Influence of grazing and fire frequency on small-scale plant community structure and resource variability in native tallgrass prairie

In grasslands worldwide, grazing by ungulates and periodic fires are important forces affecting resource availability and plant community structure. It is not clear, however, whether changes in community structure are the direct effects of the disturbance (i.e. fire and grazing) or are mediated indirectly through changes in resource abundance and availability. In North American tallgrass prairies, fire and grazing often have disparate effects on plant resources and plant diversity, yet, little is known about the individual and interactive effects of fire and grazing on resource variability and how that variability relates to heterogeneity in plant community structure, particularly at small scales. We conducted a field study to determine the interactive effects of different long-term fire regimes (annual vs four-year fire frequency) and grazing by native ungulates ( Bos bison ) on small-scale plant community structure and resource variability (N and light) in native tallgrass prairie. Grazing enhanced light and nitrogen availability, but did not affect small-scale resource variability. In addition, grazing reduced the dominance of C₄ grasses which enhanced species richness, diversity and community heterogeneity. In contrast, annual fire increased community dominance and reduced species richness and diversity, particularly in the absence of grazing, but had no effect on community heterogeneity, resource availability and resource variability. Variability in the abundance of resources showed no relationship with community heterogeneity at the scale measured in this study, however we found a relationship between community dominance and heterogeneity. Therefore, we conclude that grazing generated small-scale community heterogeneity in this mesic grassland by directly affecting plant community dominance, rather than indirectly through changes in resource variability.     

Environmental factors influencing spatial patterns of shrub diversity in chaparral,Santa Ynez Mountains,California

We examined patterns of shrub species diversity relative to landscape‐scale variability in environmental factors within two watersheds on the coastal flank of the Santa Ynez Mountains, California. Shrub species richness and dominance was sampled at a hierarchy of spatial units using a high‐powered telescope from remote vantage points. Explanatory variables included field estimates of total canopy cover and percentage rock cover, and modeled distributions of slope, elevation, photosynthetically active radiation, topographic moisture index, and local topographic variability. Correlation, multiple regression, and regression tree analyses showed consistent relationships between field‐based measurements of species richness and dominance, and topographically‐mediated environmental variables. In general, higher richness and lower dominance occurred where environmental conditions indicated greater levels of resource limitation with respect to soil moisture and substrate availability. Maximum richness in shrub species occurred on high elevation sites with low topographic moisture index, rocky substrate, and steep slopes. Maximum dominance occurred at low elevation sites with low topographic variability, high potential solar insolation, and high total shrub canopy cover. The observed patterns are evaluated with respect to studies on species‐environment relations, resource use, and regeneration of shrubs in chaparral and coastal sage scrub. The results are discussed in the context of existing species‐diversity hypotheses that hinge on reduced competitive dominance and increased resource heterogeneity under conditions of resource limitation.     

Starting points for small mammal population recovery after wildfire: recolonisation or residual populations?

Wildfire is a major driver of spatio‐temporal variation in terrestrial ecosystems. Large wildfires are predicted to occur more frequently due to climate change. The mechanisms by which post‐fire recovery proceeds are influenced by the abundance of survivors, and their distribution in relation to habitat variability and refugia. Thus, characterising early post‐fire demographic processes is critical to understanding the demographic and community‐level responses of ecosystems to fire. The Black Saturday fires of February 2009 burnt an area of approximately 3500 km² in Victoria, Australia. We quantified the effects of this high severity forest fire on the habitat, abundance, sex ratio and body mass of two small mammal species, the agile antechinus Antechinus agilis and bush rat Rattus fuscipes. We developed a hypothetical framework to distinguish in situ survival and rapid recolonisation as the processes underlying short‐term post‐fire distributions. These hypotheses were based on expected patterns of abundance over increasing distances from unburnt sources, and the estimated recolonisation capabilities of each species and sex. The agile antechinus and bush rat were present in burnt sites at 30% and 12% of the density observed in unburnt sites. In situ survival, and not recolonisation, was the most plausible explanation for our findings. Abundance and body mass data indicated a greater effect of fire on the bush rat than the agile antechinus. The bush rat showed a shift in topographic association, whereby drainage lines acted as post‐fire refugia. Our findings suggest these species do not depend on recolonisation for recovery, and that the bush rat will follow a nucleated recovery, expanding from topographic refugia. Thus, connectivity‐reducing management activities, such as salvage logging and firebreak and road construction, may not affect the early stages of population recovery in remaining stands of burnt forest. Rather, ongoing recovery is likely to be limited by demographic rates and resource availability.     

The effect of fire interval on post‐fire understorey communities in Yellowstone National Park

Questions: How does the time interval between subsequent stand‐replacing fire events affect post‐fire understorey cover and composition following the recent event? How important is fire interval relative to broad‐ or local‐scale environmental variability in structuring post‐fire understorey communities? Location: Subalpine plateaus of Yellowstone National Park (USA) that burned in 1988. Methods: In 2000, we sampled understorey cover and Pinus contorta density in pairs of 12–yr old stands at 25 locations. In each pair, the previous fire interval was either short (7–100 yr) or long (100–395 yr). We analysed variation in understorey species richness, total cover, and cover of functional groups both between site pairs (using paired t‐tests) and across sites that experienced the short fire intervals (using regression and ordination). We regressed three principal components to assess the relative importance of disturbance and broad or local environmental variability on post‐fire understorey cover and richness. Results: Between paired plots, annuals were less abundant and fire‐intolerant species (mostly slow‐growing shrubs) were more abundant following long intervals between prior fires. However, mean total cover and richness did not vary between paired interval classes. Across a gradient of fire intervals ranging from 7–100 yr, total cover, species richness, and the cover of annuals and nitrogen‐fixing species all declined while the abundance of shrubs and fire‐intolerant species increased. The few exotics showed no response to fire interval. Across all sites, broad‐scale variability related to elevation influenced total cover and richness more than fire interval. Conclusions: Significant variation in fire intervals had only minor effects on post‐fire understorey communities following the 1988 fires in Yellowstone National Park.     

Does the Type of Disturbance Matter When Restoring Disturbance‐Dependent Grasslands?

The reintroduction of burning is usually viewed as critical for grassland restoration; but its ecological necessity is often untested. On the one hand, fire may be irreplaceable because it suppresses dominant competitors, eliminates litter, and modifies resource availability. On the other hand, its impacts could be mimicked by other disturbances such as mowing or weeding that suppress dominants but without the risks sometimes associated with burning. Using a 5‐year field experiment in a degraded oak savanna, we tested the impacts of fire, cutting and raking, and weeding on two factors critical for restoration: controlling dominant invasive grasses and increasing subordinate native flora. We manipulated the season of treatment application and used sites with different soil depths because both factors influence fire behavior. We found no significant difference among the treatments—all were similarly effective at suppressing exotics and increasing native plant growth. This occurred because light is the primary limiting resource for many native species and each treatment increased its availability. The effectiveness of disturbance for restoration depended more on the timing of application and site factors than on the type of treatment used. Summer disturbances occurred near their reproductive peak of the exotics, so their mortality approached 100%. Positive responses by native species were significantly greater on shallow soils because these areas had higher native diversity prior to treatment. Although likely not applicable to all disturbance‐dependent ecosystems, these results emphasize the importance of testing the effectiveness of alternative restoration treatments prior to their application.     

Diversity patterns from sequentially restored grasslands support the ‘environmental heterogeneity hypothesis’

The ‘environmental heterogeneity hypothesis’ (EHH) has been proposed as a mechanism that enables species coexistence through resource partitioning. In accordance with this hypothesis, plant diversity is predicted to increase with variability in resources, but there has been weak support for this hypothesis from experimental studies. The objectives of this research were to 1) characterize how resource availability and heterogeneity (coefficient of variation) change as plant communities develop using sequentially restored grasslands, 2) determine if resource heterogeneity relates to plant diversity (effective number of species, richness and evenness) and 3) reveal if the strength of resource heterogeneity–diversity relationships is different among levels of resource availability. We quantified means and coefficients of variation in soil nitrate and light availability in grasslands established on former agricultural lands for different times and their relationship to plant diversity using a geostatistically‐informed design. Nitrate availability decreased exponentially with restoration age, but no directional change in nitrate heterogeneity across the chronosequence occurred due to high resource variability in some restorations. Light availability also decreased exponentially across the chronosequence, but there was no directional change in light heterogeneity. Nitrate heterogeneity was positively correlated with both plant richness and plant effective number of species at high levels of nitrate availability. However, no nitrate heterogeneity correlation was detected at low levels of nitrate availability. Light heterogeneity was positively correlated with plant effective number of species at low levels of light availability. However, no light heterogeneity correlation was detected at high levels of light availability. Plant evenness was not correlated with resource heterogeneity at any resource availability level. These results support the positive heterogeneity–diversity relationship predicted by EHH, and uniquely that this relationship develops within a decade of plant community development, but can be obscured by resource availability.     

Contrasting responses of small mammals to fire and topographic refugia

Unburnt patches within burnt landscapes are expected to provide an important resource for fauna, potentially acting as a refuge from direct effects of fire and allowing animals to persist in burnt landscapes. Nevertheless, there is little information about the way refugia are used by fauna and how populations may be affected by them. Planned burns are often patchy, with unburnt areas generally associated with gully systems providing a good opportunity to study faunal use of refugia. We used a before–after control‐impact design associated with a planned burn in south eastern Australia to investigate how two small mammal species, the bush rat Rattus fuscipes and agile antechinus Antechinus agilis, used unburnt gully systems within a larger burnt area. We tested three alternative hypotheses relating to post‐fire abundance: (i) active refugia – abundance would increase in unburnt patches because of a post‐fire shift of individuals from burnt to unburnt areas; (ii) passive refugia – abundance in unburnt patches would remain similar to pre‐fire levels; and (iii) limited or no refugia – abundance would reduce in unburnt patches related to the change induced by fire in the wider landscape. We found the two species responded differently to the presence of unburnt refugia in the landscape. Relative to controls, fire had little effect on bush rat abundance in gullies, supporting hypothesis 2. In contrast, agile antechinus abundance increased in gullies immediately post‐fire consistent with a shift of individuals from burnt parts of the landscape, supporting hypothesis 1. Differences in site fidelity, habitat use and intraspecific competition between these species are suggested as likely factors influencing responses to refugia. The way unburnt patches function as faunal refugia and the subsequent influence they have on post‐fire population dynamics, will to some extent depend on the life history attributes of individual species.     

Negative density dependence is stronger in resource‐rich environments and diversifies communities when stronger for common but not rare species

Conspecific negative density dependence is thought to maintain diversity by limiting abundances of common species. Yet the extent to which this mechanism can explain patterns of species diversity across environmental gradients is largely unknown. We examined density‐dependent recruitment of seedlings and saplings and changes in local species diversity across a soil‐resource gradient for 38 woody‐plant species in a temperate forest. At both life stages, the strength of negative density dependence increased with resource availability, becoming relatively stronger for rare species during seedling recruitment, but stronger for common species during sapling recruitment. Moreover, negative density dependence appeared to reduce diversity when stronger for rare than common species, but increase diversity when stronger for common species. Our results suggest that negative density dependence is stronger in resource‐rich environments and can either decrease or maintain diversity depending on its relative strength among common and rare species.     

Using Dominance-Diversity Curves to Assess Completion Criteria After Bauxite Mining Rehabilitation in Western Australia

Abstract Dominance‐diversity curves have been previously constructed for a range of ecosystems around the world to illustrate the dominance of particular species and show how their relative abundances compare between communities separated in time or space. We investigate the usefulness of dominance‐diversity curves in rehabilitated areas to compare the floristic composition and abundance of “undisturbed” areas with disturbed areas, using bauxite mining rehabilitation in Western Australia as an example. Rehabilitated pits (11–13 years old) subjected to prescribed fire in autumn and spring were compared with unburned rehabilitated areas and the native jarrah (Eucalyptus marginata) forest. Dominance diversity curves were constructed by ranking the log of the species density values from highest to lowest. Species were categorized according to a variety of functional responses: life form (trees, shrubs, subshrubs, and annuals), fire response syndrome (seeder or resprouter), nitrogen fixing capability, and origin (native or adventive). Exponential functions showed extremely good fits for all sites (r² = 0.939–0.995). Dominance diversity graphs showed that after burning of rehabilitated areas, sites exhibited a more similar dominance‐diversity curve than before burning. This was emphasized in a classification (UPGMA) of the regression equations from the dominance‐diversity curves that showed that sites burned in spring were more similar to the native forest than sites burned in autumn. There was no significant segregation of the nitrogen‐fixing and species origin categories, although the life form and fire response groupings showed significant segregation along the dominance‐diversity curve. Resprouters tended to be over‐represented in the lower quartiles and under‐represented in the upper quartiles of post‐burn sites. It is suggested that using dominance‐diversity curves in the monitoring of rehabilitated areas may be a useful approach because it provides an easily interpretable visual representation of both species richness and abundance relationships and may be further utilized to emphasize categories of plants that are over‐ or under‐represented in rehabilitated areas. This will assist in the post‐rehabilitation management of these sites.     

Improving restoration success through microsite selection: an example with planting sagebrush seedlings after wildfire

Post‐fire restoration of foundation plant species, particularly non‐sprouting shrubs, is critically needed in arid and semi‐arid rangeland, but is hampered by low success. Expensive and labor‐intensive methods, including planting seedlings, can improve restoration success. Prioritizing where these more intensive methods are applied may improve restoration efficiency. Shrubs in arid and semi‐arid environments can create resource islands under their canopies that may remain after fire. Seedlings planted post‐fire in former canopy and between canopies (interspace) may have different survival and growth. We compared planting Wyoming big sagebrush (Artemisia tridentata Nutt. ssp. wyomingensis Beetle & Young) seedlings post‐fire in former sagebrush canopy and interspace microsites at five locations. Four growing seasons after planting, seedling survival was 46 and 7% in canopy and interspace microsites, respectively. Sagebrush cover was 5.8 times greater in canopy compared to interspace microsites. Sagebrush survival and cover were likely greater because of less competition from herbaceous vegetation as well as benefiting from resource island effects in canopy microsites. Initially, post‐fire abundance of exotic annual grasses was less in canopy microsites, but by the third year post‐fire it was substantially greater in canopy microsites, indicating that resource availability to seedlings was greater, at least initially, in canopy microsites. These results suggest microsites with greater likelihood of success should be identified and then utilized to improve restoration success and efficiency. This is important as the need for restoration greatly exceeds resources available for restoration.     

Life‐history traits predict perennial species response to fire in a desert ecosystem

The Mojave Desert of North America has become fire‐prone in recent decades due to invasive annual grasses that fuel wildfires following years of high rainfall. Perennial species are poorly adapted to fire in this system, and post‐fire shifts in species composition have been substantial but variable across community types. To generalize across a range of conditions, we investigated whether simple life‐history traits could predict how species responded to fire. Further, we classified species into plant functional types (PFTs) based on combinations of life‐history traits and evaluated whether these groups exhibited a consistent fire‐response. Six life‐history traits varied significantly between burned and unburned areas in short (up to 4 years) or long‐term (up to 52 years) post‐fire datasets, including growth form, lifespan, seed size, seed dispersal, height, and leaf longevity. Forbs and grasses consistently increased in abundance after fire, while cacti were reduced and woody species exhibited a variable response. Woody species were classified into three PFTs based on combinations of life‐history traits. Species in Group 1 increased in abundance after fire and were characterized by short lifespans, small, wind‐dispersed seeds, low height, and deciduous leaves. Species in Group 2 were reduced by fire and distinguished from Group 1 by longer lifespans and evergreen leaves. Group 3 species, which also decreased after fire, were characterized by long lifespans, large non‐wind dispersed seeds, and taller heights. Our results show that PFTs based on life‐history traits can reliably predict the responses of most species to fire in the Mojave Desert. Dominant, long‐lived species of this region possess a combination of traits limiting their ability to recover, presenting a clear example of how a novel disturbance regime may shift selective environmental pressures to favor alternative life‐history strategies.     

Temporal variability in detritus resource maintains diversity of bacterial communities

Competition theory generally predicts that diversity is maintained by temporal environmental fluctuations. One of the many suggested mechanisms for maintaining diversity in fluctuating environments is the gleaner-opportunist trade-off, whereby gleaner species have low threshold resource levels and low maximum growth rates in high resource concentration while opportunist species show opposite characteristics. We measured the growth rates of eight heterotrophic aquatic bacteria under different concentrations of chemically complex plant detritus resource. The growth rates revealed gleaner-opportunist trade-offs. The role of environmental variability in maintaining diversity was tested in a 28-day experiment with three different resource fluctuation regimes imposed on two four-species bacterial communities in microcosms. We recorded population densities with serial dilution plating and total biomass as turbidity. Changes in resource availability were measured from filter-sterilised medium by re-introducing the consumer species and recording short-term growth rates. The type of environmental variation had no effect on resource availability, which declined slowly during the experiment and differed in level between the communities. However, the slowly fluctuating environment had the highest Shannon diversity index, biomass, and coefficient of variation of biomass in both communities. We did not find a clear link between the gleaner-opportunist trade-off and diversity in fluctuating environments. Nevertheless, our results do not exclude this explanation and support the general view that temporal environmental variation maintains species diversity also in communities feeding chemically complex resource.     

Fire as a key driver of Earth's biodiversity

Many terrestrial ecosystems are fire prone, such that their composition and structure are largely due to their fire regime. Regions subject to regular fire have exceptionally high levels of species richness and endemism, and fire has been proposed as a major driver of their diversity, within the context of climate, resource availability and environmental heterogeneity. However, current fire‐management practices rarely take into account the ecological and evolutionary roles of fire in maintaining biodiversity. Here, we focus on the mechanisms that enable fire to act as a major ecological and evolutionary force that promotes and maintains biodiversity over numerous spatiotemporal scales. From an ecological perspective, the vegetation, topography and local weather conditions during a fire generate a landscape with spatial and temporal variation in fire‐related patches (pyrodiversity), and these produce the biotic and environmental heterogeneity that drives biodiversity across local and regional scales. There have been few empirical tests of the proposition that ‘pyrodiversity begets biodiversity’ but we show that biodiversity should peak at moderately high levels of pyrodiversity. Overall species richness is greatest immediately after fire and declines monotonically over time, with postfire successional pathways dictated by animal habitat preferences and varying lifespans among resident plants. Theory and data support the ‘intermediate disturbance hypothesis’ when mean patch species diversity is correlated with mean fire intervals. Postfire persistence, recruitment and immigration allow species with different life histories to coexist. From an evolutionary perspective, fire drives population turnover and diversification by promoting a wide range of adaptive responses to particular fire regimes. Among 39 comparisons, the number of species in 26 fire‐prone lineages is much higher than that in their non‐fire‐prone sister lineages. Fire and its byproducts may have direct mutagenic effects, producing novel genotypes that can lead to trait innovation and even speciation. A paradigm shift aimed at restoring biodiversity‐maintaining fire regimes across broad landscapes is required among the fire research and management communities. This will require ecologists and other professionals to spread the burgeoning fire‐science knowledge beyond scientific publications to the broader public, politicians and media.     

What predicts the richness of seeder and resprouter species in fire‐prone Cape fynbos: Rainfall reliability or vegetation density?

In ecosystems subject to regular canopy fires, woody species have evolved two general strategies of post‐fire regeneration. Seeder species are killed by fire and populations regenerate solely by post‐fire recruitment from a seed bank. Resprouter species survive fire and regenerate by vegetative regrowth from protected organs. Interestingly, the abundance of these strategies varies along environmental gradients and across regions. Two main hypotheses have been proposed to explain this spatial variation: the gap dependence and the environmental‐variability hypotheses. The gap‐dependence model predicts that seeders are favoured in sparse vegetation (vegetation gaps allowing effective post‐fire recruitment of seedlings), while resprouters are favoured in densely vegetated sites (seedlings being outcompeted by the rapid crown regrowth of resprouters). The environmental‐variability model predicts that seeders would prevail in reliable rainfall areas, whereas resprouters would be favoured in areas under highly variable rainfall that are prone to severe dry events (leading to high post‐fire seedling mortality). We tested these two models using distribution data, captured at the scale of quarter‐degree cells, for seeder and resprouter species of two speciose shrub genera (Aspalathus and Erica) common in fire‐prone fynbos ecosystems of the mediterranean‐climate part of the Cape Floristic Region. Contrary to the predictions of the gap‐dependence model, species number of both resprouters and seeders increased with values of the Normalized Difference Vegetation Index (a widely used surrogate for vegetation density), with a more marked increase for seeders. The predictions of the environmental‐variability hypothesis, by contrast, were not refuted by this study. Seeder and resprouter species of both genera showed highest richness in environments with high rainfall reliability. However, with decreasing reliability, seeder numbers dropped more quickly than those of resprouters. We conclude that the environmental‐variability model is better able to explain the abundance of woody seeder and resprouter species in Southern Hemisphere fire‐prone shrublands (fynbos and kwongan) than the gap‐dependence model.     

Local patterns of diversity in California northern coastal scrub

Within global biodiversity hotspots such as the California Floristic Province, local patterns of diversity must be better understood to prioritize conservation for the greatest number of species. This study investigates patterns of vascular plant diversity in relation to coast–inland environmental gradients in the shrublands of Central California known as northern coastal scrub. We sampled coastal shrublands of the San Francisco Bay Area at coastal and inland locations, modeled fine‐scale climatic variables, and developed an index for local exposure to maritime salts. We compared diversity, composition, and structure of the coastal and inland plots using indirect gradient analysis and estimated species accumulation using rarefaction curves. Coastal plots were significantly higher in alpha, beta, and gamma diversity than inland plots. Plant diversity (effective species number) in coastal plots was 2.1 times greater than inland plots, and beta diversity was 1.9 times greater. Estimated richness by rarefaction was 2.05 times greater in coastal sites than inland sites. Salt deposition and water availability were the abiotic process most strongly correlated with increased maritime plant diversity and compositional differences. Stands of northern coastal scrub on the immediate coast with higher maritime influence exhibit markedly higher plant diversity than most interior stands, paralleling previous work in other vegetation types in this region. These studies suggest that the California coastline deserves special consideration for botanical conservation. Fine‐scale climatic models of cloud frequency, water availability, and the salt deposition index presented here can be used to define priority areas for plant conservation in California and other coastal regions worldwide.     

Distribution of plant life history types in California chaparral: the role of topographically‐determined drought severity

Abstract. Spatial patterns of shrub life history and Ceanothus distribution are examined in relation to topographically‐mediated differences in drought severity within 3 watersheds on the coastal and inland flank of the Santa Ynez Mountains, California. Spatially distributed fields of drought severity are simulated for the studied watersheds using high‐resolution digital terrain data and daily climate data in combination with a process‐based hydro‐ecological model (RHESSys). Field samples of species composition are spatially integrated with the distributed drought data for analysis of ecological relationships. Patterns of seedling recruitment type correspond to topographic variability in drought severity in ways that are consistent with concepts presented in the literature. Species that depend on fire for recruitment are increasingly represented with increasing drought severity, the converse also applies. Sites that experience moderate drought severity permit co‐dominance of species from both recruitment modes. Residual analysis suggests that some of the unexplained variability is related to substrate. Analyses also indicate that the distribution of 5 Ceanothus shrubs reflect differences in drought severity in ways that are consistent with their resistance to water stress‐induced xylem dysfunction. Species from the subgenus Cerastes sort in accordance with moisture availability and have unique spatial distributions. Results are evaluated and discussed with respect to studies on plant morphology, resource use and seedling establishment patterns.     

Ecological niche and phylogeny explain distribution of seed mass in the central European flora

Seed size is a crucial life‐history trait determining the amount of reserves that are available to establishing seedlings. The most frequently observed patterns in seed size distribution are a higher frequency of large‐seeded species in shaded habitats and a positive correlation of seed size with plant size. We analysed to what extent realised niche dimensions, as expressed by Ellenberg indicator values and plant functional traits such as plant height and life form, explained seed mass variation in the central European flora. By including information on phylogenetic relatedness of the species, not only contemporary ecology but also the evolutionary history of plant species could be taken into account. Seed mass evolution was slow and was best explained by selection‐inertia models with multiple adaptive peaks as a function of either habitat or life form. The highest seed mass optima were observed in the deciduous forest and saltwater and seashore habitats, and in phanerophytes in case of models with optima as a function of life form. The analyses showed that Ellenberg values were more important than habitat and life form in explaining seed mass distribution in the central European flora. The often observed relation between shade and large seeds was also evident in our study, but we found an equally important relation between large seeds and drought and a positive relation between seed mass and salinity. Our results indicate that not only plant size and competition for light but a complex set of factors influence the ecology of seed size, and that a more precise delineation of species’ niches improves the understanding of seed size evolution.     

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.     

Plant functional traits along environmental gradients in seasonally dry and fire‐prone ecosystem

Questions: How does the abundance and richness of plant assemblages with different functional (regeneration and nutrient acquisition) traits vary with fire regime, moisture availability and substrate fertility? What is the role of different functional traits in maintaining plant diversity under changing environmental conditions in seasonally dry and fire‐prone environments? Location: Southwest Western Australia. Methods: Plant species abundance and soil nutrients were determined at 16 forest sites with variable fire histories across an aridity gradient. All plant species were classified based on their functional traits as (1) perennial or annual, (2) ectomycorrhizal, arbuscular mycorrhizal, ericoid mycorrhizal, orchid mycorrhizal, proteoid or other non‐mycorrhizal, (3) resprouters or seeder, and (4) nitrogen fixer or non‐fixer. We used a multivariate (fourth‐corner) technique to simultaneously test the significance and direction of the relationship between each of these traits and fire frequency, fire interval length, aridity, and soil N, P and C fractions. Results: The functional response of the vegetation to fire regime was minor and restricted to annual species, which comprised only ～4% of taxa. Proteoid and ectomycorrhizal species dominated over species with arbuscular and orchid mycorrhizal roots, N‐fixers dominated over non‐fixers, and seeders dominated over resprouters when N fertility was low but organic labile P was high. Further, proteoid and ectomycorrhizal species richness increased with aridity, while arbuscular mycorrhizal species richness decreased. Conclusions: While the functional composition of southwest Australian vegetation is largely insensitive to changes in fire regime, nutrient acquisition and, to a lesser extent, regeneration traits provide mechanisms for the vegetation community to adjust to changes in resource availability. Thus, diversity responses to environmental change in seasonally dry and fire‐prone ecosystems are likely to be primarily mediated by the composition of nutrient acquisition traits in the vegetation community.     

Influence of fire severity on stand development of Araucaria araucana–Nothofagus pumilio stands in the Andean cordillera of south‐central Chile

Fire is the prevalent disturbance in the Araucaria–Nothofagus forested landscape in south‐central Chile. Although both surface and stand‐replacing fires are known to characterize these ecosystems, the variability of fire severity in shaping forest structure has not previously been investigated in Araucaria–Nothofagus forests. Age structures of 16 stands, in which the ages of approximately 650 trees were determined, indicate that variability in fire severity and frequency is key to explaining the mosaic of forest patches across the Araucaria–Nothofagus landscape. High levels of tree mortality in moderate‐ to high‐severity fires followed by new establishment of Nothofagus pumilio typically result in stands characterized by one or two cohorts of this species. Large Araucaria trees are highly resistant to fire, and this species typically survives moderate‐ to high‐severity fires either as dispersed individuals or as small groups of multi‐aged trees. Small post‐fire cohorts of Araucaria may establish, depending on seed availability and the effects of subsequent fires. Araucaria's great longevity (often >700 years) and resistance to fire allow some individuals to survive fires that kill and then trigger new Nothofagus cohorts. Even in relatively mesic habitats, where fires are less frequent, the oldest Araucaria–Nothofagus pumilio stands originated after high‐severity fires. Overall, stand development patterns of subalpine Araucaria–N. pumilio forests are largely controlled by moderate‐ to high‐severity fires, and therefore tree regeneration dynamics is strongly dominated by a catastrophic regeneration mode.