EARTH SURFACE PROCESSES AND LANDFORMS Earth Surf. Process. Landforms 34, 1255–1269 (2009) Copyright © 2009 John Wiley & Sons, Ltd. Published online 22 May 2009 in Wiley InterScience (www.interscience.wiley.com) DOI: 10.1002/esp.1813 Sediment transport due to tree root throw: integrating tree population dynamics, wildfire and geomorphic response Chichester, ESP EARTH The 1096-9837 0197-9337 Earth ESP1813 9999 Research Copyright John 2006 Journal Wiley Science Surf. Surface SURFACE Article Articles © Process. & UK of 2006 Sons, Processes thePROCESSES John British Ltd. Landforms Wiley and Geomorphological Landforms AND & Sons, LANDFORMS Ltd. Research Group Sediment transport due to tree root throw J. M. Gallaway,1,2 Y. E. Martin1,2* and E. A. Johnson2,3 Department of Geography, University of Calgary, Calgary, Alberta, Canada 2 Biogeoscience Institute, University of Calgary, Calgary, Alberta, Canada 3 Department of Biological Sciences, University of Calgary, Calgary, Alberta, Canada 1 Received 23 September 2008; Revised 16 February 2009; Accepted 23 February 2009 * Correspondence to: Y. E. Martin, Department of Geography, University of Calgary, Calgary, AB T2N 1N4, Canada. E-mail: [email protected] ABSTRACT: A field study was conducted to analyze root throw and associated sediment transport in Hawk Creek Watershed, Canadian Rockies. A large crown fire in 2003 allowed the opportunity to study pre-fire and post-fire root throw. Based on field data, a significant relation was found between gradient and root plate volume, as well as individual root plate dimensions. Given that tree diameters increase as trees age and that a relation in the field data was found between tree diameter and root plate volumes, sediment transport due to root throw is expected to change in response to forest disturbance and stand age. Sediment disturbance, which is the amount of sediment upheaved during tree topple and does not take into account transport distance, shows higher values on steeper gradients. Sediment transport was notable for the steepest plots, with pre-fire values of 0·016 cm3 cm–1 a–1 and post-fire values of 0·18 cm3 cm–1 a–1. A tree population dynamics model is then integrated with a root throw transport model calibrated for the Canadian Rockies to examine the temporal dynamics of sediment transport. Fire is incorporated as a disturbance that initiates development of a new forest, with the model cycling through generations of forest. Trees fall according to an exponential rate that is based on time since death, resulting in a time lag between tree mortality and sediment transport. When values of time-since-previous-fire are short, trees are generally <13 cm, and minimal sediment is upheaved during toppling. If trees reach a critical diameter at breast height (dbh) at time of fire, a pulse of sediment occurs in the immediate post-fire years due to falling of killed trees, with tree fall rates decreasing exponentially with time-since-fire. A second pulse of root throw begins at about 50 years after the previous fire, once new recruits reach a critical dbh and with initiation of competition-induced mortality. Copyright © 2009 John Wiley & Sons, Ltd. KEYWORDS: tree root throw; sediment transport; wildfire; tree topple Introduction Tree topple, which may involve stem breakage or uprooting, plays a critical role in forest population dynamics (for review see Quine and Gardiner, 2007), with events ranging in size from severe storm tree blow downs of kilometers in length to individual tree death due to competition, insects, or disease and subsequent tree topple due to decay of the supporting root structures or the stem. Root throw is defined as tree uprooting when the root plate is upheaved along with any attached sediment. Root throw is recognized as an important near-surface process affecting infiltration, air capacity, and remixing of organic material (e.g. Lutz, 1940; Meyers and McSweeney, 1995; Clinton and Baker, 2000) and is also an important sediment transporting agent on forested hillslopes (e.g. Dietrich et al., 1982; Swanson et al., 1982; Roering et al., 2002; Gabet et al., 2003; Osterkamp et al., 2006). Root throw results in vertical and horizontal displacement of sediment attached to the roots (called the root plate). The disturbed sediment often remains attached to the root plate for a period of time after root throw. Subsequent root plate disintegration due to weathering and decay of the roots leads to vertical fall of sediment, which may remain in situ or move horizontally and/or vertically due to gravity and inertia. A pitmound pair is the resulting geomorphological feature from this process (Stephens, 1956). Mound degeneration may occur through weathering and transport processes, such as rainsplash or diffusive creep. Subsequent mound disintegration is not considered as part of the root throw process herein. The volume of soil disturbed during a root throw event depends on factors including: tree species and age; whether the tree was alive or dead at time of fall; soil texture; rooting structure and depth; and moisture content of soil at time of the event (Norman et al., 1995). Root plate volumes are smaller for trees that have been dead for some time prior to falling (Cremeans and Kalisz, 1988; Norman et al., 1995; Ulanova, 2000), due to decay of fine roots and reduced cohesion between the root structure and the soil (Swanson et al., 1982). The nature of how the tree topples determines the location of the root plate relative to the pit, which in turn affects the fall 1256 EARTH SURFACE PROCESSES AND LANDFORMS location of disintegrating sediment (e.g. Beatty and Stone, 1986). The root plate may be situated either upslope or downslope of the originating pit (with some lateral component also possible), or may be situated directly above the pit. Hillslope gradient, angle of tree fall relative to contour lines, and final resting location of the root plate are key factors affecting the volume of sediment that ultimately falls either outside or within the pit (Norman et al., 1995). Sediment that falls from the root plate may contribute to downslope sediment transport, but in some cases sediment may be deposited upslope of the pit, or it may be deposited in the pit itself. Finally, the rate of sediment disintegration affects the final transport rate (assuming the transport process is only completed once sediment deposits on the ground surface), with the disintegration rate being influenced by factors such as root plate dimension, sediment particle size, and weathering intensity. How important is root throw relative to other hillslope geomorphic processes? Gabet et al. (2003) estimated sediment transport rates for root throw using a model calibrated with field data, and obtained values of the order 10–3 m3 m–1 a–1. Roering et al. (2002) estimated sediment transport rates due to biogenic processes (e.g. root growth, root throw) of the order 10–2 to 10–3 m3 m–1 a–1. For comparison, these values are close to or somewhat higher than typical soil creep rates reported in other studies [see Martin (2000) for compilation of published creep rates], and lower than mass wasting rates due to shallow landsliding in coastal British Columbia (Martin, 2000; Martin et al., 2002). To the best of our knowledge no published studies have explicitly connected the timing and rates of sediment transport due to root throw to tree population dynamics driven by wildfire disturbance. Furthermore, root throw studies have not been undertaken in the forests of the Canadian Rocky Mountains. The primary objective of this study is to develop a model that integrates tree population dynamics (i.e. tree recruitment, growth, mortality, toppling) with sediment transport due to root throw for this region. To achieve this goal, a field program was undertaken to document root throw occurrences and characteristics, and associated sediment transport, for both pre-fire and post-fire scenarios in the Canadian Rockies. In particular, a knowledge of root plate characteristics in these forests based on field evidence is essential to the calibration Figure 1. of our integrated forest population dynamics/sediment transport model. To study the temporal dynamics of root throw over scales of 103 years, the model cycles through generations of forests and associated root throw as conditioned by crown fire disturbances which cause death of the tree population. Using this integrated model, we strive for improved understanding of how tree population dynamics driven by wildfire disturbance influences the temporal dynamics of sediment transport due to root throw in the Canadian Rockies. Study Area The field program was undertaken in Hawk Creek, Kootenay National Park, south-eastern British Columbia (Figure 1). The region is underlain by folded and faulted sedimentary rocks, which were uplifted during the Cretaceous-Tertiary with elevations ranging from 800 m to 3400 m. The major valleys have been glaciated, resulting in u-shaped main valley floors and hanging valleys for many tributaries. Hawk Creek drainage basin is approximately 24 km2 in area and is a fourth order tributary of the major Vermillion River. Elevation is 1330 m at the confluence with the Vermillion River, and 3086 m in the upper slopes. Hillslope gradients are moderate (generally <30º) in the lower third of the basin, where the study plots are situated, and become steeper (up to 45°–50°) with increasing elevation. Slopes immediately adjacent to Hawk Creek have steep gradients (25°–40°) for much of the main stem length and often constrain development of a floodplain and riparian zone. The lower portion of the basin contains morainal material overlying bedrock and includes bedrock outcrops and small areas of colluvium. The soils are unconsolidated and unsorted, with a significant number of cobbles and boulders within the matrix. The matrix is a silty-sand, with clay content <2% or absent. The temperatures in Kootenay National Park are influenced by cold continental air masses from the north or maritime winds from the west. The continental divide at the northeast boundary allows for greater influence by maritime weather patterns, resulting in a somewhat milder and moister climate than east of the divide. Winter precipitation is largely in the form of snow, averaging about 170 cm annually; summer rainfall is Hawk Creek, Kootenay National Park, British Columbia, Canada. Extent of the 2003 burn is shown in the right-hand figure. Copyright © 2009 John Wiley & Sons, Ltd. Earth Surf. Process. Landforms 34, 1255–1269 (2009) DOI: 10.1002/esp SEDIMENT TRANSPORT DUE TO TREE ROOT THROW frequently delivered by convectional thunderstorms, with an average annual rainfall of about 340 mm. Precipitation increases with elevation throughout the region (Environment Canada, 2005). The vegetation for the Vermilion basin consists of subalpine forest and alpine tundra. The lower subalpine forest is composed primarily of lodgepole pine (Pinus contorta Loudon var. latifolia Engelm.) and Engelmann spruce (Picea engelmannii Parry ex. Engelm.), and the upper subalpine forest of Engelmann spruce and subalpine fir (Abies lasiocarpa [Hook.] Nutt.). The fire season is from May to September, with peak lightning activity in July and August (Masters, 1990). Crown fires are the dominant type of fire. Masters (1990) calculated a return interval for the study area of 75 years for pre-1768 data, and 267 years for post-1768 data. The change in fire interval during the 1700s is related to the Little Ice Age, as has been found for other Rocky Mountain locations (Johnson and Larsen, 1991). These fire return intervals are less than the potential lifespan of the canopy trees (250–300 years, possibly up to 375 years), making wildfire an important determinant of tree population dynamics. The duff layer (F and H organic layers of the soil) in unburned forests of this region is continuous and up to 15 cm thick. As in other Canadian Rocky Mountain locations, crown fires consume large amounts of the duff layer. In late July 2003, lightning ignited two fires in the Vermillion Valley. These fires merged to burn approximately 17 000 hectares (see Figure 1). About 80% of the forested area in Hawk Creek basin was burned by high intensity (crown) wildfire, including much of the riparian zone. Only the very upper part of the drainage basin did not burn. The previous burn date at Hawk Creek was in 1835 (Masters, 1990). Field Methods To estimate rates of root throw and associated sediment volumes, three plots, having areas of 2·5, 4·1, and 3·2 ha (Plots 1, 2, and 3, respectively), were delineated in the lower reaches of the Hawk Creek drainage basin. The plots were located within 150 m of each other, had a southwest aspect and hillslope gradients of 3°, 15° and 28°, but maintained reasonable consistency in most other physical attributes. All field measurements were made after the fire. The collection of pre-fire data for root throw was based on information derived from fallen trees and root plates, which were inferred to have been in place at the time of fire based on certain observed characteristics. Root throw frequency and characteristics were monitored in the first two years following the 1257 fire. Topples caused by a break or snap in the bole were not considered as no sediment upheaval is associated with this type of tree fall event; this study focuses exclusively on root throw events (i.e. tree topples that upheave sediment through an uprooting event). Root plates for fallen trees were surveyed and flagged to allow identification of new topples that occurred between surveys and to monitor root plate disintegration. Root plates were included in the survey if sediment was still attached to the root structure. Four root plate age classes, modified from Brown et al. (1998), were derived for the current study (see Table I). Based on the characteristics listed in Table I, root plates were assigned to one of three pre-fire age classes, or to the post-fire category. In the first-year of field work (2004), post-fire topples were identified by unburned roots at the base of the root plate. All root plates, including those which existed before the fire and those that came into existence due to post-fire toppling, were flagged to allow identification of new topples in 2005. Postfire root throw frequency was monitored by counts of tree falls in the three plots. Forest density defines the number of trees available for toppling, and was estimated by tree counts of standing trees with diameter at breast height (dbh) ≥10 cm in six sample tracts (each ~150 m2) within each plot. Surveys of existing root plates in each plot were performed in the summers of 2004 and 2005. Detailed data were collected for root plates within Age Classes 1–3. Root plates in Age Class 4 were generally more deteriorated, reducing measurement accuracy for bole, root plate, and disintegration rates; therefore, they were not included in detailed field measurements of sediment volumes or root plate disintegration, but were only included in bulk volume estimates. The detailed analysis is as follows. Fall directions for root plate bole and angle of fall relative to local contour were recorded. Fall angles were recorded in Survey Plots 2 and 3, where hillslope gradient was sufficient to allow identification of direction of steepest ascent. The two areal dimensions for a root plate are designated as width and height. Measurements of root plate width (w) were made parallel to the ground surface, and height (h) measurements were taken orthogonal to width. Measurements were normally taken on the underside of the root plate (i.e. the rounded side with newly exposed soil), but in some cases, pit infilling or pit-plate configuration forced measurements to be made on the ground surface side of the root plate. It is important to distinguish between the dimension that is parallel and that which is normal to the ground surface for later sediment transport calculations. The procedure to determine depth (d) of the root plate required two measurements. The first measurement was made by placing Table I. Root throw age classes based on bole description; modified from Brown et al. (1998) to simplify application in a post-fire environment Age class Short description 1 New 2 Recent 3 Deteriorating 4 Old Age estimates at time of first measurement Characteristics Roots at bottom of root plate not burned so tree fell after the fire Fully barked (>80%); bole solid; no checkerboard burn pattern on bole, indicating tree was not dead long prior to burning Bark 0–80%; some sapwood decay but bole generally whole; checkerboard burn pattern on some or all of bole Sapwood flaking, easily removed; settling of stem or flattening of circumference; checkerboard burn pattern on all of bole Post-fire 0–2 years before fire 2–30 years before fire 30–90 years before fire Note: Bole description includes amount of bark on the bole, burn pattern, and amount of soft or rotting bole. Copyright © 2009 John Wiley & Sons, Ltd. Earth Surf. Process. Landforms 34, 1255–1269 (2009) DOI: 10.1002/esp 1258 EARTH SURFACE PROCESSES AND LANDFORMS Figure 2. (A) Area dimensions associated with a root plate. View is shown looking at underside of root plate. (B) Left-hand photograph shows area dimensions for a root plate. Right-hand photograph shows depth of a root plate. Photograph by J. Gallaway, Kootenay National Park, British Columbia, June 2004. a rod at right angles to the bole where the ground surface intersected the trunk, and measuring from this rod to the plane estimated to pass through the outside edge of the roots. A second depth measurement was made from the rod to the maximum depth of the root plate. The difference between these two measurements defines depth for the half ellipsoid. This approach excludes sediment that may exist in the root plate directly beneath the bole, which may partially compensate for unrealistic smoothness of the half ellipsoid surface that may overstate the volume in the lower portion of the root plate. For asymmetrical root plates with considerable differences in depth across the plate, multiple depth measurements were taken and averaged. The half-ellipsoid model of Denny and Goodlett (1956) and Norman et al. (1995) was used to calculate the root plate volume (Figure 2): VRP = 2 ⎛w h d ⎞ π × × 3 ⎜⎝ 2 2 2 ⎟⎠ (1) where VRP is volume of root plate (in m3), w is width (in meters), h is height (in meters), and d is depth (in meters). Visual estimation was made of the percentage (0, 25%, 50%, Copyright © 2009 John Wiley & Sons, Ltd. 75%, 100%) of root plate sediment still attached to the root structure and the percentage (0, 25%, 50%, 75%, 100%) of root plate sediment that would eventually fall outside of the pit. Estimates were limited to these values to facilitate consistency and repeatability, and were performed by multiple personnel in the field. Older root plates were only included in total volume estimates (i.e. volume estimates summed for all root plates), and were classified based on width of root structure (small: <75 cm, medium: 75–150 cm, large: >150 cm). In addition to the collection of root plate data, a meteorological station recorded precipitation, wind speed, and wind direction over the study period. Additional data recorded for each tree topple included: (i) position on hillslope within the plot (bottom third, middle third, upper third); (ii) bole dbh (taken at 1·4 m above ground); and (iii) tree species. Field Results and Analysis Root throw frequency and direction The detailed survey included 166 root plates, with a further 143 older root plates classified as small/medium/large. No notable Earth Surf. Process. Landforms 34, 1255–1269 (2009) DOI: 10.1002/esp SEDIMENT TRANSPORT DUE TO TREE ROOT THROW 1259 Figure 3. (A) Rose diagram showing tree fall direction in degrees. (B) Histogram showing frequency of uphill and downhill topples. A value of 0º represents an uphill fall direction (perpendicular to hillslope contour lines) while a value of 180º represents a downhill fall direction, with other values falling in between. trends are found in the frequency rate per hectare of root throw amongst the three gradient classes (Table II). When tree density is taken into consideration, there are still no obvious patterns in the data amongst the gradient classes. The data are then split up and analyzed according to pre-fire and post-fire root throw events (Table III). Data for pre-fire events show no notable increase in rates with average gradient, with values ranging from 0·29 to 0·37 events annually per hectare. The post-fire data indicate increasing root throw frequency with gradient. The disparity between pre-fire and post-fire rates increases with increasing gradient, showing notable differences for the steepest gradient class of 28º. Root throw data show a mean fall direction of 29º (Figure 3A). The wind direction recorded during the study period was most frequently between 20° and 30°, a range that includes the mean direction of tree fall. This wind direction across local topography is predominantly uphill, and the effect is apparent in the distribution of tree fall angles relative to contour lines (Figure 3B). Approximately 65% of the tree falls were uphill relative to local contour lines. Root plates When all root plates are considered, the relation between dbh and root plate volume is (Figure 4): Table II. Topple survey plot 1 2 3 Figure 4. Relation between diameter at breast height (dbh) and root plate volume. Data are shown for Age Classes 1–3 (32-year period). VRP = (−5·74 × 10−4 × dbh) + (5·82 × 10−3 × dbh2) R2 = 0·45 (2) where VRP is root plate volume (in m3) and dbh is diameter at breast height (in cm). The mean value of root plate volume increases from a value of 0·2 m3 to 0·71 m3 (p < 0·01) as gradient becomes steeper (Table IV). Width and height of the root plate were defined earlier as width being parallel to the ground surface and height being Root plate counts for the three survey plots Average gradient (deg) Plot area (ha) Tree density (# ha–1) Root plate count Root plates per hectare (# ha–1) Frequency (% ha–1) 3 15 28 2·5 4·1 3·2 1665 1882 2200 85 112 112 34·0 27·3 35·0 2·0 1·5 1·6 Note: Root plate counts relative to the tree density for each plot are shown in the last column. Table III. Topple survey plot 1 2 3 Annual topple rates by gradient Average gradient (deg) Plot area (ha) Pre-fire topple rate (# ha–1 yr–1) Pre-fire topple rate (% yr–1) Post-fire topple rate (# ha–1 yr–1) Post-fire topple rate (% yr–1) 3 15 28 2·5 4·1 3·2 0·37 0·29 0·35 0·022 0·015 0·016 0·40 0·60 1·7 0·024 0·032 0·078 Note: Time span for pre-fire root plates is 90 years; time span for post-fire root plates is two years. Rates are presented per hectare, and as percent of standing tree density. Copyright © 2009 John Wiley & Sons, Ltd. Earth Surf. Process. Landforms 34, 1255–1269 (2009) DOI: 10.1002/esp 1260 Table IV. EARTH SURFACE PROCESSES AND LANDFORMS Mean values of root plate dimensions for the three survey plots Gradient class 1 (n = 45) 2 (n = 61) 3 (n = 60) Root plate volume (m3) Width (cm) Length (cm) Depth (cm) Ratio of length to width 0·22 (0·11) SE = 0·044 0·35 (0·18) SE = 0·058 0·71 (0·62) SE = 0·072 91·2 (80) SE = 6·0 115·5 (106) SE = 6·9 150·4 (155) SE = 5·6 117·1 (110) SE = 6·8 160·0 (145) SE = 10·7 219·5 (223) SE = 9·3 26·9 (25) SE = 1·9 29·7 (28) SE = 1·9 39·3 (40) SE = 2·1 1·33 (1·24) SE = 0·043 1·42 (1·30) SE = 0·0505 1·49 (1·43) SE = 0·0488 Note: Median values are given in brackets, and the standard error (SE) is indicated. normal to the ground surface. However, in that definition either width or height could be the larger of the two dimensions, likely meaning that empirical analysis of relations for changes in a particular dimension with increasing slope gradient would be obscured. Therefore, for this particular analysis and table (Table IV, columns three to five), the shorter dimension of the two is referred to as ‘width’ and the longer dimension as ‘length’; the depth dimension remains the same as defined earlier. Increases in magnitude with slope gradient for each individual dimension (width, length and depth) were found to be significant at p < 0·01. On slopes having a gradient >0°, mechanical stress is expected to be unevenly distributed around the tree diameter and the root system. Root growth may respond by increasing the density of the root system or by arranging roots in an asymmetric manner in the direction of maximum stress, both of which serve to provide increased anchorage for the tree and greater stability (Soethe et al., 2006; Coutts et al., 1999). Unfortunately, our data do not allow us to determine the direction in which our root plates were oriented when originally in the ground. Nonetheless, it is still of interest to assess changes in the ratio of length to width for root plates measured in the field. A significant increase is found for the ratio of length to width with increasing slope gradient (Table IV, column six), lending some support to the idea that asymmetry of the root plate will become more pronounced as gradient increases. Disintegration rates of root plates were determined by estimating percentages of volume removed from the root plates in 2004 and 2005 (Table V) for the three age categories (1–2 years post-fire, two years pre-fire and 30 years pre-fire; refer back to Table I). As of 2004, the youngest (most recent) root plates had similar percentages of volume removed from the root plate as the older category. Furthermore, in 2005 it was these youngest root plates that had lost the greatest amount of volume in the intervening period. For both pre-fire and post-fire scenarios, the steepest-gradient plots showed the greatest rates of sediment disintegration. A greater proportion of sediment associated with root plates falls outside the Table VI. Table V. Root plate disintegration averages by age class Age class Age maximum 1 2 3 1–2 years post-fire 2 years pre-fire 30 years pre-fire Average volume off 2004 (%) Average volume off 2005 (%) 48·3 33·3 46·7 70·8 47·7 54·0 Note: Data are for detailed root plates only. originating pit on steeper gradient slopes relative to gentler slopes, possibly due to a greater occurrence of rotational falls on steeper slopes (Table VI). For both pre-fire and post-fire root plates the average percent falling outside the pit increases from negligible values for relatively flat land surfaces, through to 20–30% for the mid-slope category and up to about 50% for the steepest gradients. Areal sediment disturbance and soil turnover The total area of ground surface disturbed by tree upheaval (i.e. total area disturbed by all events occurring during a specified time period) was based indirectly on the root plate measurements and covers all four age categories, or a period of approximately 92 years (Table I). The area subject to tree root upheaval increases with slope gradient, with annual percentages of disturbed land surface ranging from 0·003% to 0·006% (Table VII). Sediment disturbance and transport Amounts of sediment disturbance include the volume of all sediment that is uprooted, whether it is eventually returned to the pit or not. Sediment returned to the pit during root plate disintegration contributes to weathering and breakdown Root plate disintegration data for the three survey plots Topple survey plot Root plate counts Average percent falling outside pit Average percent off root plate 2004 Average percent off root plate 2005 Pre-fire root plates 1 2 3 43 56 49 5·0 30 52 46 46 33 54 53 53 Post-fire root plates 1 2 3 2 5 11 0·0 20 50 38 68 55 38 80 73 Note: Pre-fire and post-fire root plates are analyzed separately. Copyright © 2009 John Wiley & Sons, Ltd. Earth Surf. Process. Landforms 34, 1255–1269 (2009) DOI: 10.1002/esp SEDIMENT TRANSPORT DUE TO TREE ROOT THROW Table VII. Survey plot 1 2 3 Total pit area disturbed and annual rate of pit formation Average gradient (deg) Total pit area (m2 ha–1) Annual total pit area (%) 3 15 28 28·7 36·0 59·6 0·003 0·004 0·006 of the soil layer, while sediment that disintegrates and lands outside the pit contributes to sediment transport. The transport distance of sediment after uprooting becomes important in our calculations of sediment transport (see Equation 3) and is not a consideration when estimating sediment disturbance volumes. Sediment disturbance rates over a 32-year period (time period for root plates of Age Classes 1–3) are evaluated to assess if there is a gradient dependency on the total amounts of uprooted sediment over this period. Results do not show a notable difference in total volume uprooted per square meter of hillslope for the two lowest-gradient study plots, with values of 1·25 × 10–5 and 1·63 × 10–5 m3 m–2 a–1 respectively (Table VIII). However, the steepest plot, Plot 3, does show a notably higher value of 4·15 × 10–5 m3 m–2 a–1 compared to the other plots, in large part due to the higher root plate volumes associated with toppled trees at steeper gradients as discussed earlier. We now consider sediment transport rates due to root throw. It is assumed that sediment falling into the pit undergoes no net transport, and thus only sediment that is involved in forming mounds is considered in this analysis. A diffusive approach to sediment transport may be appropriate if root throw is considered to be a relatively slow, quasi-continuous process with a dependency on hillslope gradient (Norman et al., 1995), and with the potential to operate across the entire forested portion of the landscape. Adopting such an approach allows for comparison with transport rates for other processes involved in medium-term drainage basin evolution (note that the temporal dynamics of root throw transport will be explored in more detail in the modeling section of this paper). The equation used to calculate the sediment transport rate for a plot of a given gradient is (adapted from Martin and Church, 1997): Σ(Vmd ) qs = (3) Apt where qs is sediment transport rate (in m3 m–1 a–1), Vm is volume of sediment landing outside the pits and which form mounds (in m3), d is distance (in meters) that sediment is ‘transported’ along the ground (the net ground-parallel distance travel after the sediment has been upheaved and falls to the ground), Ap is area of survey plot (in m2) and t is the estimated number of years that the root plates have existed. Volume is calculated as the product of original root plate volume, the percentage that had fallen off the root plate, and the percentage falling outside the pit (this value must be calculated for uphill topples and is assumed to be 100% for Table VIII. Survey plot 1 2 3 1261 downhill topples; see explanation later). Sediment transport was calculated for each survey plot, providing a transport rate for three different gradients. This analysis is completed for root plates of Age Classes 1–3, and thus t is 32 years. Sediment transport distance was calculated using geometric models based on field measurements, similar to the approach of Gabet et al. (2003), but with two major differences. First, the current study does not consider the entire root plate volume in the transport calculations, but excludes the volume of root plate sediment that will fall back into the pit. Second, this study examines uphill and downhill sediment transport separately, rather than calculating a single net transport value. The model involves two components contributing to transport distance: (i) slope-parallel transport distance during upheaval; (ii) slopeparallel transport distance during root plate disintegration. Two assumptions are required for this model: (i) root plates are assumed to be circular, with the dimension of the root plate perpendicular to the ground representing diameter; and (ii) the root plate comes to rest with the center of mass at the edge of the pit (Figure 5A). The ground-parallel transport distance for downhill topples is now considered. The net downslope distance for upheaval associated with downhill topples is the distance between two ‘contour lines’ situated across the slope, one which crosses through the center of the pit and the other crossing through the center of the root plate mass. This plan view distance is converted into true distance by making an adjustment for slope gradient. During the disintegration phase, all sediment for downhill topples is assumed to fall outside the pit, with simple geometric calculations used to calculate the net downslope transport distance for this stage of the root throw process. The net ground-parallel transport distance used in calculations is shown in Figure 5(B). Similar geometrical considerations are applied to the model for uphill topples, but it cannot be assumed that all sediment falls outside the pit. In addition, there is now both an uphill and downhill component to the transport process, as the upheaval moves sediment uphill, while disintegration moves sediment downhill; both must be accounted for to determine if there is net uphill or downhill transport. Once again, geometric models are applied to determine the amount of sediment associated with the root plate that will fall outside the pit. A line in the direction of steepest gradient is imposed tangentially to the rim of the pit. The intersection of this line with the root plate half-ellipsoid (sitting at some angle above the pit) determines the proportion of sediment that will ultimately fall outside the pit; only this sediment is used in transport calculations (sediment landing in the pit is not considered to be part of the transport process). In Survey Plot 1, where gradient was too low to accurately determine contour line direction, only the disintegration transport distance was used, resulting in understated transport distances. The impact of this is expected to be small, as very little sediment is transported by root throw (i.e. very little sediment falls outside the originating pits) on low gradient slopes. Sediment disturbance values Average gradient (deg) Area of plot (ha) Volume uprooted over 32 years (m3) Annual volume disturbed (m3 m–2 a–1) Average depth (mm a–1) 3 15 28 2·5 4·1 3·2 10·01 21·45 42·49 1·25 × 10–5 1·63 × 10–5 4·15 × 10–5 0·013 0·016 0·041 Note: Sediment disturbance includes estimates of all sediment that was originally uprooted, although some of it may have since disintegrated and formed part of a mound. Copyright © 2009 John Wiley & Sons, Ltd. Earth Surf. Process. Landforms 34, 1255–1269 (2009) DOI: 10.1002/esp 1262 EARTH SURFACE PROCESSES AND LANDFORMS Figure 5. (A) Circular pit and root plate, with center of mass of root plate situated at edge of pit. (B) In the model, the first stage of sediment transport involves the movement of sediment from what becomes the pit to the new location of the root plate after tree topple. The second stage of transport involves the vertical disintegration of sediment to the ground, with the distance for this stage defined as the distance between the root plate centroid to the location where vertically falling sediment reaches the land surface (shown by the bold arrow). The double-headed arrow shows the ground-parallel net transport distance associated with root throw transport. Uphill and downhill sediment transport rates during the pre-fire period are shown for root plates categorized in Age Classes 2 and 3 (Figure 6A); sediment is considered as having been transported once it falls to the ground from the root plate. Sediment transport rates are also calculated for the two post-fire years (Figure 6B); these data include trees that toppled during both the post-fire and pre-fire periods (Age Classes 1–3), but for which sediment actually fell to the ground in the post-fire period. We first discuss the downslope transport rates. Negligible sediment transport occurred on the lowest gradient plot for the pre-fire period, as sediment was usually returned directly to the pit. Moreover, transport rates remained negligible for the mid-gradient plot, and it is only for our upper-gradient plot, having a slope of 28°, that notable increases in sediment transport rates were observed. There appears to be a non-linear form to the plot; however, with so few data points, broad conclusions should not be made. The post-fire results show the same pattern in the data, with the major difference being an approximately oneorder-of-magnitude increase in sediment transport for the post-fire scenario versus the pre-fire scenario. Upslope transport rates for both periods show negligible values on the low-gradient field plot, with somewhat higher values for the steepergradient field plots. Figure 6. Downslope and upslope sediment transport plots. (A) Transport results for the pre-fire period. (B) Transport results for the two post-fire years. Note scale differences on the two graphs. Copyright © 2009 John Wiley & Sons, Ltd. Earth Surf. Process. Landforms 34, 1255–1269 (2009) DOI: 10.1002/esp SEDIMENT TRANSPORT DUE TO TREE ROOT THROW Model Outline 1263 for fire intervals expresses the probability of having fires with inter-fire intervals of length t (Johnson and Van Wagner, 1985): Model overview While the field results provide an indication of sediment disturbance and transport rates due to root throw, the temporal dynamics of root throw-driven transport is directly related to the population dynamics of forests. To obtain greater insights into these temporal dynamics, we develop a combined tree population dynamics/sediment transport model that is calibrated for the central Canadian Rockies. In simple terms, this model germinates and grows trees, kills trees, and has these trees fall to the ground. A portion of these fallen trees generates root throw events. Fire events occur as a disturbance that kills all trees and results in a new forest. Thus, the model cycles through forest generations with life spans determined by the variability of fire event recurrence. Calculation of the sediment transport by root throw is organized as follows: first, between wildfires the population dynamics of the trees is modeled using the algorithms for recruitment, mortality, and growth in diameter. Next the timing of mortality of all the trees in the population from crown fire is determined based on information about fire return intervals. The tree falling (topple) rate and chance of uprooting versus breakage of the dead trees, either from fire or from inter-fire mortality, are determined. Finally, the amounts of sediment associated with root plates of toppled trees are determined and the transport rates are calculated. A list of model parameters is given in Table IX. Fire frequency The lifespan of forest trees in the study region is largely determined by fire return intervals and type of fire. Mean fire return interval in a region is estimated as the expected number of years between fires. The Weibull probability density function Table IX. Input parameters for model simulations Number of years of simulation Plot area Hillslope gradient Average fire return interval (scale parameter) (Equation 4) Shape parameter for fire (Equation 4) Duration of the fire cohorta Between-fire falling rates (F in Equation 5)b Fire cohort Understory cohort Post-fire falling rates for all fire killed trees Fraction uprooted versus broken boles Parameters to assign dbh distributions for trees of certain age Scale parameter Shape parameter dbh threshold for uprooting mass 1000 years 100 m2 20° 110 years 1·0 10 years 0·058c 0·058 0·084d 0·8 Mean dbh values (Figure 8) 1·8 13 cm a The user input for span of fire cohort will determine when the canopy is formed and new recruits are going into the understory cohort rather than the fire cohort. b Fire ‘cohort’ is defined as those trees that germinate after fire and form the canopy. Understory ‘cohort’ is defined as those trees that germinate after the canopy is formed. c The value of 0·058 is based on data for Johnson and Greene (1991) and Johnson (unpublished data) for the Kananaskis Valley. d The value of 0·084 is based on Lyon (1977). Fire killed trees may generate root throw until next fire. Copyright © 2009 John Wiley & Sons, Ltd. γ ⎛t ⎞ f (t ) = ⎜ ⎟ α ⎝α ⎠ γ −1 ⎛t ⎞ −⎜ ⎟ e ⎝α ⎠ γ (4) where t is time interval between two fires (in years), α is a scale parameter (expected fire return interval in years) and γ is a shape parameter (dimensionless). For this modeling exercise, the shape parameter γ is set to one, resulting in a negative exponential distribution, which studies in the area have shown to provide a good fit to the data (Masters, 1990; Johnson and Larsen, 1991; Reed et al., 1998). A value of 110 years is chosen for the variable α. Random numbers are selected from the Weibull distribution, which represent inter-fire time intervals for the model run. Tree population dynamics In the subalpine forests of lodgepole pine (Pinus contorta Loudon var. latifolia Engelm.) and Engelmann Spruce (Picea engelmannii Parry ex. Engelm.) found in the region under study (Johnson and Fryer, 1989; Johnson et al., 1994; Johnson et al., 2003), two groups of trees contribute to tree toppling and sediment transport. The first group consists of trees either killed by the last fire or dead standing before the last fire, and the other group consists of trees that are recruited after the last fire. The first group represents a cross-section of the live trees and standing dead trees at the time of the last fire, showing a range of diameters. The second group considers the recruitment and mortality of trees in two kinds of cohort. A cohort is a group of trees recruited at the same time and following a similar mortality schedule throughout their lives. The fire cohort is defined as those trees recruited in the fiveto 10-year period after the fire in which the canopy was killed (Johnson et al., 2003) and the forest floor was removed by smoldering combustion (Charron and Greene, 2002; Miyanishi and Johnson, 2002). The understory cohorts are those that begin to grow under the fire cohort’s canopy and on a forest floor occupied by groundcover. Both the recruitment (Figure 7A) and mortality rates (Figure 7B) used in our simulations are from Johnson and Fryer (1989) and Johnson et al. (2003). The duration of the fire cohort is the period of time for canopy trees to become established, and in our model its duration is assigned a value of 10 years. Afterwards, all new recruits are considered to be part of the understory cohort and, as seen in our recruitment curve, it generally has a lower recruitment rate. Mortality rates are specified separately for each cohort (Figure 7B). The understory cohort has a higher mortality rate than the fire cohort. Mortality rates are related to age of trees, and within one time interval different mortality rates are applied to trees of different ages and different cohorts. Root throw events Two groups of trees contribute to root throw and transport: (i) trees either killed by fire or dead standing before fire; and (ii) trees recruited after fire with different mortality rates for the two cohorts. Field data (Johnson, 1986, unpublished data) show that topples occurring in the first several years after fire of fire-killed trees or dead standing trees at the time of the fire result from having their root support weakened by removal of Earth Surf. Process. Landforms 34, 1255–1269 (2009) DOI: 10.1002/esp 1264 EARTH SURFACE PROCESSES AND LANDFORMS Figure 7. (A) Recruitment curve used in model runs. Trees recruited within the first 10 years after a fire are part of fire cohort and any trees recruited thereafter are part of understory cohort. (B) Mortality schedule for understory and fire cohorts. Based on Johnson and Fryer (1989) and Johnson et al. (2003). organic matter around their bases (Miyanishi and Johnson, 2002) and burning of roots during the fire. Subsequent postfire tree topple is due to loss of root strength (Martin and Johnson, 2004–2008, unpublished data). Dead trees will fall by uprooting or breakage. Toppling of standing dead trees occurs according to an exponential model (e.g. Lyon, 1977, who used this model for post-fire topples), whereby a constant fraction of the remaining standing dead boles falls in any given year. The fraction of dead trees that topple in a given time interval is obtained from Equation 5: FT = 1 − exp(−F ∗ dt) (5) where F is a parameterization based on field studies of fraction dead standing trees that topple and dt is the time interval (in years) used in our model runs. In our model, parameterization was based on falling rates taken from studies by Johnson and Greene (1991), Johnson (1986, unpublished data) and Lyon (1977) (see Table IX for further details). The calculated fraction is applied to the remaining standing dead boles for each time interval to determine the number of trees that topple in that period. Empirical surveys show that in our study area about 80% of trees uproot as opposed to breaking, and thus constitute root throw events (Johnson, unpublished data); this value is incorporated into the model to obtain the actual number of root throw events after the number of tree topples has been calculated. Tree fall directions are random in the model runs reported herein. Root plate volume, necessary to calculate sediment disturbance and transport, is dependent on tree size, which in turn is related to tree age when the root throw event occurred (recall that a tree topple is only considered a root throw event when sediment upheaval is involved). The distribution range for dbh is influenced by tree density in the forest (Harper, 1977); higher density forests have smaller dbh values. Tree density is dependent on factors such as soil conditions, tree species, topography, climate, and probably other factors. The dbh distribution in the present model is based on a density of 2000 trees per hectare as found in our field plots [refer to our field data and also Smithers (1961)], and requires adjustment if applying the model to different locations. For an event to be included in our sediment disturbance and transport calculations, the relevant tree must have a diameter >13 cm. This value provides an important condition that must be met in the model for notable sediment upheaval to occur when a tree topples (Johnson, unpublished data). Copyright © 2009 John Wiley & Sons, Ltd. To assign dbh in the model, mean tree diameter is first determined for a tree having a certain age in the model (Figure 8). To ensure a more realistic distribution of diameters for trees of a certain age, a two-parameter Weibull distribution is used to assign actual diameters in the model, with the scale parameter defined as the mean diameter for trees of that age and the shape parameter being assigned a value of 1·8 (tree diameter data are based on unpublished data of Johnson for the Kananaskis Valley, Alberta). Root plate volumes Two pieces of data provide the basis to assign root plate characteristics for each root throw event in the model: the age of the tree when it died and its dbh (see earlier), and the year of fall. Root plate volume is based on dbh using the regression equation obtained from field data for Hawk Creek (Gallaway, 2006) (see Equation 2). Sediment transport In the model, disintegration distributes the transfer of sediment from the root plate to the ground over 100 years and is calculated based on an empirical best fit to our field data (Gallaway, 2006): Figure 8. Mean diameters (dbh) for trees of a particular age. To ensure a realistic distribution of diameters for each age of tree in the model, a Weibull distribution is used in conjunction with the mean diameter to assign a range of diameters. Based on unpublished data by Johnson for the Kananaskis Valley, Alberta. Earth Surf. Process. Landforms 34, 1255–1269 (2009) DOI: 10.1002/esp SEDIMENT TRANSPORT DUE TO TREE ROOT THROW PV = −0·1029 + 46·19e−0·719t + 56·35e−0·02102t (6) where PV is the percent volume remaining and t is time in years. The exponential form of the equation suggests that the disintegration rate may stay constant as the volume decreases over time. The volume falling off a root plate in a time interval is the difference in the percent remaining as calculated for two points in time. The volume of interest for transport is the volume of sediment that falls outside the originating pit; this is dependent on the fall angle of the tree (i.e. if a tree falls directly upslope, it is assumed all sediment will return to the pit). Conversely, if a tree falls directly downslope, all sediment will fall outside the pit. The proportion of sediment falling outside the pit is calculated as: PS = FA/180 wRP = 0·65 + 4·65dbh R2 = 0·62 (8A) hRP = 29·24 + 0·54wRP R2 = 0·62 (8B) where dbh is diameter at breast height (in cm), wRP is root plate width (in cm), and hRP is root plate height (in cm). Transport for a particular time interval is calculated for the sum of sediment volume falling off all existing root plates in that time interval in conjunction with its transport distance. n ∑ (VRP *distRP ) i =1 At (9) where qs is sediment transport rate (in m3 m–1 a–1), VRP is volume of sediment disturbed for a root plate during a particular event (in m3), distRP is travel distance associated with that event (in meters), and At is the area of the plot being modeled (in m2). Model Results Results for an example millennial-scale model run are presented to highlight trends in temporal variations in tree populations and associated tree mortality, tree toppling, root throw, sediment upheaval, and disintegration. Values of the parameters used in the model run are given in Table IX. Fire-killed or standing dead trees at time of fire Fires are the critical disturbance process in our modeled forest, as fire events control the temporal dynamics of tree mortality, tree toppling, and root-throw sediment processes. Random Copyright © 2009 John Wiley & Sons, Ltd. Times of fire in example model runa Time of fires (years since start of model run) 50 130 235 280 580 790 805 870 920 a Model assumes a new forest begins to grow at time 0. No fire-killed trees exist at this time as no previous forest exists. (7) where PS is the proportion of sediment falling outside the pit and FA is the falling angle of the tree. Transport distance is the net slope-parallel distance after upheaval and disintegration. Direction of this net distance may be: (i) downslope; (ii) upslope, if upslope upheaval distance exceeds downslope disintegration distance; or (iii) back into the pit. Ground-parallel transport distance for sediment falling off a root plate is based on a series of geometric calculations similar to those described for the field data, and which requires a knowledge of root plate height (the pit is assumed to be circular with a diameter having this same value). The height dimension needed to calculate transport distance is obtained from the following equations: qs = Table X. 1265 fire distributions (see Equation 4) for this model run show a range of fire return intervals (time period between successive fires), with intervals as short as 15 years and up to 300 years (Table X). Figure 9 illustrates examples of tree age distributions at the time of fire for several fires with different lengths of time since the previous fire. The numbers and sizes of trees at the time of fire are a significant determinant of post-fire sediment disturbance and transport. The duration of time since the previous fire is the time period available for tree growth, and the trees are then subjected to fire-driven mortality and subsequent toppling. The number of new root throw events represents trees that not only have died and toppled, but that also have upheaved sediment (Figure 10). Important to note is that trees do not immediately topple after they die, but rather they follow an exponential toppling rate based on time since death. Therefore, we expect a time lag between tree mortality, toppling, and associated sediment disturbance and sediment transport processes. When the time interval since the previous fire is short, very large numbers of trees will be standing at the time of fire, but their dbh values will be small due to their young age and the dbh of most trees is below the threshold of 13 cm for sediment upheaval (refer to Figure 9A). Hence, the mortality and eventual toppling of these fire-killed (or dead standing) trees involve very small amounts (or no) sediment and they are not included as root throw events. For example, the fire at Year 805 represents a disturbance having a short value of timesince-previous-fire, and no root throw events for fire-killed trees occur immediately following the fire (refer to Figure 10). When the fire return interval exceeds the time needed for trees to reach the critical dbh, the fire-killed trees contribute to root throw events and to sediment disturbance (refer to Figure 9E). An example of this is the fire occurring at Year 580, which shows a notable post-fire increase in root throw events (refer to Figure 10). In such cases, both the number of trees above the critical dbh and their sizes are important in determining the total amounts of sediment upheaved. Once the time interval since the previous fire exceeds approximately 125 years, the time when tree mortality of fire cohorts decreases to very low levels in our model simulation, the actual number of trees in the plot remains approximately constant (refer to Figures 9D and 9E). However, these trees continue to grow in size, and the amount of sediment involved in the toppling of the trees will likewise increase. Sediment disturbance is directly tied to the number of new root throw events and, to a large extent, should follow the patterns of new root throw events for fire-killed or dead standing trees at the time of fire (Figure 11), as well as patterns of mortality and toppling as the forest establishes Earth Surf. Process. Landforms 34, 1255–1269 (2009) DOI: 10.1002/esp 1266 EARTH SURFACE PROCESSES AND LANDFORMS Figure 9. Tree age distributions at time of fire. The distributions are shown for fires at the following times. (A) 805 years (previous fire interval of 15 years); (B) 870 years (previous fire interval 65 years); (C) 235 years (previous fire interval of 105 years); (D) 790 years (previous fire interval of 210 years); (e) 580 years (previous fire interval of 300 years). Figure 10. New root throw events for model run. Counts are for five-year bins. Years of fire events are shown by triangular symbols. Figure 11. year bins. Volume of sediment disturbance for model run. Years of fire events are shown by triangular symbols. Results are grouped into five- itself in between fire events. Sediment is only considered as having been ‘transported’ once it disintegrates, with disintegration not being instantaneous but rather a process that is extended in time. Therefore, while we expect a broadly similar Copyright © 2009 John Wiley & Sons, Ltd. pattern for sediment transport as for sediment disturbance, the disintegration process results in a slight dampening and extending out of the temporal patterns that are found for sediment disturbance (Figure 12). Annual rates of sediment Earth Surf. Process. Landforms 34, 1255–1269 (2009) DOI: 10.1002/esp SEDIMENT TRANSPORT DUE TO TREE ROOT THROW Figure 12. year bins. 1267 Annual sediment transport rates for model run. Years of fire events are shown by triangular symbols. Results are grouped into five- Table XI. Frequency table of sediment transport rates for model run of duration 103 years Annual sediment transport rate (m3 m–1 a–1) 0–0·0005 0·0005–0·0010 0·0010–0·0015 0·0015–0·0020 0·0020–0·0025 0·0025–0·0030 0·0030–0·0035 0·0035–0·0040 0·0040–0·0045 0·0045–0·0050 0·0050–0·0055 >0·0055 Frequency 128 15 18 9 9 5 7 2 2 2 1 3 transport show marked variability that changes in response to the tree population dynamics. The frequency distribution of annual transport rates for our millennial-scale model run is shown in Table XI. The mean value for all annual transport rates is 0·0012 m3 m–1 a–1, with a standard deviation of 0·0047 m3 m–1 a–1. Values range from 0 m3 m–1 a–1 to a maximum of 0·059 m3 m–1 a–1. Understory and fire cohorts After fire occurrence, fire-killed or dead standing trees begin to topple. During this same period, new recruits begin to populate the stand in large numbers, and mortality of these trees also occurs in relatively large numbers. After about 50 to 60 years, a second pulse of root throw events begins to occur for several reasons. Firstly, the mortality rate of trees in the fire cohort increases significantly after about 60 years (see Figure 7B). Furthermore, trees that have reached this age may begin to have dbh values that are >13 cm (critical dbh for sediment upheaval). Once dbh values reach 13 cm (which occurs on average after 80 years, although the random component for tree dbh in the model allows some trees to reach this size several decades sooner), they begin to upheave notable amounts of sediment as they topple. An example can be observed in Figures 10 and 11, with notable sediment disturbance beginning at about 50 years and increasing for the next several decades after the fire event at Year 805. Prior to these conditions being met, trees in the two cohorts that are subject to mortality have little impact on sediment disturbance or transport as they are too small to upheave notable amounts of sediment. Therefore, in addition to a pulse of sediment upheaval due to fire-killed or dead standing trees in the immediate post-fire years, at about 50 years onwards one Copyright © 2009 John Wiley & Sons, Ltd. may begin to observe a second pulse of sediment upheaval and associated transport (Figures 11 and 12), with this pulse expected to last several decades, after which time tree mortality decreases to background levels. Discussion and Conclusions Sediment transport rates by root throw have been estimated in some early studies on this topic (Denny and Goodlett, 1956; Reid, 1981; Mills, 1984). However, the connection between root throw and forest population dynamics, with the latter driving the former by the process of tree toppling and root upheaval, was not fully explored in these studies. Gabet et al. (2003) expanded on earlier work by developing a model to predict medium-term rates of root throw transport as a function of hillslope gradient. Their model accounts for tree topple rates, with the assumption that uprooting rates in the forest are temporally constant at a rate of four trees per hectare per annum. Their model utilizes results of previous field studies (e.g. Norman et al., 1995) to obtain values for root plate dimensions, and uses this information in conjunction with a series of geometrical relationships pertaining to tree fall to estimate transport rates. Some additional assumptions were made in their model, such as values of dbh do not change over time as trees age, and therefore root plate volumes also do not change with time. The results of Gabet et al. (2003) provide first-order approximations of medium-term sediment transport rates due to root throw and mark notable progress in our understanding of this process. Without an explicit connection to forest disturbance and tree population dynamics, the temporal dynamics of the root throw process cannot be fully realized. To our knowledge our study represents the first attempt to explicitly combine a model of root throw with a detailed rendering of tree population dynamics to explain how ecological forcing drives the temporal aspect of sediment transport by root throw. Because there were no existing regional field data/observations of root throw on which to base model calibrations and help make inferences for our work in the Canadian Rockies, it was necessary to include a field component in our study. Tree topple rates may increase in the immediate post-fire years as root support is weakened by the removal of organic matter around the tree and due to the burning of roots during the fire. The marked post-fire increase in toppling for the steep gradient class relative to the less steep gradient classes found in our field data may occur because the structural stability of trees is lower on steeper slopes (Quine and Gardiner, 2007). Our field data identified the relation between dbh, which increases as trees age, and root plate volumes in our region; such data is critical to integrate sediment transport with tree stand ages in our model. The field data demonstrate a significant dependency between gradient and root plate volume and Earth Surf. Process. Landforms 34, 1255–1269 (2009) DOI: 10.1002/esp 1268 EARTH SURFACE PROCESSES AND LANDFORMS all individual dimensions (width, length, depth). An increase in the ratio of length to width was found at steeper gradients relative to less steep slopes (significance level of p < 0·05). It may be the case that, as hillslopes become steeper, tree roots spread perpendicular to contour lines to impart greater stability to the tree (Coutts et al., 1999). Further investigation of this idea is suggested, as such information would allow for improved understanding of the resistance offered by trees to mortality-induced toppling, wind events or landsliding. The pit-mound features formed by tree uprooting and root plate disintegration result in a unique microtopography that contributes small-scale roughness elements to the landscape. These roughness elements may affect depression storage and ponding of water during large storm events, which may affect the timing of hydrological responses in the watershed (Martin et al., 2008). The mound provides an exposed, unconsolidated supply of sediment for rainsplash or other water-driven processes prior to colonization of significant vegetation cover on the feature, while soil creep may act to diffuse the mound both prior to and after vegetation recolonization. It is recommended that future studies explore sediment transport processes operating on pit-mound features to better understand their longevity and contribution to microtopography in forests. Osterkamp et al. (2006) found an average disturbance rate of the soil by root throw of 0·0095% of the surface per year, which is within an order of magnitude of our values. Our data suggest that a complete turnover of the soil surface has not likely occurred in our study area since post-Pleistocene reforestation which began <10 000 years ago. Other studies, summarized in Schaetzl et al. (1989), report disturbance cycles from 220 to 3174 years in eastern North American forests. To allow for comparison with annual depths of sediment disturbance due to root throw for other processes reported in the literature, such as soil creep or landsliding, the volume of sediment disturbed per meters squared of land surface is converted into an annual depth of disturbance by dividing by 32 years, the approximate time period of the root plate upheaval for Age Classes 1–3. Annual disturbance values are of the order 1·0 × 10–2 mm a–1. This result is only one-order of magnitude lower than typical annual disturbance depths reported for shallow landsliding in coastal British Columbia, with a value of about 1·0 × 10–1 mm a–1 (Martin et al., 2002). This suggests that upheaval of sediment due to tree topple is a notable factor operating on forested hillslopes. When considering sediment transport (incorporating both upheaval and disintegration) a notable percentage of disturbed sediment is often returned to the pit, particularly for our lowest gradient plots, and in such situations does not contribute to net transport of sediment. There is approximately a one-order of magnitude increase in sediment transport for the post-fire versus the pre-fire rates for all gradient classes. It is only for our steepest-gradient plot that notable values of sediment transport were observed. Although considerable volumes of sediment are involved in root throw events (resulting in notable values of sediment disturbance), the associated transport distances are relatively small (5 cm to 150 cm), leading to relatively low transport rates. Given the limited time window of our field study, the millennial-scale transport estimates derived in our model, which take into consideration the temporal dynamics of sediment transport, may be preferred. In general, vegetation properties are treated very simply in many geomorphic studies. The complex effects of vegetation on sediment transport are often encapsulated in various parameters within equations, rather than being explicitly considered. However, within our model, the details of tree population dynamics (recruitment and mortality) play a very direct role in determining the timing and pulsing of root Copyright © 2009 John Wiley & Sons, Ltd. throw sediment transport. In particular, two notable pulses of transport due to root throw are evident in our results: (i) if trees are large enough to have reached a critical dbh at time of fire, then a pulse of sediment occurs in post-fire years, which decreases exponentially with time since fire; and (ii) once new recruits have reached a critical dbh and with competition mortality (thinning), then a second pulse of root throw begins at about 50 to 60 years after the previous fire. If the period of time since the previous fire is not sufficient for enough standing trees to have reached a critical dbh, then trees may topple but they will not upheave notable amounts of sediment. It is only when the time since previous fire exceeds the time needed for critical dbh to be reached that notable sediment disturbance and transport begins to occur. After about a century the number of trees does not decrease very much and dbh continues to grow, increasing root plate volumes for the trees, which eventually leads to increased sediment disturbance and transport. Transport rates for our millennial-scale model run are now compared to published results in the literature. The model of Gabet et al. (2003) estimates an annual transport rate due to root throw of 1·6 × 10–3 m3 m–1 a–1 for a 20° slope. Roering et al. (2002) estimated millennial-scale transport rates for biogenic processes (e.g. root growth, root throw) based on vertical profiles of tephra concentration and topographic derivatives. The resulting K value (i.e. the diffusion coefficient, which represents the transport rate at unit gradient or 45°) is 1·2 × 10–2 m3 m–1 a–1. Since the form of this transport equation is linear, then the transport rate for a slope gradient of 20° (the slope gradient used in our model runs) can be inferred and has a value of 4·4 × 10–3 m3 m–1 a–1. Walther et al. (submitted for publication) estimated the diffusion coefficient for soil movement by root growth, bioturbation and tree throw based on the distribution of tephra grains in soils found in southeast Washington State. They obtained a diffusivity value of 4·8 × 10–3 m3 m–1 a–1, which translates into a transport rate of 1·7 × 10–3 m3 m–1 a–1 for a slope gradient of 20°. Results for our millennialscale model run provide a mean annual transport rate of 1·2 × 10–3 m3 m–1 a–1, which compares favorably with the longer term values obtained for the above-mentioned studies (all values have the same order of magnitude). For further comparison, Martin and Church (1997) reported an average linear diffusion coefficient based on a large number of soil creep rates reported in the literature of 2 × 10–4 m3 m–1 a–1 (7·3 × 10–5 m3 m–1 a–1 for a 20° slope), and reported a diffusion coefficient for debris slides of 1 × 10–1 m3 m–1 a–1 (3·6 × 10–2 m3 m–1 a–1 for a 20° slope); our transport rates for root throw are between these two values. This study provides an example of how direct the links may be between ecology and geomorphology. In our field and modeling studies, tree age and size as determined by the forest population dynamics was shown to be a key factor affecting the timing of root throw and the amount of sediment associated with these events. It is suggested that future geomorphological studies attempt to strike a balance between incorporating vegetation in a more realistic manner than has often been the case in past geomorphic studies, while at the same time keeping our understanding of the joint processes tractable. Acknowledgements—We acknowledge the financial support of the Biogeoscience Institute and NSERC Discovery Grants to EAJ and YEM. We also thank Kootenay National Park for the support provided for the field component of this project. Numerous field assistants provided excellent assistance in the field and contributed to the success of this project. The manuscript benefited from the perceptive comments of two anonymous reviewers. Earth Surf. Process. Landforms 34, 1255–1269 (2009) DOI: 10.1002/esp SEDIMENT TRANSPORT DUE TO TREE ROOT THROW References Beatty SW, Stone EL. 1986. The variety of soil microsites created by tree falls. Canadian Journal of Forest Research 16: 539–548. 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