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.     

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.     

Does It Make Sense to Restore Wildland Fire in Changing Climate?

Forest restoration guided by historical reference conditions of fire regime, forest structure, and composition has been increasingly and successfully applied in fire‐adapted forests of western North America. But because climate change is expected to alter vegetation distributions and foster severe disturbances, does it make sense to restore the ecological role of wildland fire through management burning and related activities such as tree thinning? I suggest that some site‐ and date‐specific historical conditions may be less relevant, but reference conditions in the broad sense are still useful. Reference conditions encompass not only the recent past but also evolutionary history, reflecting the role of fire as a selective force over millennia. Taking a long‐term functional view of historical reference conditions as the result of evolutionary processes can provide insights into past forest adaptations and migrations under various climates. As future climates change, historical reference data from lower, southerly, and drier sites may be useful in places that are higher, northerly, and currently wetter. Almost all models suggest that the future will have substantial increases in wildfire occurrence, but prior to recent human‐caused fire exclusion, fire‐adapted pine forests of western North America were among the most frequently burned in the world. Restoration of patterns of burning and fuels/forest structure that reasonably emulate historical conditions prior to fire exclusion is consistent with reducing the susceptibility of these ecosystems to catastrophic loss. Priorities may include fire and thinning treatments of upper elevation ecotones to facilitate forest migration, whereas vulnerable low‐elevation forests may merit less management investment.     

Modelling Variable Fire Severity in Boreal Forests: Effects of Fire Intensity and Stand Structure

It is becoming clear that fires in boreal forests are not uniformly stand-replacing. On the contrary, marked variation in fire severity, measured as tree mortality, has been found both within and among individual fires. It is important to understand the conditions under which this variation can arise. We integrated forest sample plot data, tree allometries and historical forest fire records within a diameter class-structured model of 1.0 ha patches of mono-specific black spruce and jack pine stands in northern Québec, Canada. The model accounts for crown fire initiation and vertical spread into the canopy. It uses empirical relations between fire intensity, scorch height, the percent of crown scorched and tree mortality to simulate fire severity, specifically the percent reduction in patch basal area due to fire-caused mortality. A random forest and a regression tree analysis of a large random sample of simulated fires were used to test for an effect of fireline intensity, stand structure, species composition and pyrogeographic regions on resultant severity. Severity increased with intensity and was lower for jack pine stands. The proportion of simulated fires that burned at high severity (e.g. >75% reduction in patch basal area) was 0.80 for black spruce and 0.11 for jack pine. We identified thresholds in intensity below which there was a marked sensitivity of simulated fire severity to stand structure, and to interactions between intensity and structure. We found no evidence for a residual effect of pyrogeographic region on simulated severity, after the effects of stand structure and species composition were accounted for. The model presented here was able to produce variation in fire severity under a range of fire intensity conditions. This suggests that variation in stand structure is one of the factors causing the observed variation in boreal fire severity.     

Impacts of climate variability on tree demography in second growth tropical forests: the importance of regional context for predicting successional trajectories

Naturally regenerating and restored second growth forests account for over 70% of tropical forest cover and provide key ecosystem services. Understanding climate change impacts on successional trajectories of these ecosystems is critical for developing effective large‐scale forest landscape restoration (FLR) programs. Differences in environmental conditions, species composition, dynamics, and landscape context from old growth forests may exacerbate climate impacts on second growth stands. We compile data from 112 studies on the effects of natural climate variability, including warming, droughts, fires, and cyclonic storms, on demography and dynamics of second growth forest trees and identify variation in forest responses across biomes, regions, and landscapes. Across studies, drought decreases tree growth, survival, and recruitment, particularly during early succession, but the effects of temperature remain unexplored. Shifts in the frequency and severity of disturbance alter successional trajectories and increase the extent of second growth forests. Vulnerability to climate extremes is generally inversely related to long‐term exposure, which varies with historical climate and biogeography. The majority of studies, however, have been conducted in the Neotropics hindering generalization. Effects of fire and cyclonic storms often lead to positive feedbacks, increasing vulnerability to climate extremes and subsequent disturbance. Fragmentation increases forests’ vulnerability to fires, wind, and drought, while land use and other human activities influence the frequency and intensity of fire, potentially retarding succession. Comparative studies of climate effects on tropical forest succession across biogeographic regions are required to forecast the response of tropical forest landscapes to future climates and to implement effective FLR policies and programs in these landscapes.     

Effects of overstory and understory vegetation on the understory light environment in mixed boreal forests

Abstract. The percentage of above-canopy Photosynthetic Photon Flux Density (%PPFD) was measured at 0, 50 and 100 cm above the forest floor and above the main understory vegetation in stands of (1) pure Betula papyrifera (White birch), (2) pure Populus tremuloides (Trembling aspen), (3) mixed broad-leaf-conifer, (4) shade-tolerant conifer and (5) pure Pinus banksiana (Jack pine) occurring on both clay and till soil types. %PPFD was measured instantaneously under overcast sky conditions (nine locations within each of 29 stands) and continuously for a full day under clear sky conditions (five locations within each of eight stands). The percentage cover of the understory layer was estimated at the same locations as light measurements. Mean %PPFD varied from 2% at the forest floor under Populus forests to 15% above the understory vegetation cover under Betula forests. Percent PPFD above the understory vegetation cover was significantly higher under shade intolerant tree species such as Populus, Betula and Pinus than under shade tolerant conifers. No significant differences were found in %PPFD above the understory vegetation cover under similar tree species between clay and till soil types. The coefficient of variation in %PPFD measured in the nine locations within each stand was significantly lower under deciduous dominated forests (mean of 19%) than under coniferous dominated forests (mean of 40%). %PPFD measured at the forest floor was positively correlated with %PPFD measured above the understory vegetation and negatively correlated with cumulative total percent cover of the understory vegetation (R² = 0.852). The proportion of sunflecks above 250 and 500 mmol m^–2 s^–1 was much lower and %PPFD in shade much higher under Populus and Betula forests than under the other forests. Differences in the mean, variability and nature of the light environment found among forest and soil types are discussed in relation to their possible influences on tree succession.     

Fire characteristics and environmental conditions shape plant communities via regeneration strategy

Climate change is altering disturbance regimes outside historical norms, which can impact biodiversity by selecting for plants with particular traits. The relative impact of disturbance characteristics on plant traits and community structure may be mediated by environmental gradients. We aimed to understand how wildfire impacted understory plant communities and plant regeneration strategies along gradients of environmental conditions and wildfire characteristics in boreal forests. We established 207 plots (60 m²) in recently burned stands and 133 plots in mature stands with no recent fire history in comparable gradients of stand type, site moisture (drainage) and soil organic layer (SOL) depth in two ecozones in Canada's Northwest Territories. At each plot, we recorded all vascular plant taxa in the understory and measured the regeneration strategy (seeder, resprouter, survivor) in burned plots, along with seedbed conditions (mineral soil and bryophyte cover). Dispersal, longevity and growth form traits were determined for each taxon. Fire characteristics measured included proportion of pre-fire SOL combusted (fire severity), date of burn (fire seasonality) and pre-fire stand age (time following fire). Results showed understory community composition was altered by fire. However, burned and mature stands had similar plant communities in wet sites with deep SOL. In the burned plots, regeneration strategies were determined by fire severity, drainage and pre- and post-fire SOL depth. Resprouters were more common in wet sites with deeper SOL and lower fire severity, while seeders were associated with drier sites with thinner SOL and greater fire severity. This led to drier burned stands being compositionally different from their mature counterparts and seedbed conditions were important. Our study highlights the importance of environment–wildfire interactions in shaping plant regeneration strategies and patterns of understory plant community structure across landscapes, and the overriding importance of SOL depth and site drainage in mediating fire severity, plant regeneration and community structure.     

Stand- and plot-level changes in a boreal forest understory community following wildfire

Background: Boreal forest understory plant communities are known to be resilient to fire – the species composition of stands after a fire is quite similar to the pre-fire composition. However, we know little about recovery of individual plants within particular locations in forest stands (i.e. plot-level changes) since we usually do not have pre-fire data for plots.

Aims: We wanted to determine whether species recruited into the same or different locations in a Pinus banksiana stand that experienced a severe wildfire.

Methods: We used pre-existing permanent plots to evaluate the cover of understory after an unplanned wildfire.

Results: Across the entire stand nine of 47 species showed a significant change in cover. The largest change in a plant functional group was in the mosses, with all species present before fire being eliminated. There was no change in species diversity or total cover. At the plot level, species composition showed a much greater change. An average of 47% of the species present in a plot before the fire were absent in the same plot after the fire, and the total species turnover in plots was 88% of the species present before the fire. The plots showed a similar shift in species composition.

Conclusions: These results confirm that boreal forest communities show a high degree of resilience to fire, but within a forest stand species will be found in different locations following fire, potentially exposing them to a different set of biotic and abiotic conditions in these new locations.     

Changing growth response to wildfire in old‐growth ponderosa pine trees in montane forests of north central Idaho

North American fire‐adapted forests are experiencing changes in fire frequency and climate. These novel conditions may alter postwildfire responses of fire‐adapted trees that survive fires, a topic that has received little attention. Historical, frequent, low‐intensity wildfire in many fire‐adapted forests is generally thought to have a positive effect on the growth and vigor of trees that survive fires. Whether such positive effects can persist under current and future climate conditions is not known. Here, we evaluate long‐term responses to recurrent 20th‐century fires in ponderosa pine, a fire‐adapted tree species, in unlogged forests in north central Idaho. We also examine short‐term responses to individual 20th‐century fires and evaluate whether these responses have changed over time and whether potential variability relates to climate variables and time since last fire. Growth responses were assessed by comparing tree‐ring measurements from trees in stands burned repeatedly during the 20th century at roughly the historical fire frequency with trees in paired control stands that had not burned for at least 70 years. Contrary to expectations, only one site showed significant increases in long‐term growth responses in burned stands compared with control stands. Short‐term responses showed a trend of increasing negative effects of wildfire (reduced diameter growth in the burned stand compared with the control stand) in recent years that had drier winters and springs. There was no effect of time since the previous fire on growth responses to fire. The possible relationships of novel climate conditions with negative tree growth responses in trees that survive fire are discussed. A trend of negative growth responses to wildfire in old‐growth forests could have important ramifications for forest productivity and carbon balance under future climate scenarios.     

Effects of introduced ungulates on forest understory communities in northern Patagonia are modified by timing and severity of stand mortality

Natural disturbances such as fires, windstorms, floods, and herbivory often act on plant communities, affecting their structure and the abundance and composition of their species. Most research has focused on the effects of single disturbances on plant communities whereas the synergistic effects of several disturbances have received less attention. In this study, we evaluated how timing and severity of tree mortality modified plant use by introduced deer and early post-mortality successional trajectories in northern Patagonian conifer forests. We sampled understory composition and deer use in Austrocedrus chilensis (ciprés de la cordillera) forest stands undergoing varying timing and severity of forest mortality as reconstructed using dendroecological techniques. In addition, we evaluated the effect of fallen logs on plant composition and deer use of plants by monitoring areas of massive dieback where fallen logs had been removed for fire hazard reduction, and nearby control areas not subjected to such removal. Stepwise regression analyses showed that history and severity of tree mortality strongly influence plant composition and deer use of plants. For deer use (with pellet counts and browsing index as response variables), results showed a positive relationship with degree of stand mortality and a negative relationship with cover of fallen logs. Similarly, cover of unpalatable shrub species was explained by canopy mortality history, whereas cover of palatable shrub species was positively associated with severity of canopy mortality. In areas where fallen logs had been removed, pellet counts were six times higher than those in control areas. Though total shrub species cover was similar between log removal and control areas, proportion of unpalatable shrubs increased in areas where fallen logs had been removed. In conclusion, deer use of plants was strongly limited by tall fallen logs, allowing palatable species to establish and grow. Fallen log removal accelerated deer entrance and changed understory composition toward more browse-resistant and unpalatable species. These results underscore the importance of considering the dynamics (timing, severity, and extent) of fallen woody debris influencing understory herbivory and post-disturbance succession. In addition, experimental results underpin the importance of maintaining snags and large woody debris in disturbed landscapes where salvage logging is a routine procedure.