Background and aims: As low initial uptake and essentially zero later uptake limit efficacy of N fertilization for temperate conifers, we investigated factors limiting long-term tree uptake of ...residual super(15)N-labeled fertilizer. Methods: We used a pot bioassay to assess availability of super(15)N from soil sampled 10 years after fertilization of a Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco) stand with super(15)N-urea (200 kg N ha super(-1)). Douglas-fir seedlings were grown for 2 years in organic (designated LFH) and mineral soil (0-10 cm) layers reconstructed from control and fertilized plots; residual fertilizer N amounted to 10 % of LHF and 5 % of MIN N. Results: Percentage recovery of residual super(15)N in seedlings was not affected by the original season of fertilization (spring vs. fall), but differed by the source of super(15)N excess. LFH was a better source of residual super(15)N; 12.4 % of residual LFH super(15)N was taken up by seedlings and 7.6 % transferred to soil, whereas mineral soil yielded only 8.3 % of residual super(15)N to seedling uptake and 2.4 % to LFH. Extractable inorganic N was 2-3 orders of magnitude higher in fallow pots. Conclusions: Ten-year residual fertilizer super(15)N was clearly cycling between LFH and mineral soil and available to seedlings, indicating that other factors such as denitrification, leaching, and asynchrony of soil N mineralization and tree uptake limit long-term residual N fertilizer uptake in the field.
•Pinus ponderosa had greater drought resistance compared to Pseudotsuga menziesii.•Fuel treatments improved drought resistance.•Benefits of drought resistance from fuel treatments were dependent upon ...tree characteristics, drought severity, and species.
Drought is an important stressor in forest ecosystems that can influence tree vigor and survival. In the U.S., forest managers use two primary management techniques to promote resistance and resilience to drought: prescribed fire and mechanical thinning. Generally applied to reduce fuels and fire hazard, treatments may also reduce competition for resources that may improve tree-growth and reduce mortality during drought. A recent severe and prolonged drought in California provided a natural experiment to investigate tree-growth responses to fuel treatments and climatic stress. We assessed tree-growth from 299 ponderosa pine (Pinus ponderosa) and Douglas-fir (Pseudotsuga menziesii) in treated and untreated stands during severe drought from 2012 to 2015 in the mixed-conifer forests of Whiskeytown National Recreation Area (WNRA) in northern California. The treatment implemented at WNRA removed 34% of live basal area through mechanical thinning with a subsequent pile burning of residual fuels. Tree-growth was positively associated with crown ratio and negatively associated with competition and a 1-year lag of climate water deficit, an index of drought. Douglas-fir generally had higher annual growth than ponderosa pine, although factors affecting growth were the same for both species. Drought resistance, expressed as the ratio between mean growth during drought and mean growth pre-drought, was higher in treated stands compared to untreated stands during both years of severe drought (2014 and 2015) for ponderosa pine but only one year (2014) for Douglas-fir. Thinning improved drought resistance, but tree size, competition and species influenced this response. On-going thinning treatments focused on fuels and fire hazard reduction are likely to be effective at promoting growth and greater drought resistance in dry mixed-conifer forests. Given the likelihood of future droughts, land managers may choose to implement similar treatments to reduce potential impacts.
The role of mycorrhizal networks in forest dynamics is poorly understood because of the elusiveness of their spatial structure. We mapped the belowground distribution of the fungi Rhizopogon ...vesiculosus and Rhizopogon vinicolor and interior Douglas-fir trees (Pseudotsuga menziesii var. glauca) to determine the architecture of a mycorrhizal network in a multi-aged old-growth forest. Rhizopogon spp. mycorrhizas were collected within a 30 x 30 m plot. Trees and fungal genets were identified using multi-locus microsatellite DNA analysis. Tree genotypes from mycorrhizas were matched to reference trees aboveground. Two trees were considered linked if they shared the same fungal genet(s). The two Rhizopogon species each formed 13-14 genets, each colonizing up to 19 trees in the plot. Rhizopogon vesiculosus genets were larger, occurred at greater depths, and linked more trees than genets of R. vinicolor. Multiple tree cohorts were linked, with young saplings established within the mycorrhizal network of Douglas-fir veterans. A strong positive relationship was found between tree size and connectivity, resulting in a scale-free network architecture with small-world properties. This mycorrhizal network architecture suggests an efficient and robust network, where large trees play a foundational role in facilitating conspecific regeneration and stabilizing the ecosystem.
Separating edaphic impacts on tree distributions from those of climate and geography is notoriously difficult. Aboveground and belowground factors play important roles, and determining their relative ...contribution to tree success will greatly assist in refining predictive models and forestry strategies in a changing climate.
In a common glasshouse, seedlings of interior Douglas-fir (Pseudotsuga menziesii var. glauca) from multiple populations were grown in multiple forest soils. Fungicide was applied to half of the seedlings to separate soil fungal and nonfungal impacts on seedling performance. Soils of varying geographic and climatic distance from seed origin were compared, using a transfer function approach.
Seedling height and biomass were optimized following seed transfer into drier soils, whereas survival was optimized when elevation transfer was minimised. Fungicide application reduced ectomycorrhizal root colonization by c. 50%, with treated seedlings exhibiting greater survival but reduced biomass.
Local adaptation of Douglas-fir populations to soils was mediated by soil fungi to some extent in 56% of soil origin by response variable combinations. Mediation by edaphic factors in general occurred in 81% of combinations. Soil biota, hitherto unaccounted for in climate models, interacts with biogeography to influence plant ranges in a changing climate.
Abstract
Ectomycorrhizal (EcM) fungal communities may be shaped by both deterministic and stochastic processes, potentially influencing ecosystem development and function. We evaluated community ...assembly processes for EcM fungi of
Pseudotsuga menziesii
among 12 sites up to 400 km apart in southwest British Columbia (Canada) by investigating species turnover (β‐diversity) in relation to soil nitrogen (N) availability and physical distance. We then examined functional traits for an N‐related niche by quantifying net fluxes of
,
and protons on excised root tips from three contrasting sites using a microelectrode ion flux measurement system. EcM fungal communities were well aligned with soil N availability and
pH
, with no effect of site proximity (distance–decay curve) on species assemblages. Species turnover was significant (β
1/2
= 1.48) along soil N gradients, with many more
Tomentella
species on high N than low N soils, in contrast to
Cortinarius
species. Ammonium uptake was greatest in the spring on the medium and rich sites and averaged over 190 nmol/m
2
/s for
Tomentella
species. The lowest uptake rates of
were by nonmycorrhizal roots of axenically grown seedlings (10 nmol/m
2
/s), followed by
Cortinarius
species (60 nmol/m
2
/s). EcM roots from all sites displayed only marginal uptake of nitrate (8.3 nmol/m
2
/s). These results suggest
uptake capacity is an important functional trait influencing the assembly of EcM fungal communities. The diversity of EcM fungal species across the region arguably provides critical belowground adaptations to organic and inorganic N supply that are integral to temperate rainforest ecology.
Wood nitrogen isotope composition ( delta super(15)N) provides a potential retrospective evaluation of ecosystem N status but refinement of this index is needed. We calibrated current wood delta ...super(15)N of Douglas-fir (Pseudotsuga menziesii), an ectomycorrhizal tree species, against a productivity gradient of contrasting coastal forests of southern Vancouver Island (Canada). We then examined historical delta super(15)N via increment cores, and tested whether wood delta super(15)N corresponded with climatic fluctuations. Extractable soil N ranged from 11 to 43 kg N ha super(-1) along the productivity gradient, and was characterized by a progressive replacement of N forms (amino acids, NH sub(4) super(+) and NO sub(3) super(-)). Current wood delta super(15)N was significantly less depleted (-5.0 to -2.6 ppt) with increasing productivity, although linear correlations were stronger with Delta delta super(15)N (the difference between wood and soil delta super(15)N) to standardize the extent of isotopic fractionation by ectomycorrhizal fungi. An overall decline in wood delta super(15)N of 0.9 ppt over the years 1900-2009 was detected, but trends diverged widely among plots, including positive, negative and no trend with time. We did not detect significant correlations in detrended wood delta super(15)N with mean annual temperature or precipitation. The contemporary patterns in stand productivity, soil N supply and wood delta super(15)N were moderately strong, but interpreting historical patterns in delta super(15)N was challenging because of potential variations in N uptake related to stand dynamics. The lack of wood delta super(15)N correlations with climate may be partly due to methodological limitations, but might also reflect the relative stability in N supply due to the overriding constraints of soil organic matter quantity and quality.
Climate change is increasing fire activity in the western United States, which has the potential to accelerate climate-induced shifts in vegetation communities. Wildfire can catalyze vegetation ...change by killing adult trees that could otherwise persist in climate conditions no longer suitable for seedling establishment and survival. Recently documented declines in postfire conifer recruitment in thewestern United States may be an example of this phenomenon. However, the role of annual climate variation and its interaction with long-term climate trends in driving these changes is poorly resolved. Here we examine the relationship between annual climate and postfire tree regeneration of two dominant, low-elevation conifers (ponderosa pine and Douglas-fir) using annually resolved establishment dates from 2,935 destructively sampled trees from 33 wildfires across four regions in the western United States. We show that regeneration had a nonlinear response to annual climate conditions, with distinct thresholds for recruitment based on vapor pressure deficit, soil moisture, and maximum surface temperature. At dry sites across our study region, seasonal to annual climate conditions over the past 20 years have crossed these thresholds, such that conditions have become increasingly unsuitable for regeneration. High fire severity and low seed availability further reduced the probability of postfire regeneration. Together, our results demonstrate that climate change combined with high severity fire is leading to increasingly fewer opportunities for seedlings to establish after wildfires and may lead to ecosystem transitions in low-elevation ponderosa pine and Douglas-fir forests across the western United States.
Mycorrhizal networks (MNs) are fungal hyphae that connect roots of at least two plants. It has been suggested that these networks are ecologically relevant because they may facilitate interplant ...resource transfer and improve regeneration dynamics. This study investigated the effects of MNs on seedling survival, growth and physiological responses, interplant resource (carbon and nitrogen) transfer, and ectomycorrhizal (EM) fungal colonization of seedlings by trees in dry interior Douglas-fir (Pseudotsuga menziesii var. glauca) forests. On a large, recently harvested site that retained some older trees, we established 160 isolated plots containing pairs of older Douglas-fir "donor" trees and either manually sown seed or planted Douglas-fir "receiver" seedlings. Seed- and greenhouse-grown seedlings were sown and planted into four mesh treatments that served to restrict MN access (i.e., planted into mesh bags with 0.5-, 35-, 250-μm pores, or without mesh). Older trees were pulse labeled with carbon (¹³CO₂) and nitrogen (¹⁵NH₄¹⁵NO₃) to quantify resource transfer. After two years, seedlings grown from seed in the field had the greatest survival and received the greatest amounts of transferred carbon (0.0063% of donor photo-assimilates) and nitrogen (0.0018%) where they were grown without mesh; however, planted seedlings were not affected by access to tree roots and hyphae. Size of "donor" trees was inversely related to the amount of carbon transferred to seedlings. The potential for MNs to form was high (based on high similarity of EM communities between hosts), and MN-mediated colonization appeared only to be important for seedlings grown from seed in the field. These results demonstrate that MNs and mycorrhizal roots of trees may be ecologically important for natural regeneration in dry forests, but it is still uncertain whether resource transfer is an important mechanism underlying seedling establishment.
There is a general assumption that intraspecific populations originating from relatively arid climates will be better adapted to cope with the expected increase in drought from climate change. For ...ecologically and economically important species, more comprehensive, genecological studies that utilize large distributions of populations and direct measures of traits associated with drought‐resistance are needed to empirically support this assumption because of the implications for the natural or assisted regeneration of species. We conducted a space‐for‐time substitution, common garden experiment with 35 populations of coast Douglas‐fir (Pseudotsuga menziesii var. menziesii) growing at three test sites with distinct summer temperature and precipitation (referred to as ‘cool/moist’, ‘moderate’, or ‘warm/dry’) to test the hypotheses that (i) there is large genetic variation among populations and regions in traits associated with drought‐resistance, (ii) the patterns of genetic variation are related to the native source‐climate of each population, in particular with summer temperature and precipitation, (iii) the differences among populations and relationships with climate are stronger at the warm/dry test site owing to greater expression of drought‐resistance traits (i.e., a genotype × environment interaction). During midsummer 2012, we measured the rate of water loss after stomatal closure (transpirationₘᵢₙ), water deficit (% below turgid saturation), and specific leaf area (SLA, cm² g⁻¹) on new growth of sapling branches. There was significant genetic variation in all plant traits, with populations originating from warmer and drier climates having greater drought‐resistance (i.e., lower transpirationₘᵢₙ, water deficit and SLA), but these trends were most clearly expressed only at the warm/dry test site. Contrary to expectations, populations from cooler climates also had greater drought‐resistance across all test sites. Multiple regression analysis indicated that Douglas‐fir populations from regions with relatively cool winters and arid summers may be most adapted to cope with drought conditions that are expected in the future.
Most temperate woody plants have a winter chilling requirement to prevent budburst during mid-winter periods of warm weather. The date of spring budburst is dependent on both chilling and forcing; ...modeling this date is an important part of predicting potential effects of global warming on trees. There is no clear evidence from the literature that the curves of chilling or forcing effectiveness differ by species so we combined our data and published information to develop new curves on the effectiveness of temperature for chilling and forcing. The new curves predict effectiveness over a wide range of temperatures and we suggest both functions may be operating at the same time. We present experimental data from 13 winter environments for 5 genotypes of Douglas-fir (
Pseudotsuga menziesii var.
menziesii) and use them to test various assumptions of starting and stopping dates for accumulating chilling and forcing units and the relationship between budburst and the accumulation of chilling and forcing units. Chilling started too early to be effective in one treatment but the other 12 environments resulted in budburst from many combinations of chilling and forcing. Previous reports have suggested benefits or cancellations of effects from alternating day/night or periodic temperatures. Our simple models do not include these effects but nevertheless were effective in predicting relationships between chilling and forcing for treatments with a wide range of conditions. Overall, the date of budburst changed only slightly (+1 to −11 days) across a wide range of treatments in our colder test environment (Olympia, WA, USA) but was substantially later (+29 days) in the warmest treatment in our warmer environment (Corvallis, OR, USA). An analysis of historical climate data for both environments predicted a wide range in date to budburst could result from the same mean temperature due to the relative weightings of specific temperatures in the chilling and forcing functions. In the absence of improved understanding of the basic physiological mechanisms involved in dormancy induction and release, we suggest that simple, universal functions be considered for modeling the effectiveness of temperature for chilling and forcing. Future research should be designed to determine the exact shape of the curves; data are particularly lacking at the temperature extremes. We discuss the implications of our data and proposed functions for predicting effects of climate change. Both suggest that the trend toward earlier budburst will be reversed if winter temperatures rise substantially.
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