Ponderosa Pine Regeneration Research Articles

Abiotic Factors Modify Ponderosa Pine Regeneration Outcomes After High-Severity Fire

Abiotic Factors Modify Ponderosa Pine Regeneration Outcomes After High-Severity Fire

Environmental Influences on Density and Height Growth of Natural Ponderosa Pine Regeneration following Wildfires

Over the past century the size and severity of wildfires, as well as post-fire recovery processes (e.g., seedling establishment), have been altered from historical levels due to management policies and changing climate. Tree seedling establishment and growth drive future overstory tree dynamics after wildfire. Post-fire tree regeneration can be highly variable depending on burn severity, pre-fire forest condition, tree regeneration strategies, and climate; however, few studies have examined how different abiotic and biotic factors impact seedling density and growth and the interactions among those factors. We measured seedling density and height growth in the period 2015–2016 on three wildfires that burned in ponderosa pine (Pinus ponderosa) forests in the period 2000–2007 across broad environmental and burn severity gradients. Using a non-parametric multiplicative regression model, we found that downed woody fuel load, duff depth, and fall precipitation best explained variation in seedling density, while the distance to nearest seed tree, a soil productivity index, duff depth, and spring precipitation as snow best explained seedling height growth. Overall, results highlight the importance of burn severity and post-fire climate in tree regeneration, although the primary factors influencing seedling density and height growth vary. Drier conditions and changes to precipitation seasonality have the potential to influence tree establishment, survival, and growth in post-fire environments, which could lead to significant impacts for long-term forest recovery.

Moisture and vegetation cover limit ponderosa pine regeneration in high-severity burn patches in the southwestern US

BackgroundFire regimes are shifting in ponderosa pine (Pinus ponderosa Lawson & C. Lawson)-dominated forests, raising concern regarding future vegetation patterns and forest resilience, particularly within high-severity burn patches. The southwestern US has recently experienced a marked increase in large fires that produce large, high-severity patch interiors, with few surviving trees. These areas could be more susceptible for forest loss and conversions to alternative vegetation types than areas closer to the forest edge with more available seed sources. To better understand forest recovery, we surveyed ponderosa pine regeneration within edge and core areas (>200 m from edge) of high-severity patches in ten fires that burned between 1996 to 2008 across Arizona and New Mexico, USA. Specifically, we compared regeneration density, height, and canopy cover in patch edge and core areas and used generalized linear models to investigate the abiotic and biotic factors that contribute to ponderosa pine seedling establishment and density.ResultsHigh-severity burn-patch edge and core plots were not significantly different in seedling density, height, or canopy cover across fires. Seedling establishment was more likely at higher-elevation mesic sites and less likely when Gambel oak (Quercus gambelii Nutt.) was more abundant. Seedling density was negatively impacted by shrub, grass, and Gambel oak cover.ConclusionsRegeneration density varied among fires but analysis of regeneration in aggregated edge and core plots showed that abundance of seed availability was not the sole factor that limited ponderosa pine regeneration, probably because of surviving tree refugia within high-severity burn patches. Furthermore, our findings emphasize that ponderosa pine regeneration in our study area was significantly impacted by xeric topographic environments and vegetation competition. Continued warm and dry conditions and increased wildfire activity may delay the natural recovery of ponderosa pine forests, underscoring the importance of restoration efforts in large, high-severity burn patches.

Soil properties and climate drive ponderosa pine seedling presence in the southwestern USA

A continual improvement of our understanding about factors that influence tree regeneration is needed to inform landscape-scale restoration efforts across the western USA. This is increasingly important because changing climate and disturbance regimes will influence where restoration efforts should be implemented across landscapes. In this study, we used Forest Inventory and Analysis (FIA) data to model ponderosa pine (Pinus ponderosa C. Lawson var. scopulorum Engelm.) seedling presence across FIA plots in Arizona and New Mexico, USA. Additionally, we analyzed archived soil samples from FIA plots for percent clay, silt, and sand; samples were collected from the upper 20 cm of the mineral soil. Soil particle size is correlated with soil parent material, which has been shown to influence ponderosa pine regeneration. We used structural equation modeling to determine the relative importance of direct and indirect factors affecting ponderosa pine seedling presence. For the 0–10 cm depth increment of the mineral soil, soils with high inorganic C (carbonate) concentrations, a low percentage of coarse fragments (>2 mm) in the mineral soil, a low percentage of sand in the fine fractions (<2 mm) of the mineral soil, and low average May minimum temperature had the greatest probability of ponderosa pine seedling presence. Average annual precipitation also influenced pine regeneration through its negative correlation with inorganic C concentration. A partial explanation for the latter correlation could be that carbonates in the upper soil horizons leach into lower layers over time in areas with greater precipitation. Inorganic C concentration was also associated with soil parent material and was correlated with the soil cation exchange capacity and several macronutrient concentrations. Overall, our results indicate that soils and climate are key drivers of tree regeneration in the southwestern USA. Other factors such as the forest overstory, including the presence of seed trees, and the herbaceous plant community can also influence ponderosa pine regeneration. However, these factors were not statistically significant in the current study, likely because a small subset of FIA plots were used in the analyses. As soils are collected on additional FIA plots, new models of tree regeneration based on the current structural equation model could be evaluated and refined. Our results also indicate that information about the soil series associated with FIA plots would be useful for modeling tree regeneration, especially because soil series are commonly used by land managers in decision-making processes.

Persistent effects of fire severity on ponderosa pine regeneration niches and seedling growth

Several recent studies have documented how fire severity affects the density and spatial patterns of tree regeneration in western North American ponderosa pine forests. However, less is known about the effects of fire severity on fine-scale tree regeneration niche attributes such as understory plant composition and cover, surface fuel abundance, and soil properties, or how these attributes in turn affect regenerating ponderosa pine growth. Using 1-m2 plots centered on 360 ponderosa pine seedlings that regenerated naturally after the Pumpkin Fire in 2000 in Arizona, we quantified regeneration niche attributes 13 years post-fire and measured their associations with seedling growth 11–16 years post-fire. Plots were established in a) unburned, b) moderate-severity, and two types of high-severity (100% tree mortality) burns, either c) adjacent to residual live forest edges (high-severity edge plots), or d) >200 m from any residual live trees (high-severity interior plots). We found that all burned plots had greater understory plant species richness, percent cover of forbs, exotic plants and coarse wood, as well as higher soil pH, sand and gravel content, and lower soil clay content compared to unburned plots. High-severity burn plots had the greatest total understory plant and shrub cover, the most herbaceous fine fuel biomass, and tended to have the highest soil nitrogen content compared to other burn severity classes. Ponderosa pine growth (i.e., stem diameter at root collar (DRC) and length of terminal leader) was lowest in the unburned compared to burned plots, and ponderosa pine terminal leader growth was consistently greater in the high-severity edge plots compared to other severities. Finally, niche characteristics such as overstory tree canopy cover (−), soil phosphate (+), and cover of coarse wood (+), forbs (+), and the native grasses, mountain muhly (+) and squirreltail (−), were important explanatory variables of ponderosa pine growth. Exotic plant cover did not have a negative association with ponderosa pine growth. These results suggest that if ponderosa pine seeds can disperse and germinate, and if seedlings can survive the first few critical years after germination, then low overstory canopy cover and abundant forbs or coarse wood may be associated with increased growth rates. Alternatively, forbs may be responding to the same site benefits as the seedling; and abundant forbs, coarse wood, and fine fuels might also put seedlings at increased risk of mortality from subsequent fire, at least until they are taller and more fire resistant.

Stand density, drought, and herbivory constrain ponderosa pine regeneration pulse

Trees in dry forests often regenerate in episodic pulses when wet periods coincide with ample seed production. Factors leading to success or failure of regeneration pulses are poorly understood. We investigated the impacts of stand thinning on survival and growth of the 2013 cohort of ponderosa pine (Pinus ponderosa Douglas ex P. Lawson &amp; C. Lawson) seedlings in northern Arizona, United States. We measured seedling survival and growth over the first five growing seasons after germination in six stand basal areas (BAs; 0, 7, 14, 23, 34, and 66 (unthinned) m2·ha−1) produced by long-term experimental thinnings. Five-year survival averaged 2.5% and varied among BAs. Mean survival duration was longer in intermediate BAs (11 to 16 months) than in clearings and high BAs (5 months). The BAs of 7, 14, and 23 m2·ha−1had &gt;2600 5-year-old seedlings·ha−1. In contrast, regeneration was lower in the clearing (666 seedlings·ha−1) and failed completely in the 34 m2·ha−1and unthinned treatments. Seedling survival was highest during wet years and lowest during drought years. Many surviving seedlings had no net height growth between years 4 and 5 because of stem browsing. Results indicate that natural regeneration of ponderosa pine is influenced by stand BA, drought, herbivory, and interactions between extreme climatic events.

Regeneration patterns reveal contraction of ponderosa forests and little upward migration of pinyon-juniper woodlands

A recent severe drought caused widespread mortality of ponderosa pine (Pinus ponderosa) and pinyon pine (Pinus edulis) in forests and woodlands in the southwestern United States. The sustainability of these tree species in the region depends on adequate regeneration and perhaps movement to more climatologically favorable locations. We investigated tree regeneration and species migration along elevation gradients in three community types (pinyon-juniper woodlands, woodland-forest ecotones, and ponderosa pine forests) and three soil parent materials (sedimentary, flow basalt, and volcanic cinder) at 27 sites in northern Arizona. We measured stand characteristics, historic cone production, and density of tree regeneration and small trees. All species produced cones at sites with cinder soil parent material but had little regeneration. Woodlands and woodland-forest ecotones had regeneration of pinyon pine and juniper despite earlier mortality of some mature pinyon trees. Regeneration of ponderosa pine was nearly absent in woodland-forest ecotones, suggesting eventual loss of ponderosa pine and transition to pinyon-juniper woodland. Forests had little regeneration and few small trees of any tree species, indicating little upward migration of pinyon pine and juniper. Occurrence of pinyon pine and juniper regeneration was associated with a suite of biotic and abiotic factors, including a negative relationship with temperature and a positive relationship with precipitation. Regeneration failure of ponderosa pine at woodland-forest ecotone and low-elevation forest sites coupled with limited upward migration of pinyon pine and juniper portends future losses of tree cover as climate continues to warm.

Large, high-severity burn patches limit fungal recovery 13 years after wildfire in a ponderosa pine forest

Over the past three decades, wildfires in southwestern US ponderosa pine (Pinus ponderosa Lawson & C. Lawson) forests have increased in size and severity. These wildfires can remove large, contiguous patches of mature forests, alter dominant plant communities and increase woody debris, potentially altering fungal community composition. Additionally, post-fire conditions may shift dominant fungal functional groups from plant-symbiotic ectomycorrhizal (EM) fungi to more decomposer saprotrophic fungi. We investigated the long-term (13 years post-wildfire) effect of fire severity on 1) fungal sporocarp density, functional groups and community composition and 2) EM colonization and community composition from naturally regenerating ponderosa pine seedlings on the Pumpkin Fire that burned in 2000 in Arizona, USA. Plots were located in four burn severity classes: unburned, moderate-severity, and two high-severity (defined as 100% tree mortality) classes, either adjacent to residual live forest edges (edge plots), or >200 m from any residual live trees (interior plots). We found that high-severity burn plots had a unique sporocarp community composition, and a shift in dominant sporocarp functional groups, with 5–13 times lower EM sporocarp densities, and 4–7 times lower EM sporocarp species richness compared to unburned and moderate-severity plots. In contrast, saprotrophic sporocarp densities and richness were similar among burn severity classes, even with the large amount of coarse wood in the high-severity burn patches. Regenerating ponderosa pine seedlings had similar EM colonization and richness among severity classes, yet high-severity interior plots had a different community composition and a lower relative abundance of EM species compared to moderate-severity burn plots. Taken together, our results suggest that large patches of high-severity fire have long-term consequences for both EM sporocarp and root tip communities. Because EM fungal species vary in function, the limited species pool available in interior high-severity burn patches may influence pine recovery.

Historical and current fire regimes in ponderosa pine forests at Zion National Park, Utah: Restoration of pattern and process after a century of fire exclusion

Plant community resilience in frequent-fire forests of western North America has been compromised by over a century of fire exclusion. However, there are several western National Parks where fires have been reintroduced over the past several decades, including Zion National Park in southwest Utah. Here we reconstruct historical components of fire regimes in ponderosa pine dominated forests at Zion and compare these to recent fire frequency, current plant community and fuels structure, and potential fire behavior. Historical fires burned every 9–10 years on average up until 1879, when fires ceased contemporaneous with introduction of Euro-American livestock grazing and timber harvest in upland forests. Abundant tree regeneration occurred after fire exclusion, with tree density averaging 45 trees ha−1 in reconstructed 1880 forests versus 106 trees ha−1 today. Intervals between recent (since 1988) wildfires and prescribed fires in these same stands ranged from 7 to 13 years, similar to historical fire timing. Depending on whether plots had burned from zero to three times in recent fires, we found significant differences in canopy base heights (increased), duff and litter depths (decreased), and percent cover of grass and forbs (increased), but not tree density, tree basal area, shrub height, shrub cover, or woody fuels. Combined effects of recent fires on overstory and understory structure resulted in a significant difference in likelihood of crown fire occurrence, declining from a mean of 58% in plots with no fire since 1879 to 13% in plots with three fires since 1988. Significant effects were generally seen after two or three fires, suggesting it is the reintroduction of the fire regime and not just individual fire events that restore resiliency. Overall, effects of recent fires are building on the latent resiliency of ponderosa pine forests at Zion National Park, although questions remain about extent and future dynamics of oak and manzanita shrubfields that occupy similar environmental settings, along with a general lack of ponderosa pine regeneration across all plots.

Ponderosa Pine Regeneration, Wildland Fuels Management, and Habitat Conservation: Identifying Trade-Offs Following Wildfire

Increasing wildfires in western North American conifer forests have led to debates surrounding the application of post-fire management practices. There is a lack of consensus on whether (and to what extent) post-fire management assists or hinders managers in achieving goals, particularly in under-studied regions like eastern ponderosa pine forests. This makes it difficult for forest managers to balance among competing interests. We contrast structural and community characteristics across unburned ponderosa pine forest, severely burned ponderosa pine forest, and severely burned ponderosa pine forest treated with post-fire management with respect to three management objectives: ponderosa pine regeneration, wildland fuels control, and habitat conservation. Ponderosa pine saplings were more abundant in treated burned sites than untreated burned sites, suggesting increases in tree regeneration following tree planting; however, natural regeneration was evident in both unburned and untreated burned sites. Wildland fuels management greatly reduced snags and coarse woody debris in treated burned sites. Understory cover measurements revealed bare ground and fine woody debris were more strongly associated with untreated burned sites, and greater levels of forbs and grass were more strongly associated with treated burned sites. Wildlife habitat was greatly reduced following post-fire treatments. There were no tree cavities in treated burned sites, whereas untreated burned sites had an average of 27 ± 7.68 cavities per hectare. Correspondingly, we found almost double the avian species richness in untreated burned sites compared to treated burned sites (22 species versus 12 species). Unburned forests and untreated burned areas had the same species richness, but hosted unique avian communities. Our results indicate conflicting outcomes with respect to management objectives, most evident in the clear costs to habitat conservation following post-fire management application.

Some Lessons Learned on Early Survival and Growth of Containerized, Locally-Sourced Ponderosa Pine Seedlings in the Davis Mountains of Western Texas, US

The ponderosa pine forests in the Davis Mountains of western Texas recently experienced a major mortality event caused, in part, by an extended regional drought that predisposed trees and stands to mortality from both western pine beetle and wildfires. The loss of many overstory pines and the scarcity of natural ponderosa pine regeneration pose a considerable challenge to restoration. A commissioned study investigated artificial regeneration using containerized ponderosa pine seedlings with multiple planting seasons and vegetation management alternatives. Early survival was statistically greater for dormant season plantings than monsoon season plantings. Vegetation management treatments influenced early growth, survival, and herbivory rates. Physical weed control, which consisted of fibrous weed mats around the base of planted seedlings, showed early advantages over some vegetation management treatments in growth, survival and herbivory deterrence, but all vegetation management treatments had similar survival and herbivory results after 2.5 years. Early survival was poor in all treatments, mainly due to herbivory, which was identified as the principal short-term obstacle to artificial regeneration of ponderosa pine in the Davis Mountains. The larger question regarding feasibility of recovery in this isolated population, particularly if local climatic conditions become increasingly unfavorable, remains.

Post-fire management affects species composition but not Douglas-fir regeneration in the Klamath Mountains

Ensuring adequate conifer regeneration after high severity wildfires is a common objective for ecologists and forest managers. In the Klamath region of Oregon and California, a global hotspot of botanical biodiversity, concerns over regeneration have led to post-fire management on many sites, which involves salvage logging followed by site preparation, conifer planting, and manual shrub release. To quantify the impacts of post-fire management, we sampled 62 field sites that burned at high severity nearly 20 years ago in the Klamath-Siskiyou Mountain bioregion, stratifying by management and aspect. We measured cover of shrubs and trees and density and frequency of trees and used Nonmetric Multidimensional Scaling to compare community composition, plant community assemblage based on regenerative traits, and density of tree species between aspect and management. On average, shrub cover exceeded the cover of conifers, hardwoods or grasses, regardless of management history or aspect. The average number of species was lower and resprouting species were less abundant on south aspect sites; seed banking species were most abundant on north aspects. Post-fire management was associated with greater cover of seed banking and nitrogen-fixing species but it did not affect diversity. Management had no impact on Douglas-fir regeneration, the main species of concern in the region. Regeneration of ponderosa pine was higher on sites with post-fire management, but only on south slopes. The frequency of Douglas-fir was associated with aspect, while the frequency of ponderosa pine was associated with management. Overall, our study demonstrates the important role that aspect plays in determining the effectiveness of management after high severity wildfires. Indeed, the effect of aspect on site conditions often overwhelmed the ability of management to influence community composition (including different regenerative strategies), species diversity, and regeneration. Managed sites differed from unmanaged sites in several diverse ways with varied implications for longer-term forest development. Managed sites had taller dominant conifers, which suggests that post-fire management could hasten the period needed to achieve fire resistance. Managed sites were similar in plant community composition but had more homogeneous structure--e.g., managed sites had fewer snags, which are an important habitat feature for several bird species in the region. Finally, management was not associated with conifer regeneration success on north slopes, suggesting that interventions may not be needed uniformly across post-fire landscape. New policies of ecosystem-based management in the Klamath region should consider the important interactions between aspect and post-fire management, and tailor management practices based on specific objectives and landscape context.

Limits to Ponderosa Pine Regeneration following Large High-Severity Forest Fires in the United States Southwest

High-severity fires in dry conifer forests of the United States Southwest have created large (>1000 ha) treeless areas that are unprecedented in the regional historical record. These fires have reset extensive portions of Southwestern ponderosa pine (Pinus ponderosa Lawson & C. Lawson var. scopulorum Engelm.) forest landscapes. At least two recovery options following high-severity fire are emerging. One option is for post-fire successional pathways to move toward a return to the pre-fire forest type. Alternatively, an area may transition to persistent non-forested ecosystems. We studied regeneration patterns of ponderosa pine following eight fires in Arizona and New Mexico, USA, that burned in dry conifer forests dominated by ponderosa pine during a recent 18-year regional drought period, 1996 to 2013. Our a priori hypotheses were: 1) the most xeric areas within these severely burned dry conifer forests are least likely to regenerate to the pre-fire forest type due to persistent post-fire moisture stress; and 2) areas farther away from conifer seed sources have a lower likelihood of regeneration, even if these areas are climatically favorable for post-fire ponderosa pine establishment. We evaluated our hypotheses using empirical data and generalized linear mixed-effects models. We found that low-elevation, xeric sites are more limiting to conifer regeneration than higher-elevation mesic sites. Areas >150 m from a seed source are much less likely to have ponderosa pine regeneration. Spatial interpolations of modeled post-fire regeneration of ponderosa pine across the study landscapes indicate expansive areas with low likelihood of pine regeneration following high-severity fire. We discuss multiple post-fire successional pathways following high-severity fire, including potentially stable transitions to non-forest vegetation types that may represent long-term type conversions. These findings regarding landscape changes in Southwest forests in response to fires and post-fire regeneration patterns during early-stage climate warming contribute to the development of better-informed ecosystem management strategies for forest adaptation or mitigation under projected hotter droughts in this region.

Stand composition and aspect are related to conifer regeneration densities following hazardous fuels treatments in Colorado, USA

Abstract Hazardous fuels reduction treatments often reduce the likelihood of undesirable fire behavior (e.g. crown fire) through some combination of tree thinning and surface fuel reduction. However, fuels treatment effectiveness declines over time as fuels re-accumulate and new tree cohorts of trees establish and grow. Fuels treatment longevity therefore depends in part on post treatment tree regeneration rates, which vary depending on characteristics of sites and treatments as well as on stand composition. We used a chronosequence of 3–12 year old fuels treatments to investigate the impact of site (“wet” north aspects versus “dry” south aspects), treatment type (thinning with and without follow up slash burning), and stand composition on conifer regeneration following treatments in stands of ponderosa pine (Pinus ponderosa Lawson & C. Lawson) and Douglas-fir (Pseudotsuga menziesii [Mirb.] Franco var. glauca [Beissn.] Franco) in Colorado, USA. We found conifer regeneration density was >500 trees ha−1 on average by 10 years post treatment. While stands on north aspects had twice the regeneration density of those on south aspects, the rate of increase since treatment was similar. Instead, there was significantly more regeneration on north aspects because these stands contained a considerable amount of Douglas-fir, which was largely absent on south aspects. Ponderosa pine regeneration density increased with time since treatment, while Douglas-fir regeneration density did not, which suggests that treatments promoted reproduction of ponderosa pine. Conversely, Douglas-fir regeneration density was related to the amount of Douglas-fir basal area in stands, which was not the case for ponderosa pine, probably because all stands had ample amounts of ponderosa pine basal area. There was no difference in regeneration density between burned and unburned stands. Our results highlight the influence of stand composition and site characteristics on tree regeneration following fuels treatments. In this study, fuels treatments acted much like shelterwood regeneration treatments for ponderosa pine, pairing favorable establishment environments for this species with a seed source. This resulted in ponderosa pine trees establishing even on relatively dry south aspects, while Douglas-fir regeneration may have been inhibited by seed availability and was largely confined to wetter north aspects. Despite similar tree regeneration rates across aspects, fuels treatment effects may nonetheless deteriorate more quickly on north aspects because of this Douglas-fir regeneration, much of which predated treatments. Indeed, advance regeneration was common in all of our stands, suggesting the longevity of fuels treatment effects will depend substantially on whether these trees are eventually able to release.

Spatial patterns of ponderosa pine regeneration in high-severity burn patches

Abstract Contemporary wildfires in southwestern US ponderosa pine forests can leave uncharacteristically large patches of tree mortality, raising concerns about the lack of seed-producing trees, which can prevent or significantly delay ponderosa pine regeneration. We established 4-ha plots in high-severity burn patches in two Arizona wildfires, the 2000 Pumpkin and 2002 Rodeo-Chediski Fires, to determine if: (1) distance from forest edge influences the density and spatial patterns of regenerating ponderosa pine and sprouting tree species, (2) interactions with re-sprouting trees affect spatial patterns of ponderosa pine regeneration, and (3) distance from forest edge and species competition affect regenerating ponderosa pine height. Plots were located in high-severity burn patches (defined as 100% tree mortality) and either adjacent to residual live forest edges (edge plots), or >200 m from any residual live trees (interior plots). We found higher ponderosa pine regeneration densities in the edge plots (13–154 (median = 69) stems ha−1) than the interior plots (12–124 (median = 29) stems ha−1) on both wildfires, but no differences in spatial patterns between edge and interior plots. Ponderosa pine regeneration displayed patterns of small-scale spatial aggregation in all plots, except one edge and one interior plot on the Pumpkin Fire, which displayed random distributions. These patterns suggest both short- and long-distance dispersal play important roles in ponderosa pine regeneration in high-severity burn patches. Sprouting trees dominated tree regeneration on the Rodeo-Chediski Fire, but they were spatially independent of ponderosa pine and did not influence ponderosa pine height. Regenerating ponderosa pine height was positively correlated with neighboring ponderosa pine densities and height, suggesting that intraspecific facilitation or similar habitat preferences occur in high-severity burn patches. Collectively, these results indicate that ponderosa pines are re-establishing with heterogeneous spatial patterns in large high-severity burn patches, but often with low densities. Also, ponderosa pine regeneration could be more strongly influenced by intraspecific facilitation than interspecific competition from dense sprouting species. Future forest spatial patterns and composition are still unclear, but at this stage of development, these heterogeneous patches, characterized by drought-tolerant sprouting species or low pine densities, could be more resilient to climate change and severe wildfires than the overly-dense ponderosa pine forests that were present before the wildfires.

Long-term thinning alters ponderosa pine reproduction in northern Arizona

The future of ponderosa pine (Pinus ponderosa var. scopulorum) forests in the southwestern United States is uncertain because climate-change-induced stresses are expected to increase tree mortality and place greater constraints on regeneration. Silvicultural treatments, which include thinning, are increasingly being used to address forest health concerns by restoring ponderosa pine forests to more open conditions representative of historical forest structure. In light of the greater use of thinning and mounting concerns about the future of the species at the southern edge of its range, further investigations about impacts of thinning on ponderosa pine regeneration and underlying mechanisms are needed. We used a long-term (>50years) experiment in northern Arizona to investigate impacts of repeated stand thinning that maintained different growing stock basal areas (0, 7, 14, 23, 34, 66m2ha−1) on early seedling survival, growth, and microenvironment. Seedling survival for the first two years after germination (2013–2015), which had above-average precipitation, was higher than reported in several earlier studies and ranged between 4 and 21% among all basal areas. Seedling density exhibited a negative quadratic relationship with basal area and was positively associated with litter cover. Growing stock levels that fostered the highest seedling survival and density were those with a low density of overstory trees, low canopy cover, high cone production, coverage of soil by a thin layer of litter, and high soil water content at a depth of 15–30cm. Overstory basal area was positively associated with seedling height but negatively associated with seedling diameter. During this relatively wet period, all basal area treatments supported higher average seedling densities than those previously recommended to produce a multi-aged stand or presettlement structure in the southwestern United States. Our results show that long-term maintenance of low to intermediate basal areas (7–23m2ha−1) by thinning over the last 50years led to a favorable microenvironment for early seedling establishment of ponderosa pine.

A field experiment informs expected patterns of conifer regeneration after disturbance under changing climate conditions

Climate change may inhibit tree regeneration following disturbances such as wildfire, altering post-disturbance vegetation trajectories. We implemented a field experiment to examine the effects of manipulations of temperature and water on ponderosa pine (Pinus ponderosa Douglas ex P. Lawson &amp; C. Lawson) and Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco) seedlings planted in a low-elevation, recently disturbed setting of the Colorado Front Range. We implemented four treatments: warmed only (Wm), watered only (Wt), warmed and watered (WmWt), and control (Co). We found that measures of growth and survival varied significantly by treatment type. Average growth and survival was highest in the Wt plots, followed by the Co, WmWt, and Wm plots, respectively. This general trend was observed for both conifer species, although average growth and survival was generally higher in ponderosa pine than in Douglas-fir. Our findings suggest that warming temperatures and associated drought are likely to inhibit post-disturbance regeneration of ponderosa pine and Douglas-fir in low-elevation forests of the Colorado Front Range and that future vegetation composition and structure may differ notably from historic patterns in some areas. Our findings are relevant to other forested ecosystems in which a warming climate may similarly inhibit regeneration by dominant tree species.

Post-fire ponderosa pine regeneration with and without planting in Arizona and New Mexico

Forest fires are increasing in size and severity globally, yet the roles of natural and artificial regeneration in promoting forest recovery are poorly understood. Post-fire regeneration of ponderosa pine (Pinus ponderosa, Lawson and C. Lawson) in the southwestern U.S. is slow, episodic, and difficult to predict. Planting of ponderosa pine after wildfire may accelerate reforestation, but little is known about survival of plantings and the amount of post-fire natural regeneration. We compared ponderosa pine regeneration between paired planted and unplanted plots at eight sites in Arizona and New Mexico that recently (2002–2005) burned severely. Two sites had no natural regeneration and no survival of planted seedlings. Seedling presence increased with number of years since burning across all plots, was positively associated with forb and litter cover on planted plots, and was positively associated with litter cover on unplanted plots. Survival of planted seedlings, measured five to eight years after planting, averaged 25% (SE=8) and varied from 0% to 70% across sites resulting in seedling densities of 0–521 trees ha−1. Based on a projected 44% survival of seedlings to mature trees and target density of mature trees determined by historical range of variability and ecological restoration principles, four of eight sites have a seedling density in planted plots (125–240ha−1) that will produce a density of mature trees (55–106ha−1) close to desired levels, whereas seedlings are currently deficient at three planted sites, and in surplus at one site, which had abundant natural regeneration. Natural regeneration in unplanted plots during the first decade after burning produced seedling densities inconsistent with desired numbers of mature trees. Natural regeneration in unplanted plots produced less than 33 seedlings ha−1 at seven of eight sites, but produced 1433 seedlings ha−1 at one high-elevation site that supported a more mesic vegetation community before burning than the other sites. Our results show that current practices for planting ponderosa pine after severe fires in Arizona and New Mexico produce desired numbers of seedlings in approximately half of all projects, whereas natural regeneration rarely does within the first decade after burning.

Effects of heat and drought on carbon and water dynamics in a regenerating semi-arid pine forest: a combined experimental and modeling approach

Abstract. Predicting the net effects on the carbon and water balance of semi-arid forests under future conditions depends on ecosystem processes responding to changes in soil and atmospheric drought. Here we apply a combination of field observations and soil–plant–atmosphere modeling (SPA) to study carbon and water dynamics in a regenerating ponderosa pine forest. The effects of soil and atmospheric drought were quantified based on a field irrigation experiment combined with model simulations. To assess future effects of intensifying drought on ecosystem processes, the SPA model was run using temperature and precipitation scenarios for 2040 and 2080. Experimentally increased summer water availability clearly affected tree hydraulics and enhanced C uptake in both the observations and the model. Simulation results showed that irrigation was sufficient to eliminate soil water limitation and maintaining transpiration rates, but gross primary productivity (GPP) continued to decrease. Observations of stomatal conductance indicated a dominant role of vapor pressure deficit (VPD) in limiting C uptake. This was confirmed by running the simulation under reduced atmospheric drought (VPD of 1 kPa), which largely maintained GPP rates at pre-drought conditions. The importance of VPD as a dominant driver was underlined by simulations of extreme summer conditions. We found GPP to be affected more by summer temperatures and VPD as predicted for 2080 (−17%) than by reductions in summer precipitation (−9%). Because heterotrophic respiration responded less to heat (−1%) than to reductions in precipitation (−10%), net ecosystem C uptake declined strongest under hotter (−38%) compared to drier summer conditions (−8%). Considering warming trends across all seasons (September–May: +3 °C and June–August: +4.5 °C), the negative drought effects were largely compensated by an earlier initiation of favorable growing conditions and bud break, enhancing early season GPP and needle biomass. An adverse effect, triggered by changes in early season allocation patterns, was the decline of wood and root biomass. This imbalance may increase water stress over the long term to a threshold at which ponderosa pine may not survive, and highlights the need for an integrated process understanding of the combined effects of trends and extremes.

Factors Influencing Height-Age Relationships and Recruitment of Ponderosa Pine Regeneration in Northern Arizona

Factors Influencing Height-Age Relationships and Recruitment of Ponderosa Pine Regeneration in Northern Arizona