Geology topics

Dan Binkley

Publications and source records attributed to Dan Binkley.

15 recordsLinked to original sources

USGS2016

Ecosystem development on terraces along the Kugururok River, northwest Alaska

Riverside terraces along the Kugururok River in the Noatak National Preserve provided an opportunity to study primary succession, considering general trends that apply across all terraces, and unique events that influence individual terraces. The 30-year-old willow/poplar (Salix spp., Populus balsamifera L.) terrace had no trees taller than 1.5 m; the abundant spruce trees were not tall enough to emerge from the canopy height of the willows and poplars, and moose (Alces alces [Clinton]) browsing limited the canopy height of these plants. The 75-year-old poplar/spruce (Picea glauca [Moench] Voss) terrace had a high density of poplars (> 1000/ha) and low density of spruce (125/ha); heavy browsing by moose reduced the density of poplar by about one-half. The removal of the poplar by moose in this stand resulted in sustained increases in growth of individual spruce trees. The 100-year-old younger spruce/poplar terrace had about twice as many spruce trees (1250/ha) as poplar trees (500/ha), and the spruce trees were larger on average than the poplar trees. In the 220+ year-old older spruce/poplar type, only a few poplars remained (about 25/ha), and the number of spruce trees (600/ha) was only half that of the younger stage, either from lower initial spruce density on this terrace, or increased mortality of spruce. The 240+ year-old spruce type was a second-generation forest, characterized by a high density (1950/ha) of small spruce trees, some of which were tilted, indicating discontinuous permafrost. Plant litterfall mass showed no strong trend with terrace age, although N content of litterfall appeared to decline by about 1/3 in the spruce-dominated stages. Fungal biomass increased with ecosystem age, whereas bacterial biomass and microfauna declined. We found no evidence of declining soil N supply in older stages, but fertilization experiments would be needed to determine if N limitation of productivity changed with ecosystem development. We conclude that the general successional trend of increased spruce dominance is robust for this location, but that unique events play important roles in determining tree densities and the timing of the shift in dominance from poplar to spruce. The arrival of moose in the 1970s accelerated dominance by spruce on young terraces.

Écoscience↗

USGS2016

Expansion of forest stands into tundra in the Noatak National Preserve, northwest Alaska

Temperatures across the northern regions of North America have been increasing for 150 years, and forests have responded to this increase. In the Noatak National Preserve in Alaska, white spruce (Picea glauca [Moench] Voss) forests reach their northern limit, occurring primarily on well-drained sites and as gallery forests along streams. Rolling plateaus of tundra separate the white spruce forests into disjunct stands. We examined patterns of tree age, tree growth, and tree encroachment into tundra ecosystems in six stands along the Agashashok River. Warming over the past 150 years appears to have increased tree growth and resulted in forest expansion into adjacent tundra ecosystems. The forest/tundra ecotone shifted by about 80 to 100 m into the tundra in the past 200 years, as evidenced by declining maximum tree age with distance towards the tundra. The decadal-scale pattern of tree establishment at the farthest extent of trees into the tundra (the tundra-forest ecotone) correlated with the detrended growth index for trees within the forests; climate conditions that led to higher tree growth appeared to foster tree establishment in the tundra. This recent forest expansion has occurred across topographic boundaries, from well-drained soils on slopes onto poorly drained, flatter areas of tundra. Further expansion of the forests may be limited by more severe wind exposure and poor drainage that make the majority of tundra less suitable for trees.

Écoscience↗

USGS2016

Alder ( Alnus crispa ) effects on soils in ecosystems of the Agashashok River valley, northwest Alaska

At the northern limit of the boreal forest biome, alder ( Alnus crispa [Ait.] Pursh) shrubs occur in a variety of ecosystems. We assessed the effects of individual alder shrubs on soil properties and understory plant tissue nitrogen in floodplain terraces, valley slopes and tussock tundra ridges. The three ecosystems differed with respect to soil properties and abiotic conditions and supported distinct plant communities. Alder increased resin-exchangeable soil N and NO 3 production significantly in each ecosystem. The greatest difference between alder canopy and surrounding soil NO 3 measured both under field and laboratory conditions occurred in floodplain sites. The shrub effect on soil pH and soil organic matter was greatest on tundra ridges. Alder shrubs also influenced the nitrogen nutrition of plants growing beneath their canopies. Plants growing below alder canopies had higher foliar nitrogen concentration and natural abundance 15 N composition and lower carbon to nitrogen ratio than open-grown plants. Similar to soil N availability, understory plant leaf chemistry responded more to alder on floodplains than on slope or tundra ecosystems. This pattern suggests that understory plants rely more heavily on alder-fixed-N in this resource-poor ecosystem.

Alaska↗

USGS2015

Soil nitrogen accretion along a floodplain terrace chronosequence in northwest Alaska: Influence of the nitrogen-fixing shrub Shepherdia canadensis

Nitrogen enters terrestrial ecosystems through multiple pathways during primary succession. We measured accumulation of total soil nitrogen and changes in inorganic nitrogen (N) pools across a 300-y sequence of river terraces in northwest Alaska and assessed the contribution of the nitrogen-fixing shrub Shepherdia canadensis . Our work compared 5 stages of floodplain succession, progressing from a sparsely vegetated silt cap to dense shrubby vegetation, balsam poplar-dominated ( Populus balsamifera ) and white spruce-dominated ( Picea glauca ) mixed forests, and old-growth white spruce forest. Total soil N (0–30 cm depth) increased throughout the age sequence, initially by 2.4 g N·m −2 ·y −1 during the first 120 y of terrace development, then by 1.6 g N·m −2 ·y −1 during the subsequent 2 centuries. Labile soil N, measured by anaerobic incubation, increased most rapidly during the first 85 y of terrace formation, then remained relatively constant during further terrace development. On recently formed terraces, Shepherdia shrubs enriched soil N pools several-fold compared to soil beneath Salix spp. shrubs or intercanopy sites. Total and labile soil N accretion was proportional to Shepherdia cover during the first century of terrace development, and mineral soil δ 15 N content indicated that newly formed river terraces receive substantial N through N-fixation. About half the 600 g total N·m −2 accumulated across the river terrace chronosequence occurred during the 120 y when S. canadensis was dominant. Sediment deposited by periodic flooding continued to add N to terrace soils after the decline in Shepherdia abundance and may have contributed 25% of the total N found in the floodplain terrace soils.

Alaska↗

USGS2014

Can nitrogen fertilization aid restoration of mature tree productivity in degraded dryland riverine ecosystems?

Restoration of riparian forest productivity lost as a consequence of flow regulation is a common management goal in dryland riverine ecosystems. In the northern hemisphere, dryland river floodplain trees often include one or another species of Populus, which are fast-growing, nutrient-demanding trees. Because the trees are phreatophytic in drylands, and have water needs met in whole or in part by a shallow water table, their productivity may be limited by nitrogen (N) availability, which commonly limits primary productivity in mesic environments. We added 20 g N m −2 in a 2-m radius around the base of mature Populus fremontii along each of a regulated and free-flowing river in semiarid northwest Colorado, USA (total n = 42) in order to test whether growth is constrained by low soil N. Twelve years after fertilization, we collected increment cores from these and matched unfertilized trees and compared radial growth ratios (growth in the 3-year post-fertilization period/growth in the 3-year pre-fertilization period) in paired t tests. We expected a higher mean ratio in the fertilized trees. No effect from fertilization was detected, nor was a trend evident on either river. An alternative test using analysis of covariance (ANCOVA) produced a similar result. Our results underscore the need for additional assessment of which and to what extent factors other than water control dryland riverine productivity. Positive confirmation of adequate soil nutrients at these and other dryland riparian sites would bolster the argument that flow management is necessary and sufficient to maximize productivity and enhance resilience in affected desert riverine forests.

Colorado↗

USGS2013

Unsupported inferences of high-severity fire in historical dry forests of the western United States: Response to Williams and Baker

Reconstructions of dry western US forests in the late 19th century in Arizona, Colorado and Oregon based on General Land Office records were used by Williams & Baker (2012; Global Ecology and Biogeography, 21, 1042–1052; hereafter W&B) to infer past fire regimes with substantial moderate and high-severity burning. The authors concluded that present-day large, high-severity fires are not distinguishable from historical patterns. We present evidence of important errors in their study. First, the use of tree size distributions to reconstruct past fire severity and extent is not supported by empirical age–size relationships nor by studies that directly quantified disturbance history in these forests. Second, the fire severity classification of W&B is qualitatively different from most modern classification schemes, and is based on different types of data, leading to an inappropriate comparison. Third, we note that while W&B asserted ‘surprising’ heterogeneity in their reconstructions of stand density and species composition, their data are not substantially different from many previous studies which reached very different conclusions about subsequent forest and fire behaviour changes. Contrary to the conclusions of W&B, the preponderance of scientific evidence indicates that conservation of dry forest ecosystems in the western United States and their ecological, social and economic value is not consistent with a present-day disturbance regime of large, high-severity fires, especially under changing climate

Arizona, Colorado, Oregon↗

USGS2008

Aspen Ecology in Rocky Mountain National Park: Age Distribution, Genetics, and the Effects of Elk Herbivory

Lack of recruitment and canopy replacement of aspen (Populus tremuloides) stands that grow on the edges of grasslands on the low-elevation elk (Cervus elaphus) winter range of Rocky Mountain National Park (RMNP) in Colorado have been a cause of concern for more than 70 years. We used a combination of traditional dendrochronology and genetic techniques as well as measuring the characteristics of regenerating and nonregenerating stands on the elk winter range to determine when and under what conditions and estimated elk densities these stands established and through what mechanisms they may regenerate. The period from 1975 to 1995 at low elevation on the east side had 80-95 percent fewer aspen stems than would be expected based on the trend from 1855 through 1965. The age structure of aspen in the park indicates that the interacting effects of fires, elk population changes, and livestock grazing had more-or-less consistent effects on aspen from 1855 to 1965. The lack of a significant change in aspen numbers in recent decades in the higher elevation and west side parts of the park supports the idea that the extensive effects of elk browsing have been more important in reducing aspen numbers than other factors. The genetic variation of aspen populations in RMNP is high at the molecular level. We expected to find that most patches of aspen in the park were composed of a single clone of genetically identical trees, but in fact just 7 percent of measured aspen patches consisted of a single clone. A large frequency of polyploid (triploid and tetraploid) genotypes were found on the low elevation, east-side elk winter range. Nonregenerating aspen stands on the winter range had greater annual offtake, shorter saplings, and lower density of mid-height (1.5-2.5 m) saplings than regenerating stands. Overwinter elk browsing, however, did not appear to inhibit the leader length of aspen saplings. The winter range aspen stands of RMNP appear to be highly resilient in the face of very intense herbivory by elk and harsh environmental conditions. Conservation efforts through fencing protection and decreased elk browsing pressure are already being planned as part of the park's new elk management plan. If these efforts are undertaken, conditions that encourage stem recruitment to the tree canopy will likely result and the continued survival of these aspen stands will be enhanced.

Open-File Report↗

USGS2004

Effects of elk herbivory on vegetation and nitrogen processes

We used 35-year and 4-year ungulate exclosures to determine the effects of elk ( Cervus elaphus ) herbivory on above-ground and below-ground production and soil fertility on the elk winter range in Rocky Mountain National Park (RMNP), Colorado, USA. We used paired grazed and ungrazed plots to evaluate ungulate herbivory effects in short and tall willow ( Salix spp.), aspen ( Populus spp.), and upland grass/shrub vegetation associations. We measured nitrogen (N) fluxes (litter deposition, fecal and urinary deposition from elk, movements of N by elk, N mineralization, soil N availability, elk consumption rates) within the elk winter, above-ground and below-ground N pools (herbaceous, shrub and root biomass, %N in plants, roots, and soil), and N fluxes on and off the elk winter range (seasonal movement of N by elk). Nitrogen mineralization and soil nitrate (NO 3 ) pools were reduced in the short willow community ( P = 0.07 and 0.10, respectively; n = 4 sites) in grazed plots, but not in the upland grass/shrub community or tall willow sites ( P >0.10). Annual growth of willows was reduced by 98% in grazed plots, relative to 35-year exclosures, and 66% relative to 4-year exclosures. Thus, height, canopy size, and litter biomass of willows were reduced, and N yield of willows was 64% less in grazed plots. We evaluated movement of N by elk among 6 major vegetation associations and found that elk grazed more and bedded less in willow vegetation association compared to mixed conifer, mesic meadow, and grassland/shrub associations ( P = 0.014, 0.001, and 0.026, respectively), suggesting that elk herbivory and movement led to a net loss of N in the willow vegetation association. Elk spent less total time in willows than mesic meadow association, yet they consumed large amounts of willow plant biomass. We recommend management of elk numbers and elk herbivory that takes into consideration impacts to N process function, as negative effects from current levels of herbivory were observed in ≥1 of 3 vegetation associations studied.

Colorado↗

USGS2003

Structure, production and resource use in some old-growth spruce/fir forests in the front range of the Rocky Mountains, USA

Old-growth forests of Engelmann spruce (Picea engelmannii Parry ex. Engelm.) and subalpine fir (Abies lasiocarpa (Hook.) Nutt.) dominate much of the landscape of the Rocky Mountains. We characterized the structure, biomass and production of 18 old-growth (200-450-year-old) spruce/fir forests in Rocky Mountain National Park, Colorado, as well as the stand-level supply and use of light and nitrogen. Stands were chosen to span a broad range of elevation, aspect, and topography. Aboveground tree biomass in these old-growth forests averaged 253 Mg/ha (range 130-488 Mg/ha), with aboveground net primary production of 3700 kg ha-1 yr-1 (range from 2700 to 5200 kg ha-1 yr-1). Within stands, trees >35 cm in diameter accounted for 70% of aboveground biomass, but trees <35 cm contributed 70% of the production of woody biomass. Differences in slope and aspect among sites resulted in a range of incoming light from 58 to 74 TJ ha-1 yr-1, and tree canopies intercepted an average of 71% of incoming light (range 50-90%). Aboveground net primary production (ANPP) of trees did not relate to the supply of light or N, but ANPP correlated strongly with the amount of light and N used (r2 = 0.45-0.54, P < 0.01). Uptake of 1 kg of N was associated with about 260 kg of ANPP, and one TJ of intercepted shortwave radiation produced about 78 kg of ANPP. Across these old-growth stands, stands with greater biomass showed higher rates of both ANPP and resource use; variation in aboveground biomass was associated with 24% of the variation in N use (P = 0.04), 44% of the light use (P = 0.003), and 45% of the ANPP (P = 0.002). ?? 2002 Elsevier Science B.V. All rights reserved.

Forest Ecology and Management↗

USGS2003

Influence of elk grazing on soil properties in Rocky Mountain National Park

We used three 35-year exclosures to examine the effects of high elk populations on a variety of soil properties in three vegetation types: upland sagebrush, aspen, and meadow. Grazing and hoof action by elk significantly increased bulk density (from 0.87 kg/l ungrazed to 0.94 kg/l grazed), with greater effects on soils with fewer rocks. Grazing substantially reduced extractable calcium, magnesium, potassium and phosphorus in the sagebrush type, but not in the aspen or meadow types. The only grazing effect on pH came in aspen types, where grazing prevented aspen establishment, and kept soil pH about 0.7 units higher than under aspen inside the exclosures. Grazing had no overall effect on total soil C and N across all exclosures and vegetation types. The availability of soil nitrogen, indexed by in-field resin bags and net mineralization in soil cores, showed little overall effect of grazing. Limited data on soil leaching indicated a possibility of strong increases in nitrate leaching with grazing for an aspen vegetation type at one exclosure. Although we found little effect of grazing on soil N supply, we note that N fertilization doubled the production of grasses and shrubs; if grazing eventually led to changes in soil N supply, species composition and growth would likely change. ?? 2003 Elsevier B.V. All rights reserved.

Forest Ecology and Management↗

USGS2003

Aspen structure and variability in Rocky Mountain National Park, Colorado, USA

Elk, fire and climate have influenced aspen populations in the Rocky Mountains, but mostly subjective studies have characterized these factors. A broad-scale perspective may shed new light on the status of aspen in the region. We collected field measurements of aspen (Populus tremuloides Michx.) patches encountered within 36 randomly located belt transects in 340 km2 of Rocky Mountain National Park, Colorado, to quantify the aspen population. Aspen covered 5.6% of the area in the transects, much more than expected based on previously collected remotely sensed data. The distribution and structure of aspen patches were highly heterogeneous throughout the study area. Of the 123 aspen patches encountered in the 238 ha surveyed, all but one showed signs of elk browsing or had conifer species mixed with the aspen stems. No significant difference occurred in aspen basal area, density, regeneration, browsing of regeneration and patch size, between areas of concentrated elk use (elk winter range) and areas of dispersed elk use (elk summer range). Two-thirds of the aspen patches were mixed with conifer species. We concluded that the population of aspen in our study area is highly variable in structure and that, at a landscape-scale, evidence of elk browsing is widespread but evidence of aspen decline is not.

Landscape Ecology↗

USGS2003

Flood flows, leaf breakdown, and plant-available nitrogen on a dryland river floodplain

We tested the hypothesis that decomposition in flood-inundated patches of riparian tree leaf litter results in higher plant-available nitrogen in underlying, nutrient-poor alluvium. We used leafpacks ( n = 56) containing cottonwood ( Populus deltoides ssp. wislizenii ) leaf litter to mimic natural accumulations of leaves in an experiment conducted on the Yampa River floodplain in semi-arid northwestern Colorado, USA. One-half of the leafpacks were set on the sandy alluvial surface, and one-half were buried 5 cm below the surface. The presence of NO 3 − and NH 4 + presumed to result from a leafpack's submergence during the predictable spring flood pulse was assessed using an ion-exchange resin bag (IER) placed beneath each leafpack and at control locations. Leafpacks and IERs were collected one week after flood peak (71 days total exposure) at half the stations; the remainder were collected three weeks later (93 days exposure). A multi-peaked spring flood with above-average maximum discharge inundated leafpacks for total time periods ranging from 133 to 577 hours. Litter lost from 43 to 68 percent of its initial organic matter (OM) content. Organic matter loss increased with total time inundated and total time of exposure on the floodplain. Burial retarded OM loss if the total time inundated was relatively long, and substrate texture (sand vs. silt) affected OM loss in a complex manner through interactions with total time inundated and total time of exposure. No pulse of N attributable to leaf breakdown was detected in the IERs, and leafpack litter showed no net change in the mass of nitrogen present. Patterns of leafpack and IER nitrogen levels suggested that litter removed N from floodwater and thereby reduced N availability in underlying sediment. Immobilization of floodwater-N by litter and N mineralization outside the flood period may be important components of N flux in semi-arid and arid floodplain environments.

Colorado↗

USGS2002

Sustainability of vegetation communities grazed by elk in Rocky Mountain National Park

Current management of the worlds' grazing lands in either based on changes in plant species composition or on other management evaluation programs that emphasize changes in net aboveground production. Management is based solely on changes in aboveground production has been criticized as too limited in view, because it ignores root production, nitrogen pools, nutrient processes, and the long-term sustainability of the ecosystem. The purpose of this study was to compare the effects of elk ( Cervus elaphus ) grazing on aboveground production, internal nitrogen (N) fluxes, N pools and inputs, and elk nutrient transfers across the landscape in different vegetation types in Rocky Mountain National Park (RMNP), Colorado. Nitrogen processes and possibly N pools were significantly reduced in the willow community, but not in the upland grass/shrub community. Nitrogen mineralization rates were lower in grazed versus ungrazed short willow sites (P = 0.07; n = 4 sites), as were nitrate (NO3) pools (P = 0.10), but not in tall willow sites (P > 1.10 n = 4 sites) after 4 years. There was about half the annual N inputs to the soil surface in grazed willow sites (5.79g N/m 2 /yr = annual herbaceous biomass a?? offtake + litterfall + elk urine and feces) compared to ungrazed sites (9.66 g N/m 2 /yr = annual herbaceous biomass + litterfall), suggesting elk herbivory and movement led to a net loss of N in the willow vegetation type. Elk substantially reduced the annual growth of willows (Salix spp.) by 98% after 35 years and 66% after 4 years of treatment. Thus, height and canopy and N yield of willows were reduced as well as willow litter biomass (65 g/m2/yr in ungrazed versus 33 g/m2/yr in grazed), and N yield of willows was 64% less in grazed plots. Elk grazing had no significant effect on other soil N pools (NH4) or litter decomposition rates in either of the two willow types, nor on any nitrogen process rates or pools in the upland grass/shrub type (P > 0.10). Nitrogen concentrations in plant tissue were not influenced for the most part by elk grazing (increased N concentration found in only 4 of 13 species). Elk apparently also transported N away from aspen at an even higher rate (N inputs were 1.65 g N/m2/yr in grazed sites vs. 3.79 g N/m 2 /yr ungrazed). CENTURY modeling supported these observations. This soil N model predicted almost no change (0.5$ - 2% less) in N or carbon (C) pools in 50 years in the upland grass/shrub vegetation type, but greater losses in the willow type. If elk population levels were increased to carrying capacity in the ecosystem (about 25% more consumption) projected losses after 50 years were greater, but still <4% for soil C and 1% for spoil N in the upland type, and 6% for soil C and 2% for soil N in the willow type. Total shrub C was projected to decline 10% in 50 years in the willow type at current elk densities. We recommend conservative management of elk numbers and grazing until additional years of measures are gathered on the lower mineralization rates in the short willow type and until the projected declines in N pools can be verified.

Report↗

USGS1999

Exotic plant species invade hot spots of native plant diversity

Some theories and experimental studies suggest that areas of low plant species richness may be invaded more easily than areas of high plant species richness. We gathered nested-scale vegetation data on plant species richness, foliar cover, and frequency from 200 1-m 2 subplots (20 1000-m 2 modified-Whittaker plots) in the Colorado Rockies (USA), and 160 1-m 2 subplots (16 1000-m 2 plots) in the Central Grasslands in Colorado, Wyoming, South Dakota, and Minnesota (USA) to test the generality of this paradigm. At the 1-m 2 scale, the paradigm was supported in four prairie types in the Central Grasslands, where exotic species richness declined with increasing plant species richness and cover. At the 1-m 2 scale, five forest and meadow vegetation types in the Colorado Rockies contradicted the paradigm; exotic species richness increased with native-plant species richness and foliar cover. At the 1000-m 2 plot scale (among vegetation types), 83% of the variance in exotic species richness in the Central Grasslands was explained by the total percentage of nitrogen in the soil and the cover of native plant species. In the Colorado Rockies, 69% of the variance in exotic species richness in 1000-m 2 plots was explained by the number of native plant species and the total percentage of soil carbon. At landscape and biome scales, exotic species primarily invaded areas of high species richness in the four Central Grasslands sites and in the five Colorado Rockies vegetation types. For the nine vegetation types in both biomes, exotic species cover was positively correlated with mean foliar cover, mean soil percentage N, and the total number of exotic species. These patterns of invasibility depend on spatial scale, biome and vegetation type, spatial autocorrelation effects, availability of resources, and species-specific responses to grazing and other disturbances. We conclude that: (1) sites high in herbaceous foliar cover and soil fertility, and hot spots of plant diversity (and biodiversity), are invasible in many landscapes; and (2) this pattern may be more closely related to the degree resources are available in native plant communities, independent of species richness. Exotic plant invasions in rare habitats and distinctive plant communities pose a significant challenge to land managers and conservation biologists.

Ecological Monographs↗

USGS1998

Species-environment relationships and vegetation patterns: Effects of spatial scale and tree life-stage

Do relationships between species and environmental gradients strengthen or weaken with tree life-stage (i.e., small seedlings, large seedlings, saplings, and mature trees)? Strengthened relationships may lead to distinct forest type boundaries, or weakening connections could lead to gradual ecotones and heterogeneous forest landscapes. We quantified the changes in forest dominance (basal area of tree species by life-stage) and environmental factors (elevation, slope, aspect, intercepted photosynthetically active radiation (PAR), summer soil moisture, and soil depth and texture) across 14 forest ecotones (n = 584, 10 m x 10 m plots) in Rocky Mountain National Park, Colorado, U.S.A. Local, ecotone-specific species-environment relationships, based on multiple regression techniques, generally strengthened from the small seedling stage (multiple R2 ranged from 0.00 to 0.26) to the tree stage (multiple R2 ranged from 0.20 to 0.61). At the landscape scale, combined canonical correspondence analysis (CCA) among species and for all tree life-stages suggested that the seedlings of most species became established in lower-elevation, drier sites than where mature trees of the same species dominated. However, conflicting evidence showed that species-environment relationships may weaken with tree life-stage. Seedlings were only found in a subset of plots (habitats) occupied by mature trees of the same species. At the landscape scale, CCA results showed that species-environment relationships weakened somewhat from the small seedling stage (86.4% of the variance explained by the first two axes) to the tree stage (76.6% of variance explained). The basal area of tree species co-occurring with Pinus contorta Doug. ex. Loud declined more gradually than P. contorta basal area declined across ecotones, resulting in less-distinct forest type boundaries. We conclude that broad, gradual ecotones and heterogeneous forest landscapes are created and maintained by: (1) sporadic establishment of seedlings in sub-optimal habitats; (2) survivorship of saplings and mature trees in a wider range of environmental conditions than seedlings presently endure; and (3) the longevity of trees and persistence of tree species in a broad range of soils, climates, and disturbance regimes.

Plant Ecology↗