The population biology of Bromus tectorum in forests: effect of disturbance,grazing, and litter on seedling establishment and reproduction

Summary The effect of tree canopy, understory, herbivores, and litter depth on seedling establishment, survival, and reproduction of the alien grass, Bromus tectorum (cheatgrass), was examined in a series of experiments in four forest habitat types in western North America. Higher recruitment, survival, and reproduction on clearcuts, which would be expected if the overstory alone is limiting the distribution of cheatgrass in forests, were not observed. Removing the understory in an otherwise undisturbed Pinus ponderosa forest did, however, increase the emergence of B. tectorum, but plants in these experimentally-created openings were more vulnerable to grazing by small mammals. In contrast, removing the sparse understory in an Abies forest neither enhanced recruitment nor increased the incidence of grazing of B. tectorum seedlings. Regardless of the forest habitat, most grazed plants died before maturity; even fewer grazed plants produced seeds. Litter depth influenced both recruitment and biomass production: both the rate of germination and the size of resultant seedlings were lower on thick litter (6 cm) compared to results on thin litter (1.5 cm). In the more open Pinus ponderosa and Pseudotsuga menziesii forests, cheatgrass colonization may often occur in openings in the understory alone. Colonization in the more shady A. grandis and Thuja plicata forests is unlikely, however, unless the opening extends through both the understory and the overstory. As a result, cheatgrass is unlikely to increase in any of these forests unless the scale and incidence of disturbance increases substantially.     

Breeding Birds in Riparian and Upland Dry Forests of the Cascade Range

ABSTRACT We quantified breeding bird abundance, diversity, and indicator species in riparian and upland dry forests along 6 third- to fourth-order streams on the east slope of the Cascade Range, Washington, USA. Upland dry forest on southerly aspects was dominated by open ponderosa pine (Pinus ponderosa) and dry Douglas-fir (Pseudotsuga menziesii) plant associations. Upland mesic forest on northerly aspects was dominated by closed-canopy Douglas-fir or dry grand fir (Abies grandis) plant associations. Riparian overstory vegetation was dominated by black cottonwood (Populus trichocarpa) plant associations with a prominent hardwood tree and shrub component. We quantified bird assemblages, diversity, and abundance from parallel point transects on riparian and adjacent dry and mesic upslope forests. We detected 80 bird species from >12,000 point-transect observations during 1998–1999. Eighteen species accounted for 75% of all detections. Species richness and evenness were similar in all 3 forest types, with approximately 35 species and high evenness (0.85) in each forest type. Bird species assemblages differed among dry, mesic, and riparian forest types, with the greatest differences between riparian and both dry and mesic upland forests. Riparian forest had the greatest number (9) of strong characteristic, or indictor, species among the 3 forest types. Upland mesic forest was characterized by 7 indicator species. Upland dry forest had 4 indicator species. Our results indicate that current standards and guidelines for riparian buffers zones would allow for avian refuge and corridor functions along these streams. Forest managers could use our indicator species to predict and monitor shifts in upland forest species composition from thinning and prescribed burning practices that are used to reduce fuels in uplands and to reduce continuity of fire effects between riparian and upland zones.     

Understory vegetation indicates historic fire regimes in ponderosa pine‐dominated ecosystems in the Colorado Front Range

Question: Can current understory vegetation composition across an elevation gradient of Pinus ponderosa‐dominated forests be used to identify areas that, prior to 20th century fire suppression, were characterized by different fire frequencies and severities (i.e., historic fire regimes)? Location: P. ponderosa‐dominated forests in the montane zone of the northern Colorado Front Range, Boulder and Larimer Counties, Colorado, USA. Methods: Understory species composition and stand characteristics were sampled at 43 sites with previously determined fire histories. Indicator species analyses and indirect ordination were used to determine: (1) if stands within a particular historic fire regime had similar understory compositions, and (2) if understory vegetation was associated with the same environmental gradients that influence fire regime. Classification and regression tree analysis was used to ascertain which species could predict fire regimes. Results: Indicator species analysis identified 34 understory species as significant indicators of three distinct historic fire regimes along an elevation gradient from low‐ to high‐elevation P. ponderosa forests. A predictive model derived from a classification tree identified five species as reliable predictors of fire regime. Conclusions: P. ponderosa‐dominated forests shaped by three distinct historic fire regimes have significantly different floristic composition, and current understory compositions can be used as reliable indicators of historical differences in past fire frequency and severity. The feasibility demonstrated in the current study using current understory vegetation properties to detect different historic fire regimes, should be examined in other fire‐prone forest ecosystems.     

Inter-hemispheric comparison of fire history: The Colorado Front Range,U.S.A., and the Northern Patagonian Andes,Argentina

Fire history was compared between the Colorado Front Range (U.S.A.) and northern Patagonia (Argentina) by dating fire-scars on 525 Pinus ponderosa and 418 Austrocedrus chilensis, respectively, and determining fire weather on the basis of instrumental and tree-ring proxy records of climatic variation. Years of above average moisture availability preceding fire years, rather than drought alone, is conducive to years of widespread fire in the Colorado Front Range and the northern Patagonian study areas. Above-average precipitation promotes fire by enhancing the growth of herbaceous plants which increases the quantity of fine fuels during the fire season a few years later. The short-term variability in moisture availability that is conducive to widespread burning is strongly related to El Niño Southern Oscillation (ENSO) activity. The warm (El Niño) phase of ENSO is associated with greater moisture availability during the spring in both regions which leads to peaks in fire occurrence several years after El Niño events. The warmer and drier springs associated with la Niña events exacerbate the drying of fuels so that fire years commonly coincide with La Niña events. In both regions, there was a dramatic decline in fire occurrence after the early 1900s due to a decline in intentionally set fires by Native Americans and European settlers, fuel reduction by livestock grazing, and increasingly effective organized fire suppression activities after the 1920s. In both regions there was a marked increase in fire frequency during the mid-and late-19th centuries which coincides with increased ignitions by Native Americans and/or European settlers. However, year-to-year variability in ring widths of Pinus ponderosa and Austrocedrus chilensis also increased from relatively low values in the late 1700s and early 1800s to peaks in the 1850s and 1860s. This implies frequent alternation of years of above and below average moisture availability during the mid-19th century when the frequencies of major fire years rise. The high correlation of tree-growth variability betweem the two regions implies a strong inter-hemispheric variation in climatic variability at a centennial time scale which closely parallels a variety of proxy records of ENSO activity. Based on the relationship of fire and ENSO events documented in the current study, this long-term trend in ENSO activity probably contributed to the mid- and late-19th century increase in fire spread in both regions. These similar trends in fire occurrence have contributed to similar patterns of forest structures, forest health, and current hazard of catastrophic wildfire in the Colorado Front Range and northern Patagonia.     

Drought responses of conifers in ecotone forests of northern Arizona: tree ring growth and leaf δ<Superscript>13</Superscript>C

We sought to understand differences in tree response to meteorological drought among species and soil types at two ecotone forests in northern Arizona, the pinyon-juniper woodland/ponderosa pine ecotone, and the higher elevation, wetter, ponderosa pine/mixed conifer ecotone. We used two approaches that provide different information about drought response: the ratio of standardized radial growth in wet years to dry years (W:D) for the period between years 1950 and 2000 as a measure of growth response to drought, and ¹³C in leaves formed in non-drought (2001) and drought (2002) years as a measure of change in water use efficiency (WUE) in response to drought. W:D and leaf ¹³C response to drought for Pinus edulis and P. ponderosa did not differ for trees growing on coarse-texture soils derived from cinders compared with finer textured soils derived from flow basalts or sedimentary rocks. P. ponderosa growing near its low elevation range limit at the pinyon-juniper woodland/ponderosa pine ecotone had a greater growth response to drought (higher W:D) and a larger increase in WUE in response to drought than co-occurring P. edulis growing near its high elevation range limit. P. flexilis and Pseudotsuga menziesii growing near their low elevation range limit at the ponderosa pine/mixed conifer ecotone had a larger growth response to drought than co-occurring P. ponderosa growing near its high elevation range limit. Increases in WUE in response to drought were similar for all species at the ponderosa pine/mixed conifer ecotone. Low elevation populations of P. ponderosa had greater growth response to drought than high-elevation populations, whereas populations had a similar increase in WUE in response to drought. Our findings of different responses to drought among co-occurring tree species and between low- and high-elevation populations are interpreted in the context of drought impacts on montane coniferous forests of the southwestern USA.     

Changes in riparian forest composition along a sedimentation rate gradient

Riparian forests are highly valued for maintaining water quality through the retention of sediments and nutrients. They also provide some of the most diverse and species-rich habitats in the world. What is largely unknown, however, is how sediment deposition affects plant community composition in these forests. The objective of this study was to examine changes in plant community composition across a gradient of increasing rates of sedimentation in riparian forests in the southeastern Coastal Plain, USA. Seventeen plots were established within riparian forests receiving between 0 and 5.5 cm year⁻¹ of sediment deposits. Species density and biomass estimates were collected annually from 2002 to 2006 for overstory and mid-story plant species within each plot. Percent cover and nested frequency of understory plant species were determined annually during 2004–2006. Measures of community composition in the understory, mid-story, and overstory layers of forests were compared to changes in environmental factors associated with increased sedimentation. In the understory, annual, exotic, and upland species had higher importance values in plots receiving high sediment deposition. The densities of shade-intolerant and N-fixing species in the mid-story also increased with increasing sedimentation rates. Increased overstory mortality was associated with high sedimentation rates, though increases in understory light levels in these gaps were not the main driver of understory species changes. Edaphic factors, such as soil texture, moisture, and temperature, were significantly correlated to species composition in all three forest layers, suggesting that changes in soil physical structure due to sedimentation may drive community-level changes in these forests.     

Overstory-Understory Relationships along Forest Type and Environmental Gradients in the Spring Mountains of Southern Nevada, USA

Isolated forested mountains in deserts have numerous ecological and societal values, but land-management practices (e.g., fire-regime alteration) and climate change can affect forest composition. We analyzed tree overstory-understory relationships on 123 sites in the Spring Mountains within the Mojave Desert near Las Vegas, Nevada, USA to assess three hypotheses. We hypothesized that: the tree species comprising understories are less tolerant of fire than species in overstories, reflecting land-management practices of fire exclusion; mid-elevation forests have the lowest overstory:understory similarity because this zone could have maximum species mixing; and overstory:understory similarity is correlated with environmental gradients (consisting of 14 topographic and soil variables). We found that Pinus monophylla comprised greater relative canopy cover in understories of juniper (32% relative cover) and pinyon-juniper (78%) forests than it did in overstories of these forests (0% and 53%). Similarly, fire-intolerant Abies concolor had 6-fold greater understory than overstory cover in forests with overstories dominated by the fire-tolerant Pinus ponderosa. Overstory:understory S?rensen similarity averaged 43%?77% among six forest types, and there was little support for the supposition that similarity was lowest in mid-elevation forests. Distributions of individual overstory and understory species more closely corresponded with environmental gradients than did overstory:understory similarity. Results suggest that there is high potential for change in at least two of the six dominant forest types of the Spring Mountains. The direction of change (species of moist, higher elevation sites establishing in understories of drier forests) is the opposite of what would be expected for forest adaptation to the warmer, drier, more fire-prone conditions projected for the next century in the southwestern USA.     

Limitations to seedling establishment in a mesic Hawaiian forest

While invasive species may be visible indicators of plant community degradation, they may not constitute the only, or even the primary, limitation to stand regeneration. We used seed-augmentation and grass-removal experiments under different canopy conditions to assess the relative importance of dispersal limitation, resource availability, and competition on seedling establishment in the understory shrubs Sophora chrysophilla, Dodonea viscosa, and Pipturus albidus in a montane mesic forest in Hawaii. The study location was an Acacia koa-Metrosideros polymorpha forest at 1000–1500 m elevation on the leeward side of Hawaii Island; it is a closed-canopy forest historically subject to logging and grazing by cattle and sheep and currently dominated by the exotic grass, Ehrharta stipoides, in the herb layer. Seedling establishment after 1 and 2 years was strongly dispersal limited in Sophora and Dodonea, but not in Acacia, a non-augmented species in which abundant seedlings established, nor in Pipterus, in which only one seedling established in 2 years. Grass cover reduced seedling establishment in Acacia, Sophora, and Dodonea and, for the latter two species, seedling establishment was substantially greater in the warmer, more moist forest at the lowest elevation. Light, moisture, and resin-captured N and P were more strongly affected by elevation and canopy composition than by grass cover, but in most cases seedling establishment was not positively correlated with resource availability. Limitations to the establishment of woody seedlings in this forest-grassland mixture vary among species; however, both dispersal limitation and competition from a shade-tolerant grass are important deterrents to regeneration in these forests.     

Riverscape-level patterns of riparian plant diversity along a successional gradient, Willamette river, Oregon

Riparian forest communities dominated by Populus balsamifera ssp. trichocarpa L. (Torr. and Gray ex Hook.) Brayshaw are important contributors to biodiversity in terrestrial and aquatic ecosystems of the Western United States. Species composition along a successional gradient from stand initiation to late-succession of P. balsamifera-dominated riparian forests was investigated along 145 km of the Willamette River, Oregon. There were 151 total species encountered across 28 stands and a mean species richness of 33.3 species per stand. Young stands were dominated by P. balsamifera and Salix tree spp. and opportunistic herbaceous species. Understory trees, shrubs, and herbaceous species as well as late-successional tree species established 12–15 years after stand initiation. Fraxinus latifolia Benth. was the dominant late-successional tree species. Vertical structural diversity, P. balsamifera mean diameter at breast height, large tree biomass, and stand age were strongly correlated with understory species presence and abundance based on non-metric multidimensional scaling (NMS) ordination. There were no young stands on mid and high terraces and this was reflected in geomorphic position being strongly correlated with the stand age gradient. Abundance of Phalaris arundinacea L. an invasive grass species, was also significantly correlated with plant species composition and abundance. This study indicates that Willamette River riparian forests are diverse and therefore important to the biodiversity of the Willamette River valley and that their presence as a mosaic of communities of different successional stages may be threatened by human interventions, including influences exerted by introduced plant species.     

Late quaternary dynamics of pollen influx at mineral lake,Washington

Pollen influx analysis at Mineral Lake, Washington, indicates that immediately south of the Puget Lobe of the Fraser Glaciation, tundra was a characteristic vegetation until 16,300 years ago. Invasion ofPinus contorta began 17,500 years B.P., and boreal climax conifers (Abies, Picea andTsuga mertensiana), 16,300, but was temporarily interrupted by the Vashon advance (14,500–14,000 yr B.P.).Pseudotsuga menziesii began to grow in population 10,750 years ago, and woodland was established within a time span of 1,000 years. Modern lowland coniferous forests began to form 7,000 years ago. Logistic analysis of pollen abundance changes show that the intrinsic growth rate,r (yr⁻¹), of pioneer species (e.g. 0.024–0.026 inPteridium aquilinum) is higher than that of climax species (e.g. 0.003 inThuja plicata).P. menziesii, a subclimax species, shows an intermediater value (0.013) between these two ecologically different taxa. The absoluter value ofP. contorta (−0.011) is only slightly lower than that ofP. menziesii, although their replacement began almost simultaneously. Thus competition between these species was intense before the inflection point ofPinus curve 10,100 years ago. At this time, forest gaps became abundantly available forPseudotsuga, as indicated by a peak of the diagnostic factor (the reciprocal of the pollen influx).     

The influence of canopy cover on understorey development in forests of the western Cascade Range,Oregon, USA

Natural disturbances, especially fire and treefalls, influence tree canopy composition in the Pseudotsuga menziesii forests of the western Cascade Range, Oregon. The composition of tree, shrub, and herb assemblages in the understorey of stands with different canopy types, such as maturing Pseudotsuga, Tsuga heterophylla, or mixed species stands, also differs.Differences in both canopy type and the prevalence of canopy openings correlated with different degrees of understorey development in stands of similar ages. This suggests that understorey assemblages also reflect disturbance history. Before understorey assemblages can be used to relate community samples to community or habitat types, the extent to which their composition reflects long term influences of stand history vs. differences in site potential must be determined.     

Habitat Conditions Associated With Lynx Hunting Behavior During Winter in Northern Washington

Abstract Effectively managing habitat for threatened populations of Canada lynx (Lynx canadensis) requires knowledge of habitat conditions that provide for the ecological needs of lynx. We snow-tracked lynx to identify habitat conditions associated with hunting behavior and predation during winters of 2002–2003 and 2003–2004 in the northern Cascade Range in Washington state, USA. We recorded number and success of predation attempts, prey species killed, and trail sinuosity on 149 km of lynx trails. Lynx killed snowshoe hares (Lepus americanus), red squirrels (Tamiasciurus hudsonicus), and cricetids more than expected in Englemann spruce (Picea engelmannii) and subalpine fir (Abies lasiocarpa) forests, where snowshoe hare densities were highest. Lynx killed prey less than expected in Douglas-fir (Pseudotsuga menziesii) and ponderosa pine (Pinus ponderosa) forests and forest openings. We used the sinuosity of lynx trails as an index of quality of habitat hunted. Lynx trails that included predation attempts were more sinuous than trail segments without predation attempts. Lynx trails had greater sinuosity in forest stands with high hare densities dominated by Engelmann spruce and subalpine fir than in stands with low hare densities dominated by Douglas-fir and ponderosa pine or in forest openings. We encourage forest managers to maintain or create sufficient understory cover to support high densities of snowshoe hares as foraging habitat for lynx.     

Water flux and canopy conductance of natural versus planted forests in Patagonia, South America

In NW Patagonia, South America, natural shrublands and mixed forests of short Nothofagus antarctica (G. Forst.) Oerst. trees are currently being replaced by plantations with Pseudotsuga menziesii (Mirb) Franco. This land use change is controversial because the region is prone to drought, and replacement of native vegetation by planted forests may increase vegetation water use. The goal of this study was to examine the physiological differences, especially the response of water flux and canopy conductance to microclimate, that lead to greater water use by exotic trees compared to native trees. Meteorological variables and sapflow density of P. menziesii and four native woody species were measured in the growing season 2005–2006. Canopy conductance (gc) was estimated for both the exotic (monoculture) and native (multi-species) systems, including the individual contributions of each species of the native forest. Sapflow density, stand-level transpiration and gc were related to leaf-to-air vapor pressure difference (VPD). All native species had different magnitudes and diurnal patterns of sapflow density compared to P. menziesii, which could be explained by the different gc responses to VPD. Stomatal sensitivity to VPD suggested that all native species have a stronger stomatal control of leaf water potential and transpiration due to hydraulic limitations compared to P. menziesii. In conclusion, differences in water use between a P. menziesii plantation and a contiguous native mixed forest of similar basal area could be explained by different gc responses to VPD between species (higher sensitivity in the native species), in addition to particular characteristics of the native forest structure.     

Assessing the extent and diversity of riparian ecosystems in Sonora, Mexico

Conservation of forested riparian ecosystems is of international concern. Relatively little is known of the structure, composition, diversity, and extent of riparian ecosystems in Mexico. We used high- and low-resolution satellite imagery from 2000 to 2006, and ground-based sampling in 2006, to assess the spatial pattern, extent, and woody plant composition of riparian forests across a range of spatial scales for the state of Sonora, Mexico. For all 3rd and higher order streams, river bottomlands with riparian forests occupied a total area of 2,301 km². Where forested bottomlands remained, on average, 34% of the area had been converted to agriculture while 39% remained forested. We estimated that the total area of riparian forest along the principal streams was 897 km². Including fencerow trees, the total forested riparian area was 944 km², or 0.5% of the total land area of Sonora. Ground-based sampling of woody riparian vegetation consisted of 92, 50 m radius circular plots. About 79 woody plant species were noted. The most important tree species, based on cover and frequency, were willow species Salix spp. (primarily S. goodingii and S. bonplandiana), mesquite species Prosopis spp. (primarily P. velutina), and Fremont cottonwood Populus fremontii. Woody riparian taxa at the reach scale showed a trend of increasing diversity from north to south within Sonora. Species richness was greatest in the willow-bald cypress Taxodium distichum var. mexicanum—Mexican cottonwood P. mexicana subsp. dimorphia ecosystem. The non-native tamarisk Tamarix spp. was rare, occurring at just three study reaches. Relatively natural stream flow patterns and fluvial disturbance regimes likely limit its establishment and spread.     

Conservation value of timber quality versus associated non-timber quality stands for understory diversity in Nothofagus forests

Conservation strategies of forested landscapes must consider biodiversity of the included site types, i.e. timber-quality forests and associated non-timber-quality stands. The objectives were to characterize forest overstory structure in timber-quality versus associated non-timber-quality stands; and to compare their understory communities. Six forest types were sampled in Nothofagus forests of Tierra del Fuego (Argentina): two timber-quality N. pumilio forests, and four associated non-timber-quality stands (edge, N. antarctica, wetlands and streamside forests). Overstory structure and understory vegetation (species richness, frequencies, cover and biomass) were characterized during spring and summer seasons. Analysis of variance and multivariates were carried out. Overstory structure differed across the site types, with higher tree size, canopy closure and tree volume in timber-quality stands. Fifty-one understory plant species were observed, but understory variables varied with site types, especially wetlands (highest native and exotic richness, cover and biomass, and 25% of exclusive species). Site types were grouped in three: N. antarctica stands, streamside stands and the other N. pumilio forests according to multivariate analysis. Forty three percent of plants were distributed in all site types, and all timber-quality forest understory species were present in some associated non-timber-quality stands. Timber-quality N. pumilio forests have a marginal value for understory conservation compared to associated non-timber-quality stands, because these last include all the plants observed in timber-quality forests and also possess many exclusive species. Therefore, protection of associated non-timber-quality stands during forest management planning could increase understory conservation at landscape level, and these could be better reserves of understory diversity than retentions of timber-quality stands.     

Disturbance and climatic influences on age structure of ponderosa pine at the pine/grassland ecotone, Colorado Front Range

Abstract. In the Colorado Front Range, disturbances and climatic variation influence stand structure of ponderosa pine (Pinus ponderosa) along the lower timberline ecotone. Over the past 100 years there has been a shift to a greater density and extent of ponderosa pine at the forest-grassland boundaries. Ponderosa pine regeneration at lower timberline appears to be influenced by fires in the 1860s and decreased grazing pressure in the 1970s–1980s. Climatic variation may also have influenced age structure, even though analyses of age structure at a 10-year class scale prevented the detection of climatic influences occurring at a finer scale. These changes in disturbance regimes, possibly together with moister springs/early summers, created favourable conditions for the increase in density and extent of ponderosa pine at lower timberline ecotone.     

Loss of oak dominance in dry-mesic deciduous forests predicted by gap capture methods

Gap capture methods predict future forest canopy species composition from the tallest trees growing in canopy gaps rather than from random samples of shaded understory trees. We used gap capture methods and a simulation approach to forecast canopy composition in three old oak forests (Quercus spp.) on dry-mesic sites in southern Wisconsin, USA. In the simulation, a gap sapling is considered successful if it exceeds a threshold height of 13–17 m (height of maximum crown width of canopy trees) before its crown center can be overtopped by lateral crown growth of mature trees. The composition of both the tallest gap trees and simulated gap captures suggests that 68–90% of the next generation of canopy trees in the stands will consist of non-Quercus species, particularly Ulmus rubra, Carya ovata and Prunus serotina. Quercus species will probably remain as a lesser stand component, with Quercus alba and Quercus rubra predicted to comprise about 19% of successful gap trees across the three stands. Several methods of predicting future canopy composition gave similar results, probably because no gap opportunist species were present in these stands and there was an even distribution of species among height strata in gaps. Gap trees of competing species already average 11–13 m tall, and mean expected time for these trees to reach full canopy height is only 19 years. For these reasons, we suggest that dominance will shift from oaks to other species, even though late successional species (e.g., Acer and Tilia) are not presently common in the understories of these stands.     

The dynamics of old-growth Pseudotsuga forests in the western Cascade Range,Oregon, USA

Size and age structure, spatial analysis, and disturbance history were used to analyse the population structures and regeneration patterns of 8 conifer stands in the central western Cascade Range, Oregon, USA. Variation in forest structure reflected the effects of frequent (20–50 yr) low-intensity fires and treefalls, infrequent (100–200 yr) localised, intense fires, and extensive fires that resulted in stand replacement (every ca 400 yr?).The amount of canopy removed and the size of openings formed by fires and treefalls were important determinants of subsequent forest establishment. Single or several species stands of Pseudotsuga and/or Abies procera, or mixed species stands of Pseudotsuga, Abies procera, Tsuga heterophylla and Abies amabilis established in openings where intense fires had removed most of the canopy trees over several ha. Multi-tiered and multi-aged stands, often containing 400–500 yr-old Pseudotsuga and variously-sized more or less even-aged patches of younger shade tolerant Tsuga heterophylla and/or Abies amabilis, occurred where lower-intensity fires did not kill all overstorey trees or where treefalls occurred after the initial fire.Current regeneration processes are influenced by overstorey composition, the availability and size of canopy openings, and the availability of substrates suitable for regeneration. Tsuga heterophylla and Abies amabilis established under Pseudotsuga menziesii and Abies procera canopies and in small canopy openings (<400 m²) created by windfalls, but rarely under Tsuga. Down logs and stumps were favoured establishment sites for Tsuga.The disturbance regime of fires of low-to moderate-intensity, windfalls, and occasional fires that result in extensive stand replacement contrasts with the pattern of infrequent, catastrophic disturbances proposed for other areas of the Pacific Northwest. Although fires at stand establishment commonly determine much of the composition, structure, and subsequent stand development, canopy replacement by shade tolerant species occurs as the different life histories of the species are expressed in response to various disturbances differing in intensity and frequency. Such a non-equilibrium view of vegetation change is consistent with many other fire-dominated forests of the western United States.     

Long- and short-term effects of reindeer grazing on tundra wetland vegetation

We studied long-term (50 years) and short-term (4 years) effects of summer grazing of reindeer on subarctic tundra wetland vegetation. The long-term effects of summer grazing were studied by comparing vegetation on Finnish and Norwegian sides of the fence line separating reindeer grazing regimes. The Finnish side was intensively grazed and trampled throughout the year, whereas the Norwegian side was grazed in winter. Experimental fences were erected to examine short-term effects of grazing exclusion. Both in the long- and short-term, summer grazing decreased the height of Salix lapponum whereas the short-term effects on willow cover were less clear than the long-term effects. In contrast, Carex spp. benefited from grazing. Long-term grazing had little effect on total bryophyte cover. Grazing had negligible effects on the nutrient content of leaves of S. lapponum and Eriophorum angustifolium. We conclude that tundra wetlands can withstand moderately high grazing pressure sustained over several decades.