoses 12: 97–108.
Kenkel, N.C., C. Foster, R. Caners, R. Lastra and D. Walker. 2003. Spatial and temporal patterns of white spruce recruitment in two boreal mixedwood stands, Duck Mountain, Manitoba. Project Report 2003/2004. Sustainable Forest Management Network, Edmonton.
Kneeshaw, D.D. and P.J. Burton. 1997. Canopy and age structures of some old sub-boreal Picea stands in British Columbia. J. Veg. Sci. 8:615–626.
Kneeshaw, D.D. and Y. Bergeron. 1996. Ecological factors affecting the abundance of advance regeneration in Quebec’s southwestern boreal forest. Can. J. For. Res. 26: 888–898.
Kneeshaw, D.D. and Y. Bergeron. 1998. Canopy gap characteristics and tree replacement in the southeastern boreal forest. Ecology 79: 783–794.
Kneeshaw, D.D., Y. Bergeron and L. De Grandpré. 1998. Early response of Abies balsamea seedlings to artificially created openings. J. Veg. Sci. 9: 543–550.
Lang, A. H. 1974. Guide to the geology of Riding Mountain National Park and its vicinity - history of its upland and other scenery. Misc. Report 20. Geol. Survey Canada, Dept. Energy, Mines, and Resources, Ottawa
Larocque, I., Y. Bergeron, I.D. Campbell and R.H.W. Bradshaw. 2000. Vegetation changes through time on islands of Lake Duparquet, Abitibi, Canada. Can. J. For. Res. 30: 179–190.
Leps, J. and P. Smilauer. 2003. Multivariate Analysis of Ecological Data Using CANOCO. Cambridge Univ. Press, Cambridge.
Lieffers, V.J. and K.J. Stadt. 1994. Growth of understory Picea glauca, Calamagrostis canadensis, and Epilobium angustifolium in relation to overstory light. Can. J. For. Res. 24: 1193–1198.
Lieffers, V.J., K.J. Stadt and S. Navratil. 1996. Age structure and growth of understory white spruce under aspen. Can. J. For. Res. 26: 1002–1007.
Lorimer, C.G., J.W. Chapman and W.D. Lambert. 1994. Tall understory vegetation as a factor in the poor development of oak seedlings beneath mature stands. J. Ecol. 82: 227–237.
Maguire, D.A. and R.T.T. Forman. 1983. Herb cover effects on tree seedling patterns in amature hemlock-hardwood forest. Ecology 64: 1367–1380.
McClanahan, T.R. 1986. The effect of a seed source on primary succession on a forest ecosystem. Vegetatio 65: 175–178.
McCune, B. and T.F.H. Allen. 1985a. Will similar forests develop on similar sites?. Can. J. Bot. 63:367–376.
McCune, B. and T.F.H. Allen. 1985b. Forest dynamics in the Bitterroot Canyons, Montana. Can. J. Bot. 63:377–383.
McInnes, P.F., R.J. Naiman, J. Pastor and Y Cohen. 1992. Effects of moose browsing on vegetation and litter of the boreal forest, Isle Royale, Michigan, USA. Ecology 13: 2059–2075.
McLaren, B.E. and R.A. Janke. 1996. Seedbed and canopy cover effects on balsam fir seedling establishment in Isle Royale National Park. Can. J. For. Res. 26: 782–793.
Messier, C., S. Parent and Y. Bergeron. 1998. Characterization of understory light environment in closed mixed broadleaf conifer boreal forests: effects of overstory and understory vegetation. J. Veg Sci. 9:511–520.
Morin, H. and D. Laprise. 1997. Seedling bank dynamics in boreal balsam fir forests. Can. J. For. Res. 27: 1442–1451.
Morin, H. 1994. Dynamics of balsam fir forests in relation to spruce budworm outbreaks in the boreal zone of Quebec. Can. J. For. Res. 24: 730–741.
Naiman, R.J. 1988. Animal influences on ecosystem dynamics. Bioscience 38: 750–752.
Naiman, R.J., C.A. Johnston and J.C Kelley. 1988. Alteration of North American streams by beaver. Bioscience 38: 753–762.
Oechel, W.C. and W.T. Lawrence. 1985. Taiga. In: Chabot, B.F. and H.A. Mooney (eds.). 1985. Physiological Ecology of North American Plant Communities. University Press, Cambridge. Pages 67–94.
Okland, R.H. 1999. On the variation explained by ordination and constrained ordination axes. J. Veg. Sci. 10: 131–136.
Parks Canada. 1997. Riding Mountain National Park Ecosystem Conservation Plan. Department of Canadian Heritage, Ottawa (1997, March). [Online]. Available: http://www.parksca-nada.gc.ca/parks/manitoba/Riding_Mountain/english/ECPlan_e.htmhttp://www.parksca-nada.gc.ca/parks/manitoba/Riding_Mountain/english/ECPlan_e.htm
Pastor, J., B. Dewey, R.J. Naiman, PF. Mclnnes and Y. Cohen. 1993. Moose browsing and soil fertility in the boreal forests of Isle Royale National Park. Ecology 74:467–480.
Perala, D.A. 1990. Populus tremuloides. In: Burns, R.M. and B.H. Honkala (eds.) Silvios of North America: 2. Hardwoods. Agricultural Handbook 654. USDA, Forest Service, Washington D.C Vol.2. Pages 555–569.
Peterson, E.B. and N.M. Peterson. 1992. Ecology, management, and use of aspen and balsam poplar in the prairie provinces, Canada. Spec. Rep. 1. Can., Northwest Reg., North. For. Cent., Edmonton.
Rees, M. R. Condit, M. Crawley, S. Pacala and D. Tilman. 2001. Long-term studies of vegetation dynamics. Science 293: 650–655.
Ritchie, J.C. 1956. The vegetation of northern Manitoba I. Studies in the southern spruce forest zone. Can. J. Bot. 34: 523–561.
Ritchie, J.C. 1957. The vegetation of the Wilson Creek Watershed, Riding Mountain National Park, Manitoba. Dept. Northern Affairs and National Resources, Forestry Branch, Forest Research Division, Ottawa.
Ritchie, J.C. 1964. Contributions to the Holocene paleoecology of west-central Canada. I. The Riding Mountain Area. Can. J. Bot. 42: 181–197.
Ritchie, J.C. 1969. Absolute pollen frequencies and carbon-14 age of a section of Holocene Lake sediment from the Riding Mountain area of Manitoba. Can. J. Bot. 47: 1345–1349.
Ritchie, J.C. 1985. Quaternary pollen records from the Western Interior and the Arctic of Canada. In: Bryant, V.M. Jr. and R.G. Holloway (eds.). Pollen records of late-Quaternary North American sediments. American Association of Stratigraphie Palynologists Foundation, Dallas. Pages 327–352.
Ritchie, M.E., D. Tilman and J.M.H. Knops. 1998. Herbivore effects on plant and nitrogen dynamics in oak savanna. Ecology 79:165–177.
Rowe, J.S. 1955. Factors influencing white spruce reproduction in Manitoba and Saskatchewan. Tech. Note 3, Dept. Northern Affairs and National Resources, Forestry Branch. Ottawa
Rowe, J.S. 1956. Vegetation of the southern boreal forest in Saskatchewan and Manitoba. PhD Thesis. Univ. Manitoba, Winnipeg.
Rowe, J.S. 1961. Critique of some vegetational concepts as applied to forests of northwestern Alberta. Can. J. Bot. 39: 1007–1017.
Rowe, J.S. 1972. Forest regions of Canada. Dept. Envt., Can. For Ser., Ottawa, Publ. No. 1300.
Rudolph, TD. and PR. Laidley. 1990. Pinus banksiana. In: Burns, R.M. and B.H. Honkala (eds.). Silvios of North America: 1. Conifers. Agric. Handbook 654. USDA, Forest Service, Washington D.C. Vol. 1. Pages 280–290
Rydgren, K., G. Hestmark and R.H. Okland. 1998. Revegetation following experimental disturbance in a boreal old-growth Picea abies forest. J. Veg. Sci. 9: 763–776.
Sentar Consultants Ltd. 1992. Riding Mountain National Park: A literature review of historic timber harvesting and forest fire activity.
Shafi, M.I. and G.A. Yarranton. 1973. Vegetational heterogeneity during a secondary (postfire) succession. Can. J. Bot. 51: 73–90.
Shugart, H.H., R. Leemans and G.B. Bonan. 1992. A Systems Analysis of the Global Boreal Forest. Cambridge University Press. Cambridge.
Simard, M.-J., Y. Bergeron and L. Sirois. 1998. Conifer seedling recruitment in a southeastern Canadian boreal forest: the importance of substrate. J. Veg. Sci. 9: 575–582.
Sprugel, D.G. 1976. Dynamic structure of wave-regenerated Abies balsamea forests in the northeastern United States. J. Ecol. 64: 889–912.
Stewart, J.D., E.H. Hogg, P.A. Hurdle, K.J. Stadt, P. Tollestrup and V.J. Lieffers. 1998. Dispersal of white spruce seed in mature aspen stands. Can. J. Bot. 76: 181–188.
ter Braak, C.J.F. 1986. Canonical correspondence analysis: a new eigenvector technique for multivariate direct gradient analysis. Ecology 67: 1167–1179.
ter Braak, C.J.F. 1987. CANOCO – a FORTRAN program for canonical community by [partial] [detrended] [canonical] correspondence analysis, principal components analysis, and redundancy analysis (version 3.11). TNO Institute of Applied Computer Science, Wageningen.
Trottier, G.C. 1980. Aerial beaver censuses in Riding Mountain National Park, 1973–1980. Large Mammal System Studies Report No. 11. Canadian Wildlife Service, Ottawa.
Trottier, G.C. 1981. Beaked hazelnut - a key browse species for moose in the boreal forest region of western Canada. Alces 17: 257–281.
Trottier, G.C., S.R. Rollans and R.C. Hutchison. 1983. Range, habitat and foraging relationships of ungulates in Riding Mountain National Park. Large Mammal System Studies Report No. 14. Canadian Wildlife Service, Ottawa.
Tunstell, G. 1940. Forest working plan, Riding Mountain National Park. Report prepared for the Department of Mines and Resources, Lands, Parks, and Forests Branch, Ottawa.
Tunstell, G., C.B. Gill and G.F. Kuhring. 1922. Silvical report on Riding and Duck Mountain Forest Reserves. Dept. Interior, Forestry Branch, Ottawa.
Viereck, L.A. and W.F. Johnston. 1990. Picea mariana. In: Burns, R.M. and B.H. Honkala (eds.). Silvics of North America: 1. Conifers. Agricultural Handbook 654. USDA, Forest Service, Washington D.C. Vol. 1. Pages 227–235.
Vincent, A.B. 1965. Growth habits of mountain maple in the Ontario clay belt. For. Chron. 41: 330–344.
Waldron, R.M. 1966. Factors affecting natural white spruce regeneration on prepared seedbeds at the Riding Mountain forest experimental area, Manitoba. Dept. Publication No. 1169. Dept. Forestry and Rural Development, Forestry Branch, Ottawa.
Walker, D.J. and N.C. Kenkel. 2001. Landscape complexity in space and time. Community Ecology 2: 109–119.
Wein, R.W. and D.A. MacLean. 1983. The Role of Fire in Northern Circumpolar Ecosystems. John Wiley and Sons, New York.
Weir, J.M.H. and E.A. Johnson. 1998. Effects of escaped settlement fires and logging on forest composition in the mixedwood boreal forest. Can. J. For. Res. 28: 459–467.
Wilson, J.B, H. Gitay, S.H. Roxburgh, W.M. King and R.S. Tangney 1992. Egler’s concept of ‘initial floristic composition’ in succession - ecologists citing it don’t agree what it means. Oikos 64: 591–593.
Youngblood, A.P. 1995. Development patterns in young coniferhardwood forests of interior Alaska. J. Veg. Sci. 6: 229–236.
Zasada, J.C. and H.M. Phipps. 1990. Populus balsamifera. In: Burns, R.M. and B.H. Honkala (eds.) Silvics of North America: 2. Hardwoods. Agricultural Handbook 654. USDA, Forest Service, Washington D.C. Vol. 2. Pages 518–529.
Zasada, J.C., T.L. Sharik and M. Nygren. 1992. The reproductive process in boreal forest trees. In: Shugart, H.H., R. Leemans and G.B. Bonan (eds.). A Systems Analysis of the Global Boreal Forest. Cambridge Univ. Press, Cambridge. Pages 85–125.
Zoladeski, C.A. and P.F. Maycock. 1990. Dynamics of the boreal forest in northwestern Ontario. Am. Midl. Nat. 124: 289–300.
This research was supported by grants from Parks Canada and the Natural Sciences and Engineering Research Council (NSERC) of Canada to N. Kenkel. We thank Rod Lastra, Boyan Tracz, Heidi Wiebe and David Walker for assistance with fieldwork. Numerous personnel from Parks Canada provided additional field assistance and logistic support — we extend our special thanks to Wybo Vanderschuit for his guidance and support.
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Caners, R.T., Kenkel, N.C. Forest stand structure and dynamics at Riding Mountain National Park, Manitoba, Canada. COMMUNITY ECOLOGY 4, 185–204 (2003). https://doi.org/10.1556/ComEc.4.2003.2.6
- Advance regeneration
- Boreal forest
- Eastern deciduous