Post‐fire succession and 20th century reduction in fire frequency on xeric southern Appalachian sites

Abstract. We document post‐fire succession on xeric sites in the southern Appalachian Mountains, USA and assess effects of 20th century reduction in fire frequency on vegetation structure and composition. Successional studies over 18 yr on permanent plots that had burned in 1976–1977 indicate that tree mortality and vegetation response varied with fuel load and fire season. In the first three years after fire, hardwood sprouts dominated tree regeneration. On sites where summer and autumn fires reduced litter depth to less than 1 cm, densities of shade‐intolerant Pinus seedlings increased steadily over this period. 4 to 8 yr after fire, large numbers of newly established seedlings and sprouts had grown to 1 – 10 cm DBH. By year 18 growth of these saplings led to canopy closure on most sites. Herbaceous cover and richness peaked in the first decade after fire, then declined. On similar sites that had not burned in more than 50 yr, regeneration of shade‐intolerant Pinus spp. and mean cover and richness of herbs were considerably lower than those observed on recently burned plots. Reconstructions of landscape conditions based on observed post‐fire succession and 20th century changes in fire regime suggest that reductions in fire frequency circa 1940 led to substantial changes in forest structure and decreases in cover and richness of herbaceous species.     

Restoration of C4 grasses with seasonal fires in a C3/C4 grassland invaded by Prosopis glandulosa,a fire‐resistant shrub

Questions: Can prescribed fire restore C₄ perennial grasses in grassland ecosystems that have become dominated by fire‐resistant C₃ shrubs (Prosopis glandulosa) and C₃ grasses? Do fires in different seasons alter the direction of change in grass composition? Location: Texas, USA. Methods: We quantified short‐ and long‐term (12 yr post‐fire) herbaceous functional group cover and diversity responses to replicated seasonal fire treatments: (1) repeated‐winter fires (three in 5 yr), (2) repeated‐summer fires (two in 3 yr), and (3) alternate‐season fires (two winter and one summer in 4 yr), compared with a no‐fire control. Results: Summer fires were more intense than winter fires, but all fire treatments temporarily decreased Prosopis and C₃ annual grass cover. The alternate‐season fire treatment caused a long‐term increase in C₄ mid‐grass cover and functional group diversity. The repeated‐summer fire treatment increased C₄ short‐grass cover but also caused a long‐term increase in bare ground. The repeated winter fire treatment had no long‐term effects on perennial grass cover. Mesquite post‐fire regrowth had increasingly negative impacts on herbaceous cover in all fire treatments. Conclusions: Summer fire was necessary to shift herbaceous composition toward C₄ mid‐grasses. However, the repeated‐summer fire treatment may have been too extreme and caused post‐fire herbaceous composition to “over‐shift” toward less productive C₄ short‐grasses rather than C₄ mid‐grasses. This study provides some of the first long‐term data showing a possible benefit of mixing seasonal fires (i.e., the alternate‐season fire treatment) in a prescribed burning management plan to restore C₄ mid‐grass cover and enhance overall herbaceous diversity.     

Plant functional group responses to fire frequency and tree canopy cover gradients in oak savannas and woodlands

Questions: How do fire frequency, tree canopy cover, and their interactions influence cover of grasses, forbs and understorey woody plants in oak savannas and woodlands? Location: Minnesota, USA. Methods: We measured plant functional group cover and tree canopy cover on permanent plots within a long‐term prescribed fire frequency experiment and used hierarchical linear modeling to assess plant functional group responses to fire frequency and tree canopy cover. Results: Understorey woody plant cover was highest in unburned woodlands and was negatively correlated with fire frequency. C4‐grass cover was positively correlated with fire frequency and negatively correlated with tree canopy cover. C3‐grass cover was highest at 40% tree canopy cover on unburned sites and at 60% tree canopy cover on frequently burned sites. Total forb cover was maximized at fire frequencies of 4–7 fires per decade, but was not significantly influenced by tree canopy cover. Cover of N‐fixing forbs was highest in shaded areas, particularly on frequently burned sites, while combined cover of all other forbs was negatively correlated with tree canopy cover. Conclusions: The relative influences of fire frequency and tree canopy cover on understorey plant functional group cover vary among plant functional groups, but both play a significant role in structuring savanna and woodland understorey vegetation. When restoring degraded savannas, direct manipulation of overstorey tree canopy cover should be considered to rapidly reduce shading from fire‐resistant overstorey trees. Prescribed fires can then be used to suppress understorey woody plants and promote establishment of light‐demanding grasses and forbs.     

Change over 70 years in a southern California chaparral community related to fire history

Question: What changes in species composition and cover have occurred in chaparral as a function of fire history across an ecoregion? Location: San Diego County, California, USA. Methods: Stands in which 40 mid‐elevation chaparral vegetation plots (each 400 m² in area) were located in the 1930s were resurveyed in 2001. We stratified the stands into Infrequently versus Frequently burned (0–1 versus 2 or more fires recorded in the 91‐yr period), and Immature versus Mature (ã31 yr versus >31 yr since last fire), resulting in four groups. Ten stands were randomly selected from each of these groups for survey. Results: There were no major shifts in life form composition, e.g., live oak trees were not invading chaparral that had experienced little or no fire, nor were subshrubs or herbaceous species replacing shrubs in areas that had experienced more frequent fires. However, there was a notable increase in the frequency of the subshrub Eriogonum fasciculatum across all fire history groups. In the mature stands with infrequent fire, average cover of resprouting shrubs increased (from 72 to 91%) and cover of obligate seeding shrubs (species with fire‐cued germination) decreased (from 21 to 6%) significantly. Mature stands with frequent fire showed a significant decrease in resprouter cover (from 87 to 80%) and increase in obligate seeders (from 10 to 16%). Conclusions: While the tremendous changes in land use in southern California have been predicted to cause shifts in chaparral composition, these shifts are difficult to detect because species longevity and fire cycles are on the order of decades to a century. In this study, the expected trends could only be detected in groups that were mature at the time of the second survey.     

Effects of inter‐fire intervals on the reproductive output of resprouters and obligate seeders in the Proteaceae

Abstract Fire is often used as a management tool in fire‐prone communities to reduce fuel loads with the intention of reducing the severity and extent of unplanned fires, often resulting in the increased occurrence of fire in the dry sclerophyll vegetation of Australia. This study examined the effects of fire frequency (length of the inter‐fire interval) on the reproductive output of seven plant species in the Proteaceae, including obligate seeding shrubs (Hakea teretifolia, Petrophile pulchella), resprouting shrubs (Banksia spinulosa, Isopogon anemonifolius, Lambertia formosa) and resprouting trees (Banksia serrata, Xylomelum pyriforme). Reproductive output (measured as either number of confructescences or follicles) and relative size were estimated for 100 individuals at each of five sample sites, covering a range of past fire frequencies over 26 years including repeated short inter‐fire intervals. Patterns in reproductive output (after standardizing for size) were related to the life‐history attributes of the species. In areas that had experienced short inter‐fire intervals, obligate seeders had greater reproductive output compared with longer intervals, and the reproductive output of resprouting shrubs was less. Fire frequency did not affect reproductive output of the resprouting trees. The decreased reproductive output of the resprouting shrubs could be due to the allocation of resources to regrowth following fire rather than to reproduction. It is less clear what process resulted in the increased reproductive output of obligate seeders in high fire frequency areas, but it could be due to the most recent fires being more patchy in the areas experiencing shorter inter‐fire intervals, or it may have resulted from the selection for early reproduction in the high fire frequency areas. These results highlight the need to take into account past fire frequency at a site, in addition to time since the last fire, when planning prescribed fires.     

Anthropogenic fires increase alien and native annual species in the Chilean coastal matorral

Aim We tested the hypothesis that anthropogenic fires favour the successful establishment of alien annual species to the detriment of natives in the Chilean coastal matorral. Location Valparaíso Region, central Chile. Methods We sampled seed rain, seedbank emergence and establishment of species in four paired burned and unburned areas and compared (using GLMM) fire resistance and propagule arrival of alien and native species. To assess the relative importance of seed dispersal and seedbank survival in explaining plant establishment after fire, we compared seed rain and seedbank structure with post‐fire vegetation using ordination analyses. Results Fire did not change the proportion of alien species in the coastal matorral. However, fire increased the number of annual species (natives and aliens) of which 87% were aliens. Fire reduced the alien seedbank and not the native seedbank, but alien species remained dominant in burned soil samples (66% of the total species richness). Seed rain was higher for alien annuals than for native annuals or perennials, thus contributing to their establishment after fire. Nevertheless, seed rain was less important than seedbank survival in explaining plant establishment in burned areas. Main conclusions Anthropogenic fires favoured alien and native annuals. Thus, fire did not increase the alien/native ratio but increased the richness of alien species. The successful establishment of alien annuals was attributable to their ability to maintain rich seedbanks in burned areas and to the greater propagule arrival compared to native species. The native seedbank also survived fire, indicating that the herbaceous community has become highly resilient after centuries of human disturbances. Our results demonstrate that fire is a relevant factor for the maintenance of alien‐dominated grasslands in the matorral and highlight the importance of considering the interactive effect of seed rain and seedbank survival to understand plant invasion patterns in fire‐prone ecosystems.     

Impact of two wildfires on endemic granite outcrop vegetation in Western Australia

Abstract. In seasonally dry regions of the world fire is a recurring disturbance but little is known of how fire interacts with granite outcrop vegetation. We hypothesize that the floristic composition in granite vegetation, usually attributed to the edaphic environment, may also reflect the impact of disturbances such as fire. Dramatic differences in floristic composition and cover over 13 years and two fires were observed in vegetation on a Western Australian granite outcrop. This was very marked in the first year following the two fires, with annuals and geophytes showing the greatest turnover of species. Even among the perennial shrubs there was considerable turnover in a number of obligate seeders. After the first fire the number of species declined for woody perennials, herbaceous perennials and annuals, remained unchanged for perennial grasses and sedges, and varied with highest richness 4 yr after fire for geophytes. Demographic studies of two endemic woody obligate seeders and three endemic mallee eucalypt resprouters similarly showed dramatic differences within and between species in seedling recruitment following the two fires. Fire does have a significant impact on the floristic composition of semi‐arid granite outcrop vegetation communities. Studies on other granite outcrop systems are needed to test the generality of this conclusion.     

Response of plant species and life form diversity to variable fire histories and biomass in the jarrah forest of south‐west Australia

Frequent fires reduce the abundance of woody plant species and favour herbaceous species. Plant species richness also tends to increase with decreasing vegetation biomass and cover due to reduced competition for light. We assessed the influence of variable fire histories and site biomass on the following diversity measures: woody and herbaceous species richness, overall species richness and evenness, and life form evenness (i.e. the relative abundance or dominance among six herbaceous and six woody plant life forms), across 16 mixed jarrah (Eucalyptus marginata) and marri (Corymbia calophylla) forest stands in south‐west Australia. Fire frequency was defined as the total number of fires over a 30‐year period. Overall species richness and species evenness did not vary with fire frequency or biomass. However, there were more herbaceous species (particularly rushes, geophytes and herbs) where there were fewer shrubs and low biomass, suggesting that more herbaceous species coexist where dominance by shrubs is low. Frequently burnt plots also had lower number and abundance of shrub species. Life form evenness was also higher at both high fire frequency and low biomass sites. These results suggest that the impact of fire frequency and biomass on vegetation composition is mediated by local interactions among different life forms rather than among individual species. Our results demonstrate that measuring the variation in the relative diversity of different woody and herbaceous life forms is crucial to understanding the compositional response of forests and other structurally complex vegetation communities to changes in disturbance regime such as increased fire frequency.     

Impact of shrub canopies on understorey vegetation in western Eurasian tundra

Question: How does the composition and species richness of understorey vegetation associate with changing abundance of deciduous shrub canopies? What are the species‐specific associations between shrubs and understorey plants? Location: Tundra habitats along an over 1000‐km long range, spanning from NW Fennoscandia to the Yamal Peninsula in northwest Russia. Methods: The data from 758 vegetation sample plots from 12 sites comprised cover estimates of all plant species, including bryophytes and lichens, and canopy height of deciduous shrubs. The relationships between shrub volume and cover of plant groups and species richness of vegetation were investigated. In addition, species‐specific associations between understorey species and shrub volume were analysed. Results: Shrub abundance was shown to be associated with the composition of understorey vegetation, and the association patterns were consistent across the study sites. Increased forb cover was positively associated with shrub volume, whereas bryophyte, lichen, dwarf shrub and graminoid cover decreased in association with increasing volume of deciduous shrubs. The total species richness of vegetation declined with increasing shrub volume. Conclusions: The results suggest that an increase of shrubs – due to climatic warming or a decrease in grazing pressure – is likely to have strong effects on plant–plant interactions and lead to a decrease in the diversity of understorey vegetation.     

Population size and fire intensity determine post‐fire abundance in grassland lichens

Questions: What is the variability in abundance of lichens on grassland soil between and within fields after prescribed fire? Is post‐fire lichen abundance an effect of pre‐fire population size? Location: Cedar Creek Natural History Area, Minnesota, USA. Methods: Lichen abundance, estimated as ground cover and dominated by Cladonia spp., was mapped in plots in two fields before prescribed burning on 06.10.2003 and 15.10.2003 for the first time since abandonment in the 1950s. The plots were resurveyed one year post‐fire. Results: Post‐fire cover of Cladonia spp. varied strongly between the fields, most likely due to different weather conditions between the burn events, which resulted in different fire intensities, one of low and one of high intensity. In the field that experienced the low intensity fire, post‐fire cover of Cladonia spp. was still relatively high, and showed a positive relationship with pre‐fire cover, while no such relationship was found after the high intensity fire. In that field Cladonia spp. experienced high mortality rates irrespective of pre‐fire cover. Conclusions: This study provides an example of how species response to disturbance can be a function of population size, but that this relationship can be non‐linear; lichens in grassland can survive a low intensity fire proportionally to pre‐fire population size, but experience high mortality rates above a fire intensity threshold. The applications of these results are that fire intensity matters to species response to prescribed fire, and that the persistence of climax lichen communities and biodiversity in the study system needs a broad range of fire intervals.     

Fire and competition in Australian heath: a conceptual model and field investigations

Abstract. We describe a model of heath vegetation, in which species were classified into five functional groups based on characteristics of their propagule pools, post-fire growth, timing and mode of reproduction and competitive status. The model assumes no recruitment without fire and a simple competitive hierarchy based on vertical stature. A critical feature of the model is an initial post-fire window of 5–6 yr in which competition from overstorey species on understorey species is reduced. Understorey functional groups differ in their ability to exploit this window. In the field, we tested five predictions derived from the model: (a) overall species richness of understorey varies inversely with overstorey density as a result of a trend in richness of woody species, but not in herbaceous species; (b) where an overstorey was present in the previous fire interval, post-fire population density is reduced in a functional group of understorey serotinous resprouting shrubs, but not in a group of understorey obligate-seeding shrubs with soil seed banks; (c) in understorey serotinous resprouting shrubs, post-fire regrowth in resprouting individuals is adversely affected by the presence of an overstorey in the preceding fire interval; (d) in understorey serotinous resprouting shrubs, levels of pre-fire propagules are lower in the presence of an overstorey, reducing the density of post-fire recruits; and (e) in understorey serotinous resprouting shrubs, recruitment relative to the pre-fire population is unaffected by overstorey species within the window of reduced competition. Of these, three tests (a,b,d) supported the model, one (e) may support the model, but the results were inconclusive and one (c) did not support the model. Limitations and further applications of the model are discussed. Our results suggest that maintenance of high densities of overstorey populations is in conflict with conservation of some understorey species. Models of the type we propose will help identify and resolve such conflicts and promote the judicious use of fire to maintain full species diversity of plant communities.     

Interaction of multiple disturbances: importance of disturbance interval in the effects of fire on rehabilitating mined areas

Abstract Multiple disturbance regimes are increasingly common as novel anthropogenic disturbances are added to existing natural disturbances. However, it is generally unknown whether simultaneous or sequential effects of different forms of disturbance are predictable from the independent effects of each disturbance. This study examines the short‐term effects of sequential disturbance by mineral sand‐mining followed by fire in a forest community in south‐eastern Australia. Four combinations of disturbance were sampled: unburned mined, burned mined, unburned forest (unmined) and burned forest (unmined, with between‐fire interval matching the disturbance interval between mining and fire of the burned mined treatment). All combinations were sampled approximately 12 months following fire on the burned sites. The impact of fire after mining depended on disturbance interval. Sites burned 0.5–2.4 years since mining had fewer native vascular plant species than unburned mined sites of the same mined age, whereas sites with 10–16 years or 20–26 years between mining and fire had greater native species richness than unburned mined sites of the same age. Burning 20–26 years after mining brought native species richness within the range of burned forest. For both unmined and mined sites native seedling densities increased with burning, and with longer disturbance intervals. Weed species richness and weed seedling densities were greater on mined sites than in forest, and burning mined sites elevated weed seedling densities further, particularly for short intervals. Both disturbance interval and fire intensity are likely to have contributed to these results, as intensity on mined areas increased with interval, and at 20–26 years post‐mining was equivalent to unmined forest. These results suggest that fire could be used to promote rehabilitation of these mined areas after at least 10 years, but should be excluded from earlier stages of post‐mining regeneration. However, other sources of spatial and temporal variability should be considered in addition to interval and intensity, as variation among mined areas was correlated with post‐fire weather conditions and available weed sources. Finally, the combined effects of mining and fire could not be predicted from knowledge of the disturbances operating separately, indicating that effects of multiple disturbance may be synergistic rather than additive.     

Are long‐unburnt eucalypt forest patches important for the conservation of plant species diversity?

Question: Are long‐unburnt patches of eucalypt forest important for maintaining floristic diversity? Location: Eucalyptus forests of southeastern New South Wales, Australia. Methods: Data from 976 sites representing a range of fire history from three major vegetation formations – shrubby dry sclerophyll forest (SF), grassy dry SF and wet SF – were analysed. Generalized linear models were used to examine changes in species richness with increasing time since wildfire and analysis of similarities to examine changes in community composition. Chi‐squared tests were conducted to examine the distribution of individual species across four time since fire categories. Results: Plant species relationships to fire varied between the three formations. Shrubby dry SF supported lower plant species richness with increasing time since wildfire and this was associated with shifts in community composition. Grassy dry SF showed significant shifts in community composition and species richness in relation to time, with a peak in plant species richness 20–30 yr post fire (either prescribed fire or wildfire). Wet SF increased in species richness until 10–20 yr post wildfire then displayed a general declining trend. Species richness in each vegetation type was not related to the fire frequencies and fire intervals observed in this study. Conclusions: Long‐unburnt (30–50 yr post wildfire) forests appeared to play a minor role in the maintenance of plant species diversity in dry forest systems, although this was more significant in wet forests. Maintenance of a range of fire ages within each vegetation formation will assist in maintaining floristic diversity within regions.     

Longleaf pine (Pinus palustris Mill.) fire scars reveal new details of a frequent fire regime

Question: How frequent and variable were fire disturbances in longleaf pine ecosystems? Has the frequency and seasonality of fire events changed during the past few centuries? Location: Kisatchie National Forest, Western Gulf Coastal Plain, longleaf pine–bluestem ecosystem, in relatively rough topography adjacent to the Red River, Louisiana, USA. Methods: Cross‐sections of 19 remnant pines exhibiting 190 fire scars were collected from a 1.2‐km² area. Tree‐rings and fire scars were precisely dated and analysed for the purpose of characterizing past changes in fire and tree growth. Temporal variability in fire occurrence and seasonality was described for the pre‐ and post‐European settlement periods. Seasonality of historic fires was determined by the scar position within the rings. The relationship between fire and drought was investigated using correlation and superposed epoch analysis. Results: The mean fire return interval for the period 1650‐1905 was 2.2 years (range 0.5 to 12 yr). Significant new findings include: evidence for years of biannual burning, temporal variability in fire seasonality, an increase in fire frequency and percentage of trees scarred circa 1790, and synchronous growth suppression and subsequent release of trees coinciding with land‐use changes near the turn of the 20th century. Drought conditions appeared unrelated to the occurrence of fire events or fire seasonality. Conclusions: Multi‐century fire history records from longleaf pine ecosystems are difficult to obtain due to historic land‐use practices and the species high resistance to scarring; however, our results indicate potential for reconstructing detailed fire histories in this ecosystem. Fire scars quantitatively documented one of the most frequent fire regimes known. Fire regime information, such as the temporal variability in fire intervals, prevalence of late‐growing season fire events and biannual burning, provide a new perspective on the dynamics of longleaf pine fire regimes.     

Influence of fire on critically endangered Swartland Shale Renosterveld in the Cape Floristic Region

Questions

The degree to which renosterveld shrublands are fire‐dependent is currently unclear. To address this issue, the following questions were asked: (1) does smoke stimulate germination of soil‐stored seeds in renosterveld; (2) does recently‐burned renosterveld display changed composition and higher diversity than unburned vegetation; and (3) how do the species compositions of renosterveld soil seed banks and standing vegetation compare?

Location

Swartland, Cape Floristic Region, South Africa.

Methods

Soil seed bank samples from a north‐ and south‐facing slope were smoke‐treated and germinated to test for smoke‐stimulated germination. Burned standing vegetation was surveyed 16 months post‐fire, as was unburned vegetation on the same slopes. Seed bank species richness and density were compared between smoke‐treated and untreated samples within and between slopes. Burned and unburned standing vegetation were compared within and between slopes in terms of species richness, abundance and aerial cover. Compositional similarity of the seed banks and standing vegetation was assessed.

Results

Seed banks were dominated by annuals and graminoids. Smoke treatment had no effect, except for driving significantly higher species richness and seedling density in south‐facing slope perennial shrubs. Species richness and seedling density were significantly higher in seed banks on the south‐facing slope compared to the north‐facing slope. Burned standing vegetation exhibited significantly higher diversity than unburned vegetation. Annuals and graminoids displayed significantly higher species richness and aerial cover in burned renosterveld. The north‐facing slope contained less than half the number of species/m² compared to the south‐facing slope. The seed banks and standing vegetation showed low to intermediate similarity (Sørensen = 31%–53%), but grouped close together on an NMDS plot, suggesting intermediate similarity overall.

Conclusions

Elevated germination of perennial shrubs in smoke‐treated seed bank samples and increased diversity of post‐fire standing vegetation suggest the renosterveld in this study shows elements of a fire‐driven system. Certain species only recruited in burned sites, suggesting fire‐stimulated germination. Aspect had a major influence on plant community composition, with the mesic south‐facing slope being more diverse than the xeric north‐facing slope. The similarity between the seed banks and standing vegetation was higher than previously shown for renosterveld, and appears to be higher than for fynbos.     

Rainforest expansion reduces understorey plant diversity and density in open forest of eastern Australia

The expansion of rainforest pioneer trees into long‐unburnt open forests has become increasingly widespread across high rainfall regions of Australia. Increasing tree cover can limit resource availability for understorey plant communities and reduce understorey diversity. However, it remains unclear if sclerophyll and rainforest trees differ in their competitive exclusion of understory plant communities, which contain most of the floristic diversity of open forests. Here, we examine dry open forest across contrasting fire histories (burnt and unburnt) and levels of rainforest invasion (sclerophyll or rainforest midstorey) to hindcast changes in understorey plant density, richness and composition. The influence of these treatments and other site variables (midstorey structure, midstorey composition and soil parameters) on understorey plant communities were all examined. This study is the first to demonstrate significantly greater losses of understorey species richness, particularly of dry open‐forest specialists, under an invading rainforest midstorey compared to a typical sclerophyll midstorey. Rainforest pioneers displaced over half of the understorey plant species, and reduced ground cover and density of dry forest specialists by ~90%. Significant understorey declines also occurred with increased sclerophyll midstorey cover following fire exclusion, although losses were typically less than half that of rainforest‐invaded sites over the same period. Understorey declines were closely related to leaf area index and basal area of rainforest and wattle trees, suggesting competitive exclusion through shading and potentially belowground competition for water. Around 20% of displaced species lacked any capacity for population recovery, while transient seed banks or distance‐limited dispersal may hinder recovery for a further 68%. We conclude that rainforest invasion leads to significant declines in understorey plant diversity and cover in open forests. To avoid elimination of local native plant populations in open forests, fires should occur with sufficient frequency to prevent overstorey cover from reaching a level where shade‐intolerant species fail to thrive.     

Spatial patterns of desert annuals in relation to shrub effects on soil moisture

Questions: What are the effects of a shrub (Haloxylon ammodendron) on spatial patterns of soil moisture in different seasons? How does productivity of understorey annuals respond to these effects? Are such effects always positive for annuals under shrubs? Location: South Gurbantunggut Desert, northwest China. Methods: Using geostatistics, we explored seasonal patterns of topsoil moisture in a 12 × 9‐m plot over the growing season. To determine spatial patterns of understorey annuals in response to H. ammodendron presence, biomass of annuals was recorded in four 0.2 × 5.0‐m transects from the centre of a shrub to the space between shrubs (interspace). We also investigated vertical distribution of root biomass for annuals and soil moisture dynamics across soil profiles in shrub‐canopied areas and interspaces. Results: Topsoil moisture changed from autocorrelation in the wet spring to random structure in the dry season, while soil moisture below 20 cm was higher in shrub‐canopied areas. Across all microhabitats, soil moisture in upper soil layers was higher than in deeper soil layers during the spring wet season, but lower during summer drought. Topsoil was close to air‐dry during the dry season and developed a ‘dry sand layer’ that reduced evaporative loss of soil water from deeper layers recharged by snowmelt in spring. Aboveground biomass of understorey annuals was lowest adjacent to shrub stems and peaked at the shrub margin, forming a ‘ring’ of high herbaceous productivity surrounding individual shrubs. To acclimate to drier conditions, annuals in interspaces invested more root biomass in deeper soil with a root/shoot ratio (R/S) twice that in canopied areas. Conclusions: Positive and negative effects of shrubs on understorey plants in arid ecosystems are commonly related to nature of the environmental stress and tested species. Our results suggest there is also microhabitat‐dependence in the Gurbantunggut Desert. Soil water under H. ammodendron is seasonally enriched in topsoil and deeper layers. Understorey annuals respond to the effect of shrubs on soil water availability with lower R/S and less root biomass in deeper soil layers and develop a ‘ring’ of high productivity at the shrub patch margin where positive and negative effects of shrubs are balanced.     

The effect of short fire cycles on the cover and density of understorey sprouting species in South African mountain fynbos

Abstract. Two South African mountain fynbos sites were studied to determine the effect of short fire cycles on the cover and density of understorey sprouting species and their subsequent effect on plant-species richness. Frequent fires (4–6 years between burns) increased the cover of sprouting species by 32% when compared to an adjacent site where the penultimate fire was 28 years previously. There was little or no effect of fire frequency on the densities of understorey sprouters; however, individuals were larger at sites with short fire cycles. The response of individual species of sprouters was variable with one species, Hypodiscus striatus , showing no response to fire frequency. The impact of sprouting species on the species richness of the plant community was great. The mean number of species recorded in quadrats with a high cover of sprouters was 60% lower in comparison to quadrats with low covers or under the burned skeletons of overstorey proteas. The effect of sprouters was consistent for all functional groups of species (i.e. sprouters, non-sprouters, short-lived and long-lived species), in each case reducing the number of species present.     

Initial response of understorey vegetation to fire severity and salvage‐logging in the southern boreal forest of Québec

Abstract. In this study we compared the effects of fire on understorey vegetation in the Québec southern boreal forest with effects of salvage‐logging (clear‐cutting after fire). All 61 400‐m² sampling sites were controlled for overstorey composition (Deciduous, Mixed and Coniferous) and disturbance type, which consisted of three fire impact severity (FIS) classes (Light, Moderate and Extreme) and two harvesting techniques (Stem‐only and Whole‐tree Harvesting). Percent‐cover data of vegetation and post‐disturbance environmental characteristics were recorded in the field during the first two years after fire as well as soil texture. Ordination of fire alone demonstrated that, on Coniferous sites, fire initiates a succession whereby the understorey Coniferous sites approaches that of Deciduous‐Mixed sites, due to the release of the understorey from Sphagnum spp. dominance, this pattern being a function of FIS. On Deciduous‐Mixed stands, increased FIS resulted in a transition from herb to shrub dominance. Ordination of all five disturbance types showed that the impact of salvage‐logging on understorey composition was within the range of fire, but marginalized to the extreme end of the FIS spectrum. Variance partitioning demonstrated that overstorey and soil texture were the most important explanatory variables of fire alone, while disturbance type explained the largest independent fraction of understorey variation when salvage‐logging was introduced. Salvage‐logging also results in significant reductions in understorey abundance, richness and diversity, while indicator species analysis suggests that it favours mesoxerophytic to xeric species. Results are interpreted in light of shade‐tolerance dynamics, forest floor disturbance and soil moisture regimes. Implications for sustainable forest management are discussed.     

Using biodiversity deconstruction to disentangle assembly and diversity dynamics of understorey plants along post‐fire succession in boreal forest

Aim The study aims to decipher the co‐occurrence of understorey plant assemblages and, accordingly, to identify a set of species groups (diversity deconstruction) to better understand the multiple causal processes underlying post‐fire succession and diversity patterns in boreal forest. Location North‐eastern Canadian boreal forest (49°07′–51°44′ N; 70°13′–65°15′ W). Methods Data on understorey plant communities and habitat factors were collected from 1097 plots. Species co‐occurrence was analysed using null model analysis. We derive species groups (i.e. biodiversity deconstruction) using the strength of pairwise species co‐occurrences after accounting for random expectation under a null model and cluster analyses. We examine the influence of a set of spatiotemporal environmental variables (overstorey composition, time‐since‐fire, spatial location and topography) on richness of species groups using Bayesian model averaging, and their relative influence through hierarchical partitioning of variance. Results Understorey plant assemblages were highly structured, with co‐occurrence‐based classification providing species groups that were coherently aggregated within, but variably segregated between, species groups. Group richness models indicate both common and distinct responses to factors affecting plant succession. For example, Group 2 (e.g. Rhododendron groenlandicum and Cladina rangiferina) showed concurrent contrasting responses to overstorey composition and was strongly segregated from Groups 3 (e.g. Clintonia borealis and Maianthenum canadense) and 4 (e.g. Epilobium angustifolium and Alnus rugosa). Groups 3 and 4 showed partial similarity, but they differed in their response to time‐since‐fire, drainage and latitude, which were more important for Group 1 (e.g. Ptilium crista‐castrensis and Empetrum nigrum). A single successional model based on total richness masked crucial group‐level relationships with factors that we examined, such as latitude. Main conclusions By demonstrating the co‐occurrence structure and linking to causal factors, the results from this study characterize both common and distinct responses of understorey plants to biophysical attributes of sites, and potential interspecific interactions, behind non‐random assemblage structure during post‐fire succession. A biodiversity deconstruction approach could offer a concise and explicit framework to gain a better understanding of the complex assembly of ecological communities during succession.