Differences in insect species richness and faunal composition of birch seedlings, saplings and trees: the importance of plant architecture

Abstract. 1. Data are presented on the species richness and faunal composition of herbivorous insects on birch seedlings, saplings and trees at one site in Northern England.
2. Species richness of insect herbivores in equal-sized samples from birch seedlings and trees was similar through most of the season.
3. Effects of plant architecture were confined to the first sampling date, when seedling faunas were species poor compared with trees – possibly due to safe overwintering sites on the extensive bark, twigs and buds of trees.
4. The faunal composition of birch seedlings, saplings and trees was also similar. Out of a total of 112 recorded species of herbivores, only one aphid species was confined to seedlings.
5. Similarly, no evidence for clear-cut vertical stratification of insects within trees was found.
6. Species turnover as host plants mature ('horizontal' stratification) and vertical stratification within trees add little to the high overall species richness of birch-feeding insects in Britain, contrary to the predictions of Lawton (1983).     

Comparative anatomy of resin ducts of the Pinaceae

 Resin ducts are common in the Pinaceae. The comparative anatomy of stems and leaves of 50 species and two varieties from ten genera has been investigated. The structure and distribution of resin ducts differ among genera. Resin ducts occur in foliage leaves of ten genera of Pinaceae. Cortical resin ducts are absent in the stems of Pseudolarix and Larix. Resin ducts only occur in the secondary xylem of stems of Pinus, Picea, Cathaya, Larix, Pseudotsuga and some Keteleeria species. All of the epithelial and sheath cells are alive and thin-walled in the resin ducts of stem cortex and mesophyll. Except for Pinus the epithelial cells of resin ducts in the secondary xylem of stems have thick, lignified walls. Comparative study shows there are obvious differences in the resin ducts of different genera; apparent differences do not exist, however, in the resin ducts of different species of the same genus. According to the structure and distribution of the resin ducts in ten genera of Pinaceae, a synoptical arrangement of the genera is given and generic relationships of the Pinaceae are discussed. Received: 12 September 1995 / Accepted: 14 March 1996     

The ubiquity of natural adsorbable organic halogen production among basidiomycetes

 Recently, several species of basidiomycetes were shown to produce de novo high concentrations of chloroaromatic metabolites. Since these lignocellulose-degrading fungi play a major role in the ecosphere, the purpose of this study was to determine the ubiquity of organohalogen production among basidiomycetes. A total of 191 fungal strains were monitored for adsorbable organic halogen (AOX) production when grown on defined liquid media. Approximately 50% of the strains tested and 55% of the genera tested produced AOX. A low production of 0.1–0.5 mg AOX/l was observed among 25% of the strains, a moderate production of 0.5–5.0 mg AOX/l was observed among 16% of the strains and 9% of the strains produced high levels (5–67 mg AOX/l). The latter group was dominated by species belonging to the genera Hypholoma, Mycena and Bjerkandera, showing specific AOX productions in the range 1074–30893 mg AOX/kg dry weight of mycelial biomass. Many highly ecologically significant fungal species were identified among the moderate to high producers. These species were also able to produce AOX when cultivated on natural lignocellulosic substrates. Hypholoma fasciculare and Mycena metata respectively produced up to 132 mg and 193 mg AOX/kg dry weight of forest litter substrate in 6 weeks. Received: 5 October 1995/Received revision: 28 December 1995/Accepted: 12 February 1996     

Differential drought responses between saplings and adult trees in four co-occurring species of New England

Tree-ring characteristics in four species were examined to address whether co-occurring mature trees of different successional status respond differently to drought, and whether saplings of these species have a greater response to drought than mature trees. We examined saplings and mature trees of paper birch, yellow birch, red maple and sugar maple, which varied in successional status (shade-tolerance) and co-occurred at Harvard Forest, Petersham, Mass., USA. Three drought events in 1964–1966, 1981 and 1995 were identified using climate data. For mature trees, there was no significant interspecific difference in relative changes in ring-width index (RWI) during the 1964–1966 and 1995 drought events. However, the interspecific difference was significant in the 1981 drought event. Response function analysis for mature trees showed that the radial growth of sugar maple was mainly controlled by spring and summer precipitation, red maple by spring and summer precipitation and temperature, yellow birch by winter and summer precipitation, and spring and summer temperature, and paper birch by spring and summer precipitation and spring temperature. Saplings of sugar maple and yellow birch, but not red maple and paper birch, showed significant positive correlations between RWI and annual total precipitation. In the 1995 drought event, saplings and mature trees of red maple and paper birch differed significantly in drought responses, but this was not true in sugar maple and yellow birch. Our results do not support a generally greater response in saplings than in mature trees, nor an early- versus late successional difference in drought responses.     

Effect of pH on transfructosylation and hydrolysis by β-fructofuranosidase from Aspergillus oryzae

 β-Fructofuranosidase was purified from commercial alkaline protease (Aspergillus oryzae origin). The optimal pH of its transfructosylating activity was more alkaline (pH 8) than that of its hydrolyzing activity (pH 5). In the case of a 24-h reaction with sucrose, the hydrolysis and transfructosylation reaction were optimal at pH 4–5 and pH 8, respectively. In the reaction at pH 8 1-kestose and nystose were the main fructooligosaccharides produced. The transfer ratio was hardly different between pH 5 and pH 8 early in the reaction, but the transfer products (1-kestose and nystose) were decreased at pH 5 as the reaction proceeded because of their hydrolysis. Received: 18 January 1995/Received last revision: 23 August 1995/Accepted: 13 September 1995     

Seedling growth strategies and seed size effects in fourteen oak species native to different soil moisture habitats

 Seedling growth and morphology are thought to reflect evolutionary responses to habitat or influences of seed size. To test these hypotheses, we selected fourteen species of North American oaks differing in soil moisture habitat preference and seed size. Seedlings were grown for 1 – 2 years with abundant soil water and moderate soil nutrition in pots placed outdoors and in a common garden. Oak species native to xeric environments produced the smallest seedlings. Oaks from hydric soils had more shoot weight per unit of root weight and more height per unit of total plant weight than did mesic or xeric oaks. Essentially no differences in leaf area per unit of total plant weight were detected. Species with thinner and larger individual leaves tended to produce larger seedlings. Within species, seed size was generally unrelated to seedling growth, although results may have been complicated by uncontrolled genotypic variability. However, when species were compared, those with larger mean seed size produced larger seedlings. Root/shoot allometry, height growth and leaf thickness in the tested species may reflect evolutionary responses to soil moisture and flooding. Although seed size influenced seedling growth, no clear relationship between seed size and soil moisture habitat was found. Received: 26 March 1995 / Accepted: 30 November 1995     

The use of RAPD for verifying the apomictic status of seedlings of Malus species

 The lack of red-purple pigmentation in seedlings obtained from crosses between M. cv Baskatong, carrying a dominant homozygous gene for red-purple pigmentation, and other species has been used for the detection of apomictic plants in Malus species. RAPD marker techniques were employed to evaluate the genetic similarity between putative apomictic seedlings and their female parents. From the selected set of 20 (OPA) primers about half were able to detect hybrids from the apomictic seedlings, if present. RAPD analyses confirmed the usefulness of the colour-marker gene in detecting the hybrids in vitro for seedlings of M. toringoides×M. cv Baskatong, but not for crosses involving M. hupehensis×M. cv Baskatong where in vitro colour-based selection was not possible (due to red stems in all cases). The set of primers (OPA-01, 02, 08, 09, 10, 12, 13, 14, 16, 18 and 20) clearly determined the hybrid nature of seedlings and allowed the selection of apomictic ones. Therefore, although cv Baskatong is useful as an indicator, these data show that this technique is not applicable in all cases. Received: 22 May 1997 / Accepted: 28 July 1997     

Vegetation trajectories of Korean red pine (Pinus densiflora Sieb. et Zucc.) forests at Mt. Seorak, Korea

The vegetation dynamics of Korean red pine (Pinus densiflora) forests were investigated at Mt. Seorak, Korea. Our Detrended Correspondence Analysis (DCA) classified the forests into four types: ridge top, upper slope, lower slope/hill, and streamside. The ridge top forests were likely to sustain themselves, as suggested by the large proportion of seedlings and saplings (89% at <25 years old) and the relatively high density ofP. densiflora (2388 stems ha^-1). Periodic disturbances, such as flash floods, made the streamside inhospitable to late-successional species. Such conditions may have provided a favorable environment for the recruitment ofP. densiflora seedlings through increased solar radiation and decreased competition with other species. On the upper slopes, the dominance ofQuercus seedlings and saplings (63% at >25 years old, and a density of 3263 stems ha^-1) suggests a transition from pine to oak forest. Extensive human interventions appeared to arrest the natural succession from pine to oak forests on the lower slope/hill, while encouraging invasions by forest-edge and introduced species (e.g.,Rosa multiflora andRobinia pseudoacacia).     

Tracing water uptake by jarrah (Eucalyptus marginata) trees using natural abundances of deuterium

 Seasonal change in the δ²H content of water from twig sap, soil, rainfall and groundwater were measured to determine the water sources accessed by jarrah (Eucalyptus marginata) trees at three sites in Western Australia with differing soils and depths to water table. During winter and spring the main contributor to the water uptake of the trees was stored water in the surface layers of the soil replenished by predominantly winter rainfall. With the onset of summer drought jarrah became more reliant on water from deeper down the profile. There was no clear evidence that jarrah could tap water from groundwater more than 14 m deep in deep sands. Defining the source of water for trees in deep lateritic soils using stable isotopes is hampered by the uniform deuterium profiles down most of the unsaturated zone and into the groundwater. There was a limited response in the δ²H values of sapwater in twigs to changes in the δ²H of the upper layers of the deep sand following input of rainfall in autumn. The damped response was related to the small variation in the δ²H composition of rainfall in most events during the year and the mixing in the tree of water extracted from different locations in the soil profile. Received: 21 August 1995 / Accepted: 3 December 1995     

Hydrogen fluoride effects on plasma membrane composition and ATPase activity in needles of white pine (Pinus strobus) seedlings pretreated with 12 h photoperiod

 Eastern white pine (Pinus strobus L.) seedlings were pretreated with 12 h photoperiod to induce dormancy. Dormant plants were fumigated with 0.5 ppb (0.4 μg m^–3) or 2.0 ppb (1.6 μg m^–3) hydrogen fluoride (HF) for 2 – 28 days. Plasma membranes were isolated from needles of treated and control seedlings to determine their chemical composition and ATPase activity. For all analyses, only those plants which did not show needle necrosis were selected. The amount of plasma membrane phospholipid expressed on a plasma membrane protein basis was higher after 2 days in the 0.5 ppb HF treatment as compared to controls. After 2 days of 2.0 ppb HF treatment as well as after 8 and 28 days of both HF treatments phospholipid to protein ratios in fluoride treated seedlings were lower as compared to control levels. A decrease in sterol levels could be observed after 2 days in both HF treatments. A large increase in the ratio of sterols to proteins was observed in plasma membranes of eastern white pine seedlings treated with 0.5 ppb HF for 28 days. Increased sterol to phospholipid ratios were observed after 8 and 28 days in 0.5 ppb and after 2 and 8 days of 2.0 ppb HF treatment. A decrease in ATPase activity was observed after 8 days with both fluoride treatments. Drastic increase of ATPase activity was observed after 28 days of HF treated plants. Observed changes of sterol and phospholipid levels after only 2 days of fumigation suggest early fluoride effects on plasma membrane composition during plant dormancy. Received: 25 October 1995 / Accepted: 24 May 1996     

Effects of increasing saturation vapour pressure deficit on growth and ABA levels in black spruce and jack pine

 Plant responses to saturation vapour pressure deficit (SVPD) were studied by subjecting black spruce [Picea mariana (Mill) B.S.P.] and jack pine seedlings (Pinus banksiana Lamb.) to humid (0.3 – 0.8 kPa) or dry (2.0 – 2.5 kPa SVPD) regimes for 4 weeks using a computer-controlled environmental system to control diurnal variation in SVPD. Dry matter accumulation in needles was not altered by increasing SVPD. However, root growth declined by 60% which increased shoot to root ratio and reduced total seedling dry weight in both black spruce and jack pine. Relative growth rate of jack pine also declined to about half the rate of plants grown under humid conditions. In situ root marking studies showed that the decline in root growth of jack pine under the high SVPD was the result of reduced lateral root initiation, whereas root elongation was unaffected by humidity. A 4-week exposure to dry air increased abscisic acid (ABA) levels in needles, but not roots, of jack pine whereas ABA levels in black spruce were not altered. A short (3-day) exposure failed to increase needle ABA levels in either species. These results suggest that the responses of conifers to dry air were not the result of ABA accumulation. Received: 24 March 1996 / Accepted: 30 May 1996     

Taxonomy of Pinus species based on the seed oil fatty acid compositions

 The fatty acid compositions of the seed oils from ten pine species have been established by capillary gas-liquid chromatography of the methyl esters. With regard to either normal fatty acids or Δ5-olefinic acids, the general pattern of fatty acids did not differ from that of other pine seed oils reported previously. The main fatty acid was linoleic (9,12–18:2) acid (44.4–57.1%), followed by either oleic (9–18:1) acid (13.4–24.5%) or pinolenic (5,9,12–18:3) acid (1.5–25.2%). When applying multivariate analyses to the chemometric data (13 variables) of 49 pine species (ca. 40% of the living pine species), it was possible to distinguish between several sections: Pinea, Longifolia, Halepensis, Ponderosa-Banksiana, Sylvestris, and Cembra. The latter section was clearly divided into two sub-groups. A few species that presented a low overall content of Δ5-olefinic acids, and that grow in warm-temperate regions, were isolated from the bulk of other pine species. It is hypothesized that Δ5-olefinic acids might be related to cold-acclimation. Received: 5 June 1997 / Accepted: 17 August 1997     

Phytohormones in needles of healthy and declining silver fir (Abies alba Mill.): I. Indole-3-acetic acid

 Levels of indole-3-acetic acid (IAA) were determined in needles from silver fir (Abies alba Mill.) trees in the northern Black Forest. IAA was quantified by gas chromatography (GC) as 1-heptafluorobutyryl-IAA-methylester (HFB-IAA-ME) using electron capture detection. Prior to GC analysis, extensive purification of needle extracts was performed employing two HPLC steps. Peak identity of HFB-IAA-ME was confirmed by combined gas chromatography-mass spectrometry in selected samples. Levels of IAA in needles belonging to different needle age-classes exhibited a cyclic seasonal pattern with highest concentrations in winter and lowest levels in spring when bud-break occurred. Such a cyclic seasonal pattern of IAA levels was also observed in needles from declining fir trees or fir trees suffering from a strong sulfur impact (S-impact) in the field due to a local SO₂ source. Levels of IAA increased with increasing needle age. This age dependency of IAA concentrations was most pronounced in late autumn when IAA levels were high and nearly disappeared in spring when IAA levels reached their minimum. In needles from declining fir trees or fir trees suffering from a strong S-impact in the field, IAA levels hardly increased with increasing needle age. It is suggested that in healthy trees high levels of IAA protect older needles from abscission and that the considerable losses of older needles of declining fir trees or of fir trees under S-impact are a consequence of the low levels of IAA found in older needles of such trees. Received: 4 May 1995 / Accepted: 29 August 1995     

The Janzen-Connell effect on the population dynamics of a Fagus engleriana- Cyclobalanopsis oxyodon community in a subtropical zone of China

The Janzen-Connell (J-C) hypothesis provides a mechanism explaining the high species diversity in tropical rainforests. It postulates that predation could cause greater mortality on seeds and seedlings near their parental trees. In this study, we tested the hypothesis in a subtropical zone, a mixed evergreen-deciduous broad-leaved forest dominated by the Fagus engleriana and Cyclobalanopsis oxyodon. The study area was in the Shennongjia region, a key area of biodiversity conservation in both China and the world. The recruitment probability index was used to detect the J-C effect on nine species of the community, which were more than 50 individuals. Six large adults of each species were selected, and the numbers of saplings and adults were counted at the distance intervals of 0–5, 5–10, 10–15, 15–20, and 20–25 m from each focal tree. Two species in saplings stage and six in adult stage supported the J-C hypothesis, but their χ² was not significant. Three species, the F. engleriana, Rhododendron hypoglaucum, and Toona sinensis, showed a strong Hubbell pattern in the adult stage. Because of these results, we reject the J-C hypothesis and conclude that species could recruit near the conspecific trees in subtropical forest. The reasons why the J-C hypothesis fails to explain the species diversity in this community are the shortage of seed-consuming agents of subtropical forest and the influence of microsite topographic variation.     

Shake-flask culture of Laccaria laccata,an ectomycorrhizal basidiomycete

 Large-scale exploitation of the potential benefits of ectomycorrhizal fungi in improving plantation yields means that fermentation techniques for these fungi will be required. Starting with a base performance on a rich, complex medium, the effect of variations in some physicochemical culture parameters on biomass yield was studied. It was possible to reduce the amount of phosphate salts (to 1/9th) and other ingredients (to 1/3rd) in the medium. A shaking speed of either 100 rpm or 200 rpm in an orbital incubator was satisfactory and biomass yield responded to an increase in carbon substrate (glucose, from 10 g l^-1 and 20 g l^-1) though Y _x/s declined. An increase in inoculum size shortened culture time but decreased biomass yield. The upper limit of the incubation temperature was between 25°C and 30°C. Biomass yields were about 12 g l^-1 dry weight (Y _x/s=0.63) when 20 g l^-1 glucose was supplied, and about 7 g l^-1 (Y _x/s=0.74) when 10 g l^-1 glucose was supplied. Received: 9 October 1995/Accepted: 4 December 1995     

Response of seedling communities to mammalian pest eradication on Ulva Island, Rakiura National Park, New Zealand

Norway rats (Rattus norvegicus) were eradicated from Ulva Island, Rakiura National Park, in 1996. The aim of our work was to determine if seedlings and saplings increased in density and/or species richness following this eradication. In 2003, we took advantage of eight permanent plots (5 × 5 m) that had been established on Ulva Island in 1991, by counting seedlings and saplings of woody species, including tree ferns. Over this period, total numbers of woody seedlings (< 30 cm tall), and saplings (30 cm – 2 m tall) did not increase significantly (P > 0.05). However, seedlings of two species, Dicksonia squarrosa and Pittosporum colensoi, and saplings of one species, Coprosma colensoi, did increase in numbers significantly. Little change in the species richness of seedlings or saplings was recorded. The removal of Norway rats may have been partly responsible for the recorded increases in density, though the eradication of white-tailed deer (Odocoileus virginianus borealis) from Ulva Island approximately two decades prior to this work is an important confounding factor.     

Regulation of hydrogen metabolism in Butyribacterium methylotrophicum by substrate and pH

 Exogenous H₂/CO₂ and glucose were consumed simultaneously by Butyribacterium methylotrophicum when grown under glucose-limited conditions. CO₂ reduction to acetate was coupled to H₂ consumption. The addition of either H₂ or CO₂ to glucose batch fermentation resulted in an increase in cell density, hydrogenase (H₂-consuming and -producing) activities and fatty acid production by B. methylotrophicum as compared to when N₂ was the feed gas. Hydrogenase activities appeared to be tightly regulated and were produced at higher rates during the exponential phase when CO₂ was the feed gas as compared to H₂ or N₂. The increase in H₂-consuming activity and decrease in H₂-producing activity was correlated with an increase in butyrate synthesis. H₂-consuming and ferredoxin (Fd)–NAD reductase activities increased while H₂-producing and NADH–Fd reductase activities decreased in cells grown at pH 5.5 compared to those at pH 7.0. The molar ratio of butyrate/acetate was shifted from 0.35 at pH 7.0 to 1.22 at pH 5.5. The addition of exogenous H₂ did not decrease the butyrate/acetate ratio at pH 7.0 nor at pH 5.5. The results indicated that growth pH values regulated both hydrogenase and Fd–NAD oxidoreductase activities such that, at acid pH, more intermediary electron flow was directed towards butyrate synthesis than H₂ production. Received: 22 August 1995/Received revision: 18 December 1995/Accepted: 22 January 1996     

Stomatal and nonstomatal limitations of photosynthesis in relation to the drought and shade tolerance of tree species in open and understory environments

 Light saturated photosynthesis (A) in field saplings of shade tolerant, intermediate, and intolerant tree species was analyzed for stomatal and nonstomatal limitations to test differences between species and sun and shade phenotypes during drought. Throughout the study, photosynthesis was highest and mesophyll limitations of A (L_m) lowest in the intolerant species in both open and understory habitats. The shade tolerant species exhibited the only drought-related decreased A and increased L_m in the open, and the greatest drought-related decreased A and increased L_m in the understory. Few species exhibited significant habitat or drought-related differences in stomatal conductance to CO₂ (g_c), but even slight decreases in g_c during drought were associated with large increases in stomatal limitations to A (L_g). Combined changes in L_m and L_g resulted in increased relative stomatal limitation to A (l _g) in several species during drought. Nevertheless, the overall lack of stomatal closure allowed for nonstomatal limitations to play a major role in reduced A during drought. Higher leaf N was associated with shallower slope of the l _g versus g_c relationship, an indication of greater A capacity. Photosynthetic capacity tended to be greater in the intolerant species than the tolerant species, and it tended to decrease during drought primarily in the shade tolerant species in the understory. Findings in the literature suggest that carbon reduction reactions may be more susceptible to drought than photosynthetic light reactions. If so, reduced carbon reduction capacity of shade tolerant species or shade phenotypes may predispose them to drought conditions, which suggests a mechanism behind the well-recognized tradeoff between drought tolerance and shade tolerance of temperate tree species. Received: 20 October 1995 / Accepted: 20 February 1996     

Simultaneous utilization of pyridine and fructose by Rhodococcus opacus UFZ B 408 without an external nitrogen source

 A bacterium classified as Rhodococcus opacus, which is able to use pyridine (a potentially growth-inhibiting substrate) as its sole source of carbon, energy and nitrogen, was isolated. In a carbon-limited chemostat culture, the kinetics was determined for growth on both pyridine and a mixture of pyridine and fructose (9 mM/22.15 mM). With growth on pyridine, stable steady states were achieved up to dilution rates of about 0.1 h^-1. A further increase in the dilution rate resulted in the progressive accumulation of pyridine in the culture liquid and the cells were washed out. The maximum specific growth rate (μ_max = 0.23 h^-1) and the K _S value (0.22 mM) for growth on pyridine were determined from the residual pyridine concentrations measured within the range of stable steady states. With growth on the substrate mixture, the specific pyridine consumption rates and the residual pyridine concentrations were lower at similar dilution rates than with growth on pyridine alone, and stable steady states were established at dilution rates of up to 0.13 h^-1. The maximum pyridine degradation rate was enhanced to 270 mg pyridine l^-1 h^-1 compared to 210 mg pyridine l^-1 h^-1with growth on pyridine as a single substrate. An external nitrogen source did not need to be added in the case of growth on the substrate mixture. Fructose was assimilated by means of ammonium released from pyridine. Analysis of the nitrogen balance furnished proof that pyridine is an energy-deficient substrate; pyridine was assimilated and dissimilated at a ratio of 1 mol/0.67 mol respectively. The resulting yield coefficient was about 0.55 g dry weight/g pyridine. Moreover, it was demonstrated that, in regard to the biologically usable energy, 1 mol pyridine corresponds to 0.43 mol fructose. Received: 3 July 1995/Received revision: 19 October 1995/Accepted: 24 October 1995     

Competition and regeneration in quaking aspen–white fir (Populus tremuloides–Abies concolor) forests in the Northern Sierra Nevada,USA

Question: How does competition between quaking aspen (Populus tremuloides) and white fir (Abies concolor) affect growth and spatial pattern of each species? Location: The northern Sierra Nevada, California, USA. Methods: In paired plots in mixed aspen‐ (n=3) or white fir‐dominated (n=2) stands, we mapped trees and saplings and recorded DBH, height, species, and condition and took increment cores. We tallied seedlings by species. Tree ring widths were used as a measure of basal area change over the last decade, and canopy openness was identified using hemispherical photographs. Linear mixed models were used to relate neighborhood indices of competition, stand, and tree‐level variables to diameter increment. Spatial patterns of stems were identified using the Neighborhood Density Function. Results: White fir radial growth was higher in aspen‐ than white fir‐dominated plots. Individual‐level variables were more important for white fir than for aspen growth, while variables representing competitive neighborhood were important only for aspen. The forest canopy was more open in aspen‐ than white fir‐dominated stands, but ample aspen seedlings were observed in all stands. Canopy stems of aspen and white fir were randomly distributed, but saplings and small trees were clumped. Aspen saplings were repelled by canopy aspen stems. Conclusions: Variation in canopy openness explained more stand–stand variation in white fir than aspen growth, but high light levels were correlated with recruitment of aspen seedlings to the sapling class. Radial growth of aspen was predicted by indices of neighborhood competition but not radial growth of white fir, indicating that spacing and stem arrangement was more important for aspen than white fir growth. Fire suppression has removed a major disturbance mechanism that promoted aspen persistence and reduced competition from encroaching conifers, and current forests favor species that regenerate best by advance regeneration (white fir).