Changes in fire regime break the legacy lock on successional trajectories in Alaskan boreal forest

Predicting plant community responses to changing environmental conditions is a key element of forecasting and mitigating the effects of global change. Disturbance can play an important role in these dynamics, by initiating cycles of secondary succession and generating opportunities for communities of long‐lived organisms to reorganize in alternative configurations. This study used landscape‐scale variations in environmental conditions, stand structure, and disturbance from an extreme fire year in Alaska to examine how these factors affected successional trajectories in boreal forests dominated by black spruce. Because fire intervals in interior Alaska are typically too short to allow relay succession, the initial cohorts of seedlings that recruit after fire largely determine future canopy composition. Consequently, in a dynamically stable landscape, postfire tree seedling composition should resemble that of the prefire forest stands, with little net change in tree composition after fire. Seedling recruitment data from 90 burned stands indicated that postfire establishment of black spruce was strongly linked to environmental conditions and was highest at sites that were moist and had high densities of prefire spruce. Although deciduous broadleaf trees were absent from most prefire stands, deciduous trees recruited from seed at many sites and were most abundant at sites where the fires burned severely, consuming much of the surface organic layer. Comparison of pre‐ and postfire tree composition in the burned stands indicated that the expected trajectory of black spruce self‐replacement was typical only at moist sites that burned with low fire severity. At severely burned sites, deciduous trees dominated the postfire tree seedling community, suggesting these sites will follow alternative, deciduous‐dominated trajectories of succession. Increases in the severity of boreal fires with climate warming may catalyze shifts to an increasingly deciduous‐dominated landscape, substantially altering landscape dynamics and ecosystem services in this part of the boreal forest.     

Ten years of vegetation assembly after a North American mega fire

Altered fuels and climate change are transforming fire regimes in many of Earth's biomes. Postfire reassembly of vegetation – paramount to C storage and biodiversity conservation – frequently remains unpredictable and complicated by rapid global change. Using a unique data set of pre and long‐term postfire data, combined with long‐term data from nearby unburned areas, we examined 10 years of understory vegetation assembly after the 2002 Hayman Fire. This fire was the largest wildfire in recorded history in Colorado, USA. Resistance (initial postfire deviance from prefire condition) and resilience (return to prefire condition) declined with increasing fire severity. However, via both resistance and resilience, ‘legacy’ species of the prefire community constituted >75% of total plant cover within 3 years even in severely burned areas. Perseverance of legacy species, coupled with new colonizers, created a persistent increase in community species richness and cover over prefire levels. This was driven by a first‐year increase (maintained over time) in forbs with short life spans; a 2–3‐year delayed surge in long‐lived forbs; and a consistent increase in graminoids through the 10th postfire year. Burning increased exotic plant invasion relative to prefire and unburned areas, but burned communities always were >89% native. This study informs debate in the literature regarding whether these increasingly large fires are ‘ecological catastrophes.’ Landscape‐scale severe burning was catastrophic from a tree overstory perspective, but from an understory perspective, burning promoted rich and productive native understories, despite the entire 10‐year postfire period receiving below‐average precipitation.     

Modeling the effects of fire and climate change on carbon and nitrogen storage in lodgepole pine (Pinus contorta) stands

The interaction between disturbance and climate change and resultant effects on ecosystem carbon (C) and nitrogen (N) fluxes are poorly understood. Here, we model (using CENTURY version 4.5) how climate change may affect C and N fluxes among mature and regenerating lodgepole pine ( Pinus contorta var. latifolia Engelm. ex S. Wats.) stands that vary in postfire tree density following stand-replacing fire. Both young (postfire) and mature stands had elevated forest production and net N mineralization under future climate scenarios relative to current climate. Forest production increased 25% [Hadley (HAD)] to 36% [Canadian Climate Center (CCC)], compared with 2% under current climate, among stands that varied in stand age and postfire density. Net N mineralization increased under both climate scenarios, e.g., +19% to 37% (HAD) and +11% to 23% (CCC), with greatest increases for young stands with sparse tree regeneration. By 2100, total ecosystem carbon (live+dead+soils) in mature stands was higher than prefire levels, e.g., +16% to 19% (HAD) and +24% to 28% (CCC). For stands regenerating following fire in 1988, total C storage was 0–9% higher under the CCC climate model, but 5–6% lower under the HAD model and 20–37% lower under the Control. These patterns, which reflect variation in stand age, postfire tree density, and climate model, suggest that although there were strong positive responses of lodgepole pine productivity to future changes in climate, C flux over the next century will reflect complex relationships between climate, age structure, and disturbance-recovery patterns of the landscape.     

Resistance and resilience of stream insect communities to repeated hydrologic disturbances after a wildfire

1. Wildfires are often followed by severe, sediment‐laden floods in burned catchments. In this study, we documented resistance and resilience of stream insect communities to repeated postfire flash floods in a ‘burned stream’. We employed a before‐after‐control‐impact (BACI) design, where communities in comparable reaches of a burned stream and a reference stream were sampled from 2 years before, to 6 years after, a crown wildfire in north‐central New Mexico. 2. The first 100‐year flood following the 1996 Dome wildfire reduced total insect density and taxon richness to near zero in the burned stream. Despite showing low resistance, density returned rapidly to prefire levels because of colonisation by simuliids, chironomids and the mayfly Baetis tricaudatus. In general, taxa that were generalist feeders (collectors) with strong larval dispersal dominated communities in early postfire years with repeated, moderate flash floods. 3. Taxon richness and community composition were less resilient to postfire hydrologic disturbances. Taxon richness did not recover until floods dampened 4 years after the fire. Despite hydrologic recovery, composition in the burned stream still differed from prefire and reference stream compositions after 6 years postfire. A unique assemblage, dominated by taxa with strong larval or adult dispersal, was established after flash floods abated. Specialist feeders (shredders and grazers) that were common in prefire years were reduced or absent in the postfire assemblage. 4. Community succession in the burned stream was explained by the interaction between species traits, geographic barriers to colonisation and hydrologic conditions after the fire. Comparable changes in insect density, taxon richness, community composition and trait representation were not found in the reference stream, providing strong evidence that repeated postfire flash floods shaped community responses in the burned stream.     

Postfire Vegetation Recovery in Highland Pine Forests of the Dominican Republic

We surveyed postfire vegetation at five sites at high elevations (> 2000 m) in the Cordillera Central, Dominican Republic. Highlands of the Cordillera Central are dominated by a single pine species, Pinus occidentalis, but plant communities are rich with endemics and conservation and fire management efforts in these systems are ongoing. The burns were 2–7 yr in age and had consumed nearly all shrub crowns. Pines suffered high mortality (> 50%, all sites combined), but shrubs resprouted at high rates (88%, N = 957) after fire. All shrub taxa produced basal resprouts; eight of 11 shrub taxa measured had resprouting rates > 90 percent, while Baccharis myrsinites had the lowest (56%). Most taxa grew to prefire height quickly (within 5–7 yr), with regrowth of stem diameters lagging behind. Patterns and rates of shrub recovery resembled those documented in high elevation shrublands in Costa Rica and Brazil. Pinus occidentalis does not resprout, but larger individuals can survive fire. Survival increases dramatically when trees attain > 13-cm dbh, when bark becomes thick enough to protect cambial tissue. Overall, pines are regenerating much more slowly than shrubs, but seedling establishment varied considerably between sites. Frequent fires may cause a decline in pines and an increase in shrub- or grass-dominated communities. Succession in these high elevation fire-dependent pine forests favors taxa already present in the preburn vegetation, with woody composition changing little after fire, in contrast to lower-elevation cloud forest, where postfire vegetation has been shown to bear little resemblance to mature forest even after several years.     

Recovery of ground vegetation at the initial stage of fire succession

The impact of surface fires varying intensity on the living ground vegetation in the middle-taiga forests of Central Siberia are studied. It is revealed that fires, regardless of their intensity, decrease the percentage cover and the biomass of living ground vegetation; they also destroy the moss and lichen layer. The postfire recovery of the ground vegetation at the initial stage of postfire succession is determined by the prefire forest type, the fire intensity, and the burn depth of the litter.     

Succession of vegetation after a high-intensity fire in a pine forest with lichens

This paper presents the results of 20-year studies into the impact made by an experimental high-intensity fire on ecosystem components and postfire succession in a middle-taiga pine forest. About 44% of forest fuel loads burned down during the fire, and the emission of carbon was more than 18 t C/ha. As a result of the fire impact, trees died within 3 years after the fire, and this resulted in a significant accumulation of fuel loads. Twenty years after the fire, the biomass of forest fuel loads surpassed the prefire values 4 times, which led to the possibility of the origin of a repeated high-intensity fire. The initial stage of postfire succession in the pine forest is determined by forest vegetation conditions and takes place with the replacement of dominant grass and shrubs. The agrochemical and hydrothermal soil indicators were revealed to be changed after the fire, and this promoted improved conditions for the origin and development of natural regeneration sufficient for the formation of forest stand.     

Differences in bird communities in postfire silvicultural practices stands within pine forest of South Korea

We examined differences in bird communities in relation to characteristics of habitat structure in a pine forest, Samcheok, South Korea. An unburned stand, a stand burned 7 years earlier and then naturally restored, and a stand where Japanese red pine Pinus densiflora seedlings were planted after the fire were used for the survey. Habitat structure was dramatically changed by postfire silvicultural practices. Number of stand trees, shrubs, seedlings, snags, and vegetation coverage were significantly different among study stands. We made 1,421 detections of 46 bird species during 23 separate line transect surveys per stand between February 2007 and December 2008. The mean number of observed bird species and individuals, bird species diversity index (H′), and Simpson’s diversity index (D _s) were highest in the unburned stand and lowest in the pine seedling stand. There were more species and individuals of forest-dwelling birds in the unburned stand than both burned stands. Canopy and cavity nesters, foliage searchers, bark gleaners, and timber drillers were significantly higher in the unburned stand. In the pine seedling stand, densities of birds that prefer open field and shrub cover were higher. Stand structure was simplified in the pine seedling stand by postfire practices. Because of differences in habitat structure and bird communities, postfire practices in the burned stand should be re-evaluated. Also, management strategies for pine forest after forest fires are needed based on results of long-term experiments.     

Resistance to wildfire and early regeneration in natural broadleaved forest and pine plantation

The response of an ecosystem to disturbance reflects its stability, which is determined by two components: resistance and resilience. We addressed both components in a study of early post-fire response of natural broadleaved forest (Quercus robur, Ilex aquifolium) and pine plantation (Pinus pinaster, Pinus sylvestris) to a wildfire that burned over 6000 ha in NW Portugal. Fire resistance was assessed from fire severity, tree mortality and sapling persistence. Understory fire resistance was similar between forests: fire severity at the surface level was moderate to low, and sapling persistence was low. At the canopy level, fire severity was generally low in broadleaved forest but heterogeneous in pine forest, and mean tree mortality was significantly higher in pine forest. Forest resilience was assessed by the comparison of the understory composition, species diversity and seedling abundance in unburned and burned plots in each forest type. Unburned broadleaved communities were dominated by perennial herbs (e.g., Arrhenatherum elatius) and woody species (e.g., Hedera hibernica, Erica arborea), all able to regenerate vegetatively. Unburned pine communities presented a higher abundance of shrubs, and most dominant species relied on post-fire seeding, with some species also being able to regenerate vegetatively (e.g., Ulex minor, Daboecia cantabrica). There were no differences in diversity measures in broadleaved forest, but burned communities in pine forest shared less species and were less rich and diverse than unburned communities. Seedling abundance was similar in burned and unburned plots in both forests. The slower reestablishment of understory pine communities is probably explained by the slower recovery rate of dominant species. These findings are ecologically relevant: the higher resistance and resilience of native broadleaved forest implies a higher stability in the maintenance of forest processes and the delivery of ecosystem services.     

Spatial and temporal variation in historic fire regimes in subalpine forests across the Colorado Front Range in Rocky Mountain National Park, Colorado, USA

Aim The historical variability of fire regimes must be understood in the context of drivers of the occurrence of fire operating at a range of spatial scales from local site conditions to broad‐scale climatic variation. In the present study we examine fire history and variations in the fire regime at multiple spatial and temporal scales for subalpine forests of Engelmann spruce–subalpine fir (Picea engelmannii, Abies lasiocarpa) and lodgepole pine (Pinus contorta) of the southern Rocky Mountains. Location The study area is the subalpine zone of spruce–fir and lodgepole pine forests in the southern sector of Rocky Mountain National Park (ROMO), Colorado, USA, which straddles the continental divide of the northern Colorado Front Range (40°20′ N and 105°40′ W). Methods We used a combination of dendroecological and Geographic Information System methods to reconstruct fire history, including fire year, severity and extent at the forest patch level, for c. 30,000 ha of subalpine forest. We aggregated fire history information at appropriate spatial scales to test for drivers of the fire regime at local, meso, and regional scales. Results The fire histories covered c. 30,000 ha of forest and were based on a total of 676 partial cross‐sections of fire‐scarred trees and 6152 tree‐core age samples. The subalpine forest fire regime of ROMO is dominated by infrequent, extensive, stand‐replacing fire events, whereas surface fires affected only 1–3% of the forested area. Main conclusions Local‐scale influences on fire regimes are reflected by differences in the relative proportions of stands of different ages between the lodgepole pine and spruce–fir forest types. Lodgepole pine stands all originated following fires in the last 400 years; in contrast, large areas of spruce–fir forests consisted of stands not affected by fire in the past 400 years. Meso‐scale influences on fire regimes are reflected by fewer but larger fires on the west vs. east side of the continental divide. These differences appear to be explained by less frequent and severe drought on the west side, and by the spread of fires from lower‐elevation mixed‐conifer montane forests on the east side. Regional‐scale climatic variation is the primary driver of infrequent, large fire events, but its effects are modulated by local‐ and meso‐scale abiotic and biotic factors. The low incidence of fire during the period of fire‐suppression policy in the twentieth century is not unique in comparison with the previous 300 years of fire history. There is no evidence that fire suppression has resulted in either the fire regime or current forest conditions being outside their historic ranges of variability during the past 400 years. Furthermore, in the context of fuel treatments to reduce fire hazard, regardless of restoration goals, the association of extremely large and severe fires with infrequent and exceptional drought calls into question the future effectiveness of tree thinning to mitigate fire hazard in the subalpine zone.     

Rapid Increases in Forest Understory Diversity and Productivity following a Mountain Pine Beetle (Dendroctonus ponderosae) Outbreak in Pine Forests

The current unprecedented outbreak of mountain pine beetle (Dendroctonus ponderosae) in lodgepole pine (Pinus contorta) forests of western Canada has resulted in a landscape consisting of a mosaic of forest stands at different stages of mortality. Within forest stands, understory communities are the reservoir of the majority of plant species diversity and influence the composition of future forests in response to disturbance. Although changes to stand composition following beetle outbreaks are well documented, information on immediate responses of forest understory plant communities is limited. The objective of this study was to examine the effects of D. ponderosae-induced tree mortality on initial changes in diversity and productivity of understory plant communities. We established a total of 110 1-m² plots across eleven mature lodgepole pine forests to measure changes in understory diversity and productivity as a function of tree mortality and below ground resource availability across multiple years. Overall, understory community diversity and productivity increased across the gradient of increased tree mortality. Richness of herbaceous perennials increased with tree mortality as well as soil moisture and nutrient levels. In contrast, the diversity of woody perennials did not change across the gradient of tree mortality. Understory vegetation, namely herbaceous perennials, showed an immediate response to improved growing conditions caused by increases in tree mortality. How this increased pulse in understory richness and productivity affects future forest trajectories in a novel system is unknown.     

Aspen succession and nitrogen loading: a case for epiphytic lichens as bioindicators in the Rocky Mountains,USA

Question: Can lichen communities be used to assess short‐ and long‐term factors affecting seral quaking aspen (Populus tremuloides) communities at the landscape scale? Location: Bear River Range, within the Rocky Mountains, in northern Utah and southern Idaho, USA. Method: Forty‐seven randomly selected mid‐elevation aspen stands were sampled for lichens and stand conditions. Plots were characterized according to tree species cover, basal area, stand age, bole scarring, tree damage, and presence of lichen species. We also recorded ammonia emissions with passive sensors at 25 urban and agricultural sites throughout an adjacent populated valley upwind of the forest stands. Nonmetric multidimensional scaling (NMS) ordination was used to evaluate an array of 20 variables suspected to influence lichen communities. Results: In NMS, forest succession explained most variance in lichen composition and abundance, although atmospheric nitrogen from local agricultural and urban sources also significantly influenced the lichen communities. Abundance of nitrophilous lichen species decreased with distance from peak ammonia sources and the urban center in all aspen succession classes. One lichen, Phaeophyscia nigricans, was found to be an effective bioindicator of nitrogen loading. Conclusions: Lichen communities in this landscape assessment of aspen forests showed clear responses to long‐term (stand succession) and short‐term (nitrogen deposition) influences. At the same time, several environmental factors (e.g. tree damage and scarring, distance to valley, topography, and stand age) had little influence on these same lichen communities. We recommend further use of epiphytic lichens as bioindicators of dynamic forest conditions.     

The role of succession in the evolution of flammability

Fire-prone ecosystems contain plants that are both fire-adapted and flammable. It has been hypothesized that these plants were under selection to become more flammable, but it is unclear whether this could be adaptive for an individual plant. We propose arrested succession as a robust mechanism that supports the evolution of flammability in surface fire ecosystems without the need to invoke group selection or additional fitness benefits. We used the natural history of lodgepole pine (Pinus ponderosa) forests, longleaf pine (Pinus palustris) forests, and tall grass prairies to create a general mathematical model of surface fire ecosystems and solved for the evolutionarily stable strategy (ESS) level of flammability. In our model, fires always kill understory plants and only sometimes kill overstory plants. Thus, more flammable plants suffer increased mortality due to fires, but also more frequently arrest succession by clearing their understory of late successional competitors. Increased flammability was selected for when the probability of an overstory plant dying from an individual fire was below a maximum threshold and the rate of succession relative to fires was above a minimum threshold. Future studies can test our model predictions and help resolve whether or not plants have been selected to be more flammable.     

Effects of high‐severity fire drove the population collapse of the subalpine Tasmanian endemic conifer Athrotaxis cupressoides

Athrotaxis cupressoides is a slow‐growing and long‐lived conifer that occurs in the subalpine temperate forests of Tasmania, a continental island to the south of Australia. In 1960–1961, human‐ignited wildfires occurred during an extremely dry summer that killed many A. cupressoides stands on the high plateau in the center of Tasmania. That fire year, coupled with subsequent regeneration failure, caused a loss of ca. 10% of the geographic extent of this endemic Tasmanian forest type. To provide historical context for these large‐scale fire events, we (i) collected dendroecological, floristic, and structural data, (ii) documented the postfire survival and regeneration of A. cupressoides and co‐occurring understory species, and (iii) assessed postfire understory plant community composition and flammability. We found that fire frequency did not vary following the arrival of European settlers, and that A. cupressoides populations were able to persist under a regime of low‐to‐mid severity fires prior to the 1960 fires. Our data indicate that the 1960 fires were (i) of greater severity than previous fires, (ii) herbivory by native marsupials may limit seedling survival in both burned and unburned A. cupressoides stands, and (iii) the loss of A. cupressoides populations is largely irreversible given the relatively high fuel loads of postfire vegetation communities that are dominated by resprouting shrubs. We suggest that the feedback between regeneration failure and increased flammability will be further exacerbated by a warmer and drier climate causing A. cupressoides to contract to the most fire‐proof landscape settings.     

Understory Plant Community Composition Is Associated with Fine-Scale Above- and Below-Ground Resource Heterogeneity in Mature Lodgepole Pine (Pinus contorta) Forests

Understory plant communities play critical ecological roles in forest ecosystems. Both above- and below-ground ecosystem properties and processes influence these communities but relatively little is known about such effects at fine (i.e., one to several meters within-stand) scales, particularly for forests in which the canopy is dominated by a single species. An improved understanding of these effects is critical for understanding how understory biodiversity is regulated in such forests and for anticipating impacts of changing disturbance regimes. Our primary objective was to examine the patterns of fine-scale variation in understory plant communities and their relationships to above- and below-ground resource and environmental heterogeneity within mature lodgepole pine forests. We assessed composition and diversity of understory vegetation in relation to heterogeneity of both the above-ground (canopy tree density, canopy and tall shrub basal area and cover, downed wood biomass, litter cover) and below-ground (soil nutrient availability, decomposition, forest floor thickness, pH, and phospholipid fatty acids (PLFAs) and multiple carbon-source substrate-induced respiration (MSIR) of the forest floor microbial community) environment. There was notable variation in fine-scale plant community composition; cluster and indicator species analyses of the 24 most commonly occurring understory species distinguished four assemblages, one for which a pioneer forb species had the highest cover levels, and three others that were characterized by different bryophyte species having the highest cover. Constrained ordination (distance-based redundancy analysis) showed that two above-ground (mean tree diameter, litter cover) and eight below-ground (forest floor pH, plant available boron, microbial community composition and function as indicated by MSIR and PLFAs) properties were associated with variation in understory plant community composition. These results provide novel insights into the important ecological associations between understory plant community composition and heterogeneity in ecosystem properties and processes within forests dominated by a single canopy species.     

Forest crown density restoration and influencing factors in the burned area of northern Great Hing′an Mountains of China

Fire in the Great Hing′an Mountains in 1987 affected an area of more than 1.33×10⁶ hm², creating a mosaic of burn severities across the landscape, which strongly affected the postfire vegetation succession. In addition, undulate landform and anthropogenic disturbance inevitably influenced the postfire vegetation succession. In this paper, a typical area was selected for a case study, including two forest farms, covering more than 1.2×10⁵ hm². In order to reveal how the forest changed in 2000 (13 years after the fire) by comparing with 1987 (prefire) and to find out the relationship between the forest succession and the affecting factors, forest crown density was selected as the criterion, and forest type, fire severity, silviculture practice, elevation and topography gradients were designed as the affecting variables. With the support of GIS software, each variable was classified and entered into the multivariate regression model. The result showed that the forest crown density changed notably in 2000 compared with that of the prefire, and all the variables significantly affected the forest crown density. The most important affecting variable was elevation, which was positively correlated with the forest crown density. The next was fire severity, which was negatively related with the forest succession. The effects of topographic factors and silviculture practices on forest crown density were relatively small.     

Prefire grazing by cattle increases postfire resistance to exotic annual grass (Bromus tectorum) invasion and dominance for decades

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Aboveground Net Primary Production and Leaf-Area Index in Early Postfire Vegetation in Yellowstone National Park

Aboveground net primary production (ANPP) and leaf-area index (LAI) of lodgepole pine (Pinus contorta var. latifolia Engelm. ex Wats.) saplings and aboveground productivity of herbaceous vegetation components were determined 9 years after the 1988 fires in Yellowstone National Park (YNP). Measurements were made in four sites representing a wide range of early postfire vegetation present in YNP, including high-density lodgepole pine, low-density lodgepole pine, and two nonforest stands. LAI of the pine saplings and total ANPP (trees plus herbs) generally increased with increasing sapling density, from 0.002 m² m^{− 2} and 0.25 Mg ha^{− 1} year^{− 1} in the infertile nonforest stand (100 pine saplings ha^{− 1}) to 1.8 m² m^{− 2} and 4.01 Mg ha^{− 1} year^{− 1} in the high-density pine stand (62,800 saplings ha^{− 1}). Aboveground herbaceous productivity was not strongly correlated with sapling density, but appeared to be influenced by soil fertility. In the high-density pine stand, tree ANPP and LAI were within the lower range of values reported for similar mature coniferous forests. This finding suggests that at least some ecosystem processes (related to ANPP and LAI) may have nearly recovered after only 9 years of postfire succession, in at least some of the young forests developing after the 1988 Yellowstone fires. Received 7 April 1998; accepted 1 December 1998.     

Interactions of Fire and Nonnative Species Across an Elevation/Plant Community Gradient in Hawaii Volcanoes National Park

Invasive species interacting with fires pose a relatively unknown, but potentially serious, threat to the tropical forests of Hawaii. Fires may create conditions that facilitate species invasions, but the degree to which this occurs in different tropical plant communities has not been quantified. We documented the survival and establishment of plant species for 2 yr following 2003 wildfires in tropical moist and wet forest life zones in Hawaii Volcanoes National Park, Hawaii. Fires were ignited by lava flows and burned across a steep environmental gradient encompassing two previously burned shrub-dominated communities and three Metrosideros polymorpha forest communities. Fires in all community types were stand replacing, where >95 percent of overstory trees were top killed. Over half (>57%) of the trees survived via basal sprouting, but sprout growth differed among forest communities. Sprout growth (>250,000 cm³) was greatest in the forest community where postfire understory cover was lowest presumably due to thick native Dicranopteris linearis fern litter that remained postfire. In contrast, M. polymorpha sprout growth was much slower (<100,000 cm³) in the two forest communities where there was rapid understory recovery of nonnative ferns Nephrolepis multiflora and invasive grasses Paspalum conjugatum. These results suggest that the rapid establishment of an invasive-dominated understory limited recovery of the overstory dominant M. polymorpha. In contrast to the three forest communities, there were few changes in vegetation composition in the shrubland communities. Nonnative species invasions coupled with repeated fires selectively eliminated fire-sensitive species thereby maintaining these communities in dominance of primarily nonnative, fire-resilient, species.