Scott Rupp
Interior Alaska is characterized by a mosaic of vegetation, which reflects the disturbance history of the landscape (Viereck 1973). In the uplands, fire resets the successional clock (Van Cleve et al. 1991). Fire frequency for interior white spruce ecosystems is estimated at 105 yr (Yarie 1981). These white spruce ecosystems are both commercially and socially important components of the boreal forest. The wise management and continued persistence of these systems are important issues in the management of interior Alaska forests. Simulating early establishment patterns of white spruce can provide information helpful in the management of these systems and may provide for a decreased dependence upon planting of both burned and harvested sites.
A geographic information system (GIS) provides an excellent framework for developing a spatially explicit model of early post-disturbance seedling establishment patterns. The strict geographic representation allows the identification of specific seed sources, description of the seedbed, and simulation of the effects of competing vegetation and environmental constraints upon seedling establishment. Creation of the model within a GIS provides for spatially explicit representation of key biological processes and offers the land manager relevant information on the regeneration patterns of a defined landscape.
The objective of this study was to develop a geographic model of white spruce seedling establishment patterns within an upland white spruce community following a wildfire event. The model was developed entirely within ArcInfo using AML's and the GRID environment. Routines were developed to simulate the production and dispersal of seed upon the landscape, to simulate the effects of fire upon the seedbed and competing vegetation, and simulate the initial establishment pattern of seedlings within a burnt upland white spruce ecosystem.
This paper provides a general description of the model and details individual model routines. A brief comparison of model results against both field observations and the literature is included. The discussion focuses on potential management applications.
The model subroutines are a simplistic representation of Harper's model (1977), focusing on the simulation of white spruce seedling establishment patterns. ABFEM simulates the production of seed (white spruce and paper birch), the reproductive potential of the landscape including both the seed (white spruce and paper birch wind dispersed seed) and vegetative bank (paper birch basal sprouts and aspen root suckers), and the early establishment of seedlings (white spruce, paper birch, and aspen) conditioned by the environmental sieve described by Harper (1977).
Empirical data used to develop and calibrate individual routines were measured in the Fairbanks area of interior Alaska over the past 4 decades. Data from the literature was used to fill certain information gaps. Model validation was carried out through comparison of model predictions with the literature and observations from interior Alaska.
Using probabilities and random numbers, the cone/catkin crop is rated as poor, moderate-good, or excellent. The total within stand seed fall per unit area (100 m^2 ) and percent viability associated with the cone/catkin crop rating are calculated. The NORMAL function in GRID is used to develop normal distributions following the production trends observed from long-term records of seed production, in the Fairbanks area of interior Alaska. NORMAL develops a normal distribution and then randomly choses a value within the distribution for output. A normal distribution of total seed density is developed based upon the observed means and standard deviations from local production records for the given crop rating. A value within the distribution is randomly chosen and assigned to the parameters of total seed and percent viable. Total within stand viable seed is then calculated. The routine output is the number of viable seed per unit area (100 m^2 ) produced within the source stand for each species.
DISPERSE calculates the Euclidean distance of each cell within the disturbed area to the closest source cell, using the EUCDISTANCE function. This distance grid is input into the dispersal equation along with the viable seed parameter to create an output grid of dispersed viable seed.
Reports from the literature suggest the influence of winds upon the "seed shadow" (Harris 1967, Schlesinger 1970, Janzen 1971, Sharpe and Fields 1982, Zasada and Lovig 1983, Youngblood and Max 1992, Greene and Johnson 1995, 1996, Rupp personal communication). The WINDS subroutine provides the option for the simulation of potential seed dispersing wind effects upon the spatial distribution of white spruce seed about its source. The euclidean direction of each cell within the disturbed area to the closest source cell is calculated and used to determine the proper dispersal equation to be applied. Separate dispersal curves were developed for white spruce seed dispersing with the wind and seed dispersing against the wind.
ESTABLISH uses a dynamic seed:seedling index and seedbed information along with the grid of dispersed seed to determine successfully established seedlings on a cell by cell basis. The dynamic seed:seedling index is an index that calculates the number of viable seed needed to produce one established 5 year old seedling given the time since disturbance and the seedbed conditions encountered by the dispersed seed. Seedlings originating from seed (white spruce and paper birch) are calculated using the seed:seedling index. The seed:seedling index uses seed:seedling ratios developed for white spruce (Ackerman 1957, Eis 1967, Zasada 1971, Clautice 1974, Ganns 1977, Zasada et al. 1978, Walker 1986, Walker et al. 1986, Zasada et al. 1992, Zasada personal communication) and paper birch (Marquis et al. 1964, Horsley and Abbott 1970, Zasada 1971, Clautice 1974, Zasada et al. 1978, Zasada 1985, Perala 1987, Perala and Alm 1989) associated with seedbed substrate characteristics, topography, and time since disturbance to determine the number of 5 yr seedlings established within a cell.
Seedbed substrate characteristics are simulated by the subroutines DISTURB and SPROUT. The DISTURB subroutine simulates the effects of a disturbance (wildfire) upon the forest floor and subsequent seedbed. Fire in interior Alaska, as well as in most fire-dominated systems, is heterogeneous in nature; producing a spatially variable pattern in both areal extent and intensity/severity (Lutz 1956, Quirk and Sykes 1971, Viereck 1973, Heinselman 1981, Thomas and Wein 1985, Hobbs and Atkins 1988, Ratz 1995). DISTURB produces a heterogeneous burn pattern across the landscape, resulting in various degrees of forest floor consumption. A grid of random numbers is produced for the disturbed area and classified into 1 of 5 burn severity classes, following Dyrness and Norum (1983). The probability of occurrence of a specific severity class is input by the user. Several heterogeneous burn patterns can be defined by the user, which are created using the initial classified disturbance grid and various focal functions (i.e. FOCALMEAN and FOCALMAJORITY) within GRID.
The subroutine SPROUT simulates the occupation of disturbed cells by aspen root suckers and paper birch basal sprouts. This subroutine provides information on both potential establishment patterns of aspen clones and paper birch sprouts and provides information on specific vegetative competition, in terms of site occupation, experienced by white spruce seedlings. SPROUT uses pre-disturbance vegetation patterns and output from DISTURB to determine post-disturbance vegetative reproduction patterns.
From a functional standpoint, each model routine has some major shortcomings. SEEDS performs well simulating general trends in production periodicity and variability. However, the routine lacks the predictive ability to identify actual occurrence of future crops, important to natural regeneration management.
DISPERSE describes the general pattern of seed dispersal for white spruce and paper birch and provides some indication of the influence winds may have upon the seed shadow. However, more information is needed to accurately model dispersion of seed upon the landscape and identify specific effects of topography and wind upon the spatial distribution of seed. For example, research regarding aspen regeneration has focused on vegetative reproduction (Peterson and Peterson 1992) and little information exists on aspen seed dispersal. Also, certain irregular seed source shapes result in discontinuity of the seed shadow, due to the manner in which the euclidean direction of each cell is calculated by the EUCDISTANCE function in GRID.
The ESTABLISH routine is a very simplistic representation of the establishment process and over-generalizes the complexity of the germination, survival, growth, and establishment of seedlings. This approach was intentional, with the objective of describing seedling establishment patterns in a manner that would provide practical information to the land manager. Seed:seedling ratios provide this practicality, but the data needed to develop the proper level of detail, in terms of viable seed needed to produce an established seedling for a specific seedbed environment, is lacking. Furthermore, representation of the spatial pattern of fire effects by DISTURB lacks empirical data. SPROUT also lacks empirical data, particularly for interior Alaska, and provides only a speculative generalized view of the vegetative reproduction process in burnt upland white spruce communities in interior Alaska.
The simulation involved an area of the BCEF uplands containing stands of pure white spruce and white spruce/mixed hardwoods. A 35 ha island of white spruce/mixed hardwoods survived the fire and served as a major seed source (Figure 1).
Other regions of the North American boreal forest provide for only a very generalized comparison, due to significant differences in both site and environmental conditions. Studies from Canada report a range of seedling densities from 0 to over 9000 seedlings per 100 m2 on disturbed sites (Phelps 1948, Timoney and Peterson 1996).
The BCEF study site can be used to demonstrate potential management applications. Using the island of surviving white spruce/mixed hardwoods (Figure 1), we can provide an example of the potential use of ABFEM in silvicultural planning and decision making. The island serves as a hypothetical seed source following a harvest operation (i.e. clearcut, strip cutting, etc.). ABFEM allows for investigation of the influence of seed source orientation and its interaction with wind upon the seed shadow. Figure 3 provides an example of the influence seed source orientation can have upon the seed shadow of white spruce.
The model can also be utilized to investigate the potential influence of site preparation upon seedling establishment patterns. Figure 4 provides an example of the influence seedbed preparation can have upon establishment patterns.
The model has several shortcomings related to the lack of empirical data for key processes and due to limitations in the GRID environment and AML. Future research will concentrate on gathering the empirical data needed to properly parameterize and calibrate the model and to further develop the model structure towards a user friendly format for use by the land manager.
Bjorkbom, J.C. 1971. Production and germination of paper birch seed and its dispersal into a forest opening. USDA For. Serv. Northeast For. Exp. Sta. Res. Pap. NE-209. 14p.
Bjorkbom, J.C., D.A. Marquis, and F.E. Cunningham. 1965. The variability of paper birch seed production, dispersal, and germination. USDA For. Serv. Northeast For. Exp. Sta. Res. Pap. NE-41. 8p.
Clautice, S.F. 1974. Spruce and birch germination on different seedbeds and aspects after fire in interior Alaska. Masters Thesis. University of Alaska Fairbanks. 94 p.
Dobbs, R.C. 1976. White spruce seed dispersal in central British Columbia. For. Chron. 52: 225-228.
Dyrness, C.T., and R.A. Norum. 1983. The effects of experimental fires on black spruce forest floors in interior Alaska. Can J. For. Res. 13: 879-893.
Eis, S. 1967. Establishment and early development of white spruce in the interior of British Columbia. For. Chron. 43: 174-177.
Farmer, R.E. 1997. Seed ecophysiology of temperate and boreal zone forest trees. St. Lucie Press, Delray Beach, Florida. 253p.
Fox, J.D., J.C. Zasada, A.F. Gasbarro, and R. Van Veldhuizen. 1984. Monte Carlo simulation of white spruce regeneration after logging in interior Alaska. Can J. For. Res. 14: 617-622.
Ganns, R.C. 1977. Germination and survival of artificially seeded white spruce on prepared seedbeds on an interior Alaska floodplain site. Masters Thesis. University of Alaska Fairbanks. 81p.
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.
Harper, J.L. 1977 Population biology of plants. Acedemic Press, London. 892p.
Harris, A.S. 1967. Natural reforestation on a mile-square clearcut in southeast Alaska. USDA For. Serv. Res. Pap. No. PNW-52.
Heinselman, M.L. 1981. Fire intensity and frequency as factors in the distribution and structure of northern ecosystems. In Fire regimes and ecosystem properties. Proceedings of the conference, December 1978, Honolulu, HI. U.S. For. Serv. Gen. Tech. Rep. WO-26. pp. 7-57.
Hobbs, R.J., and L. Atkins. 1988. Spatial variability of experimental fires in south-west Western Australia. Australian J. Ecol. 13: 295-299.
Horsley, S.B., and H.G. Abbott. 1970. Direct seeding of paper birch in strip clearcutting. J. For. 68: 635-638.
Janzen, D.H. 1971. Seed predation by animals. Ann. Rev. of Ecol. and Sys. 2: 465-492.
Juday, G.P. and C.T. Dyrness (eds.). 1985. Early Results of the Rosie Creek fire research project 1984. School Agr. Land Resources Manage. Univ. Alaska, Fairbanks, Alaska. Misc. Publ. 85-2.
Lutz, H.J. 1956. Ecological effects of forest fires in the interior of Alaska. Tech. Bull. 1133. Washington, D.C.: U.S. Department of Agriculture. 121p.
Marquis, D.A., J.C. Bjorkbom, and G. Yelenosky. 1964. The effect of seedbed condition and light exposure on paper birch regeneration. J. For. 62: 876-881.
Okubo, A., and S.A. Levin. 1989. A theoretical framework for data analysis of wind dispersal of seeds and pollen. Ecology. 70: 329-338.
Packee, E.C. 1990. White spruce regeneration on a blade-scarified Alaskan loess soil. Northern J. Appl. For. 7: 121-123.
Perala, D.A. 1987. Regenerating the birches: ecology and cultural regquirements. PhD. Dissertation. University of Minnesota. 215p.
Perala, D.A., and A.A. Alm. 1989. Regenerating paper birch in the Lake States with the shelterwood method. North J. Appl. For. 6: 151-153.
Peterson, E.B., and N.M. Peterson. 1992. Ecology, management, and use of aspen and balsam poplar in the prairie provinces, Canada. For. Can., Northwest Reg., North. For. Cent. Edmonton, Alberta. Spec. Rep. 1. 252p.
Phelps, 1948. White spruce reproduction in Manitoba and Saskatchewan. Can. Dep. Mines and Resources, Dominion For. Serv., Res. Note 86.
Quirk, W.A., and D.J. Sykes. 1971. White spruce stringers in a fire-patterned landscape in interior Alaska. In Proceedings, fire in the northern environment, a symposium. Pacific Northwest For. and Ran. Exp. Sta., Portland, Oregon. pp. 179-197.
Ratz, A. 1995. Long-term spatial patterns created by fire: a model oriented towards boreal forests. Int. J. Wildland Fire. 5: 25-34.
Rowe, J.S. 1955. Factors influencing white spruce reproduction in Manitoba and Saskatchewan. Canada Dep. North. Affairs and Nat. Resources, For. Br., Res. Div., Tech. Note 3.
Safford, L.O., J.C. Bjorkbom, and J.C. Zasada. 1990. Paper birch (Betula papyrifera Marsh.). In Silvics of native and naturalized trees of the United States and Puerto Rico. R.M. Burns (Tech. Compiler). USDA For. Serv. Washington, DC. Agric. Handb. 271.
Schlesinger, R.C. 1970. Diffusion models applied to seed dispersal. Ph.D. dissertation. Syracuse Univ. 103p.
Sharpe, D.M., and D.E. Fields. 1982. Integrating the effects of climate and seed fall velocities on seed dispersal by wind: a model and application. Ecol. Modelling. 17: 297-310.
Thomas, P.A., and R.W. Wein. 1985. The influence of shelter and the hypothetical effect of fire severity on the postfire establishment of conifers from seed. Can. J. For. Res. 15: 148-155.
Timoney, K.P., and G. Peterson. 1996. Failure of natural regeneration after clearcut logging in Wood Buffalo National Park, Canada. For. Ecol. and Management. 87: 89-105.
Van Cleve, K., F.S. Chapin, III, C.T. Dyrness, and L.A. Viereck. 1991. Element cycling in taiga forests: State-factor control. BioScience. 41: 78-88.
Viereck, L.A. 1973. Wildfire in the taiga of Alaska. Quat. Res. 3:465-495.
Walker, L.R., and F.S. Chapin III. 1986. Physiological controls over seedling growth in primary succession on an Alaskan floodplain. Ecology. 67: 1508-1523.
Walker, L.R., J.C. Zasada, and F.S. Chapin III. 1986. The role of life history processes in primary succession on an Alaskan floodplain. Ecology. 67: 1243-1253.
Wurtz, T.L., and J.C. Zasada. 1987. An exceptional case of natural regeneration of white spruce in interior Alaska. In Current topics in forest research: emphasis on contributions by women scientists. Proc. National Symp. U.S. Dep. Agric. For. Serv., Gen. Tech. Rep. SE-46. pp. 86-88.
Willson, M.F. 1992. The ecology of seed dispersal. In Seeds: the ecology of regeneration in plant communities. M. Fenner (ed.). C.A.B. International, Oxen, U.K. pp. 61-85.
Yarie, J. 1981. Forest fire cycles and life tables: a case study from interior Alaska. Can. J. For. Res. 11: 554-562.
Youngblood, A., and T.A. Max. 1992. Dispersal of white spruce seed on Willow Island in interior Alaska. USDA For. Serv. Res. Pap. PNW-RP-443. Portland, OR. 17p.
Zasada, J.C. 1971. Natural regeneration of interior Alaska forests - seed, seedbed, and vegetative reproduction considerations. In Proceedings, fire in the northern environment, a symposium. Pacific Northwest For. and Ran. Exp. Sta., Portland, Oregon. pp. 231-246.
Zasada, J.C. 1985. Production, dispersal, and germination of white spruce and paper birch and first-year seedling establishment after the Rosie Creek fire. School Agr. Land Resources Manage. Univ. Alaska, Fairbanks, Alaska. Misc. Publ. 85-2: 34-37.
Zasada, J.C., M.J. Foote, F.J. Deneke, and R.H. Parkerson. 1978. Case history of an excellent white spruce cone and seed crop in interior Alaska: cone and seed production, germination, and seedling survival. U.S. Dep. Agric. For. Serv., Pac. NW For. Exp. Sta., Gen. Tech. Rep. PNW-65.
Zasada, J.C., and D.F. Grigal. 1978. The effects of silvicultural system and seed bed preparation on natural regeneration of white spruce and associated species in interior Alaska. In Proc. 5th North American forest biology workshop. C.A. Hollis and A.E. Squillace (eds.). School of Forest Resources and Conservation. University of Florida, Gainesville, FL. pp. 213-220.
Zasada, J.C., and D. Lovig. 1983. Observations on primary dispersal of white spruce, Picea glauca. Can. Field Naturalist. 97: 104-106.
Zasada, J.C., Sharik, T.L., and M. Nygren. 1992. The reproductive process in boreal forest trees. In H. Shugart, R. Leemans, and G. Bonan (eds). A system analysis of the global boreal forest. Cambridge Univ. Press, Cambridge, UK.