Fuels and landscape flammability in an Australian alpine environment

Factors governing landscape‐scale flammability are poorly understood, yet critical to managing fire regimes. Studies of the extent and severity of the 2003 Australian alpine fires revealed marked differences in flammability between major alpine plant communities, with the occurrence and severity of fire greater in heathland compared to grassland. To understand this spatial variation in landscape flammability, we documented variation in two physical properties of fuel – load and bulk density – at the life‐form and plant community scale. We measured the load (mass per unit area) and bulk density (mass per unit volume) of fine fuels (<6 mm) at 56 sites across the Bogong High Plains, southeastern Australia. Fine fuel load was positively correlated with shrub cover, and fine fuel bulk density was negatively correlated with shrub cover. Furthermore, fine fuel load and bulk density were accurately predicted using simple measures of canopy height and shrub cover. We also conducted a burning experiment on individual shrubs and snowgrass (Poa spp.) patches to assess comparative differences in flammability between these life‐forms. The burning experiment revealed that shrubs were more flammable than snowgrass as measured by a range of flammability variables. Consequently, our results indicate that treeless alpine landscapes of southeastern Australia are differentially flammable because of inherent life‐form differences in both fine fuel load and bulk density. If shrub cover increases in these alpine landscapes, as projected under climate change, then they are likely to become more flammable and may experience more frequent and/or severe fires.     

Removal of invasive shrubs alters light but not leaf litter inputs in a deciduous forest understory

The establishment and spread of non‐native, invasive shrubs in forests poses an important obstacle to natural resource conservation and management. This study assesses the impacts of the physical removal of a complex of woody invasive shrub species on deciduous forest understory resources. We compared leaf litter quantity and quality and understory light transmittance in five pairs of invaded and removal plots in an oak‐dominated suburban mature forest. Removal plots were cleared of all non‐native invasive shrubs. The invasive shrubs were abundant (143,456 stems/ha) and diverse, dominated by species in the genera Ligustrum, Viburnum, Lonicera, and Euonymus. Annual leaf litter biomass and carbon inputs of invaded plots were not different from removal plots due to low leaf litter biomass of invasive shrubs. Invasive shrub litter had higher nitrogen (N) concentrations than native species; however, low biomass of invasive litter led to low N inputs by litter of invasive species compared to native. Light transmittance at the forest floor and at 2 m was lower in invaded plots than in removal plots. We conclude that the removal of the abundant invasive shrubs from a native deciduous forest understory did not alter litter quantity or N inputs, one measure of litter quality, and increased forest understory light availability. More light in the forest understory could facilitate the restoration of forest understory dynamics.     

Shrubs as ecosystem engineers in a coastal dune: influences on plant populations,communities and ecosystems

Question: How do two shrubs with contrasting life‐history characteristics influence abundance of dominant plant taxa, species richness and aboveground biomass of grasses and forbs, litter accumulation, nitrogen pools and mineralization rates? How are these shrubs – and thus their effects on populations, communities and ecosystems – distributed spatially across the landscape? Location: Coastal hind‐dune system, Bodega Head, northern California. Methods: In each of 4 years, we compared vegetation, leaf litter and soil nitrogen under canopies of two native shrubs –Ericameria ericoides and the nitrogen‐fixing Lupinus chamissonis– with those in adjacent open dunes. Results: At the population level, density and cover of the native forb Claytonia perfoliata and the exotic grass Bromus diandrus were higher under shrubs than in shrub‐free areas, whereas they were lower under shrubs for the exotic grass Vulpia bromoides. In contrast, cover of three native moss species was highest under Ericameria and equally low under Lupinus and shrub‐free areas. At community level, species richness and aboveground biomass of herbaceous dicots was lower beneath shrubs, whereas no pattern emerged for grasses. At ecosystem level, areas beneath shrubs accumulated more leaf litter and had larger pools of soil ammonium and nitrate. Rates of nitrate mineralization were higher under Lupinus, followed by Ericameria and then open dune. At landscape level, the two shrubs – and their distinctive vegetation and soils – frequently had uniform spatial distributions, and the distance separating neighbouring shrubs increased as their combined sizes increased. Conclusions: Collectively, these data suggest that both shrubs serve as ecosystem engineers in this coastal dune, having influences at multiple levels of biological organization. Our data also suggest that intraspecific competition influenced the spatial distributions of these shrubs and thus altered the distribution of their effects throughout the landscape.     

Effects of Prescribed Burning on Herbaceous and Woody Vegetation in Northern Lowland Meadows

Abstract Spring burning of sedge‐grass meadows in the Slave River Lowlands (SRL), Northwest Territories, Canada was applied between 1992 and 1998 to reduce shrub encroachment and enhance Bison bison (bison) habitat, although the impact of fire on preferred bison forage was unknown before management. In the summer of 1998 we conducted a study in the Hook Lake area of the SRL to test the effect of burn frequency (unburned, burned once, or burned three times since 1992) on herbaceous plant community composition and Salix spp. L. (willow) shrub vigor. Plant species abundance, litter biomass, soil pH, and depth of the organic soil horizon were measured in 300 1‐m² quadrats nested within 30 1,000‐m² plots in both burned and unburned dry meadows. To test the relationship between frequency and willow vigor, all willow shrubs within the plots were assigned a vigor score from I (dead) to IV (flourishing). The spring burns appear to have reduced willow vigor; however, shrub survival remained high (76%) on the most frequently burned meadows. Ordination plots resulting from canonical correspondence analysis suggest that multiple spring burns influenced plant community composition in dry meadow areas and that less palatable bison forage species (e.g., Carex aenea Fern. and Juncus balticus L.) were correlated with a regime of three spring burns. Our results suggest that frequent spring fires in the Hook Lake area have only a small negative effect on willow cover but may reduce the abundance of primary bison forage plants compared with less frequently burned meadows.     

Fire frequency influences composition and structure of the shrub layer in an Australian subcoastal temperate grassy woodland

Little is known about the relationship between fire regimes and plant diversity in Australia's temperate grassy woodlands. The effect of fire frequency on shrubs in grassy woodland remnants across Western Sydney's Cumberland Plain was examined. Shrub species richness and composition were compared in sites that had experienced a high, moderate or low frequency of fire over the previous 20 years. Nine sites were surveyed, three in each fire frequency category; most sites, including all low‐fire‐frequency sites, had burnt 9–36 months prior to sampling. Fire frequency had a profound effect on the composition and structure of the shrub layer. Per cent frequency and density of the prickly shrub Bursaria spinosa (Pittosporaceae) was considerably higher in low‐fire‐frequency sites than where fires had occurred at least once a decade. In sites where fire had been absent for decades prior to a recent fire, this species dominated the landscape, while elsewhere it occurred as clumps in a grassy matrix. Per cent frequency of other native shrubs, particularly obligate seeders, was greatest at moderate fire frequencies. Exotic shrubs were recorded most often where fire had been rare. While ordination clearly separated out the low‐fire‐frequency sites, complete separation between high‐ and moderate‐fire‐frequency blocks was not achieved. The increase in Bursaria in the absence of fire mirrors the encroachment of woody plants into a range of grassy ecosystems around the world. The sensitivity of obligate seeder species, many of them short‐lived legumes with fire‐cued seeds, to both very frequent and very infrequent fire shows the vulnerability of these species to extreme fire regimes, despite the safeguards conferred by hard‐seededness. Competition from Bursaria, as well as loss of viable seed in the soil, may have contributed to the low frequency of these species after a long inter‐fire interval.     

Short‐term effects of herbicides and a prescribed fire on restoration of a shrub‐encroached pine savanna

Shrub encroachment occurring worldwide in savannas and grasslands has commonly been hypothesized to result from anthropogenically altered environments. Two disturbance‐based approaches to restoration have involved: (1) application of selective herbicides to reduce density/cover of shrubs; (2) reinstatement of natural fire regimes to generate environmental conditions favoring herbaceous species. We studied short‐term responses of native shrubs, vines, and grasses in a Louisiana pine savanna to herbicides followed by a prescribed fire and fire alone. Plots established in the summer, 2013, were hand‐sprayed in the fall with Imazapyr and Triclopyr, Triclopyr alone, or no herbicide, then prescribed burned the following spring. Numbers of species of shrubs and vines at scales of 1 and 100 m², numbers of stems and regrowth of stems produced by six common species of shrubs, and the number of flowering culms of perennial C₄ grasses were assessed postfire in 2014. Compared with fire alone, herbicides followed by fire resulted in (1) small reductions in species richness of shrubs and no effects on vines, (2) fewer stems comprising shrub genets, but similar postfire regrowth of resprouting shrub stems, and (3) fewer flowering culms of C₄ grasses. Underground storage organs of savanna shrubs and vines survived both aboveground disturbances. Thus, single applications of herbicides followed by fires reduced, but did not reverse shrub encroachment and negatively affected grasses. Because effects of herbicides overrode those of prescribed fires, these disturbances did not act synergistically, suggesting that reinstating natural fire regimes should be a priority in restoration of savannas and grasslands.     

Abrupt fire regime change may cause landscape‐wide loss of mature obligate seeder forests

Obligate seeder trees requiring high‐severity fires to regenerate may be vulnerable to population collapse if fire frequency increases abruptly. We tested this proposition using a long‐lived obligate seeding forest tree, alpine ash (Eucalyptus delegatensis), in the Australian Alps. Since 2002, 85% of the Alps bioregion has been burnt by several very large fires, tracking the regional trend of more frequent extreme fire weather. High‐severity fires removed 25% of aboveground tree biomass, and switched fuel arrays from low loads of herbaceous and litter fuels to high loads of flammable shrubs and juvenile trees, priming regenerating stands for subsequent fires. Single high‐severity fires caused adult mortality and triggered mass regeneration, but a second fire in quick succession killed 97% of the regenerating alpine ash. Our results indicate that without interventions to reduce fire severity, interactions between flammability of regenerating stands and increased extreme fire weather will eliminate much of the remaining mature alpine ash forest.     

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.     

Fire season affects size and architecture of Colophospermum mopane in southern African savannas

Wildfires may be started naturally by lightning or artificially by humans. In the savanna regions of southern Africa, lightning fires tend to occur at the start of the wet season, during October and November, while anthropogenic fires are usually started during the dry season, between July and August. A long-term field manipulation experiment initiated in the Kruger National Park in 1952 was used to explore whether this seasonal divergence affects tree abundance, spatial pattern, size and architecture. After 44 years of prescribed burning treatments that simulated the seasonal incidence of lightning and anthropogenic fires, mean densities of the locally-dominant shrub, Colophospermum mopane, were 638 and 500 trees ha^–1 respectively. Trees in burnt plots had aggregated distributions while trees in unburnt plots had random distributions. Significant differences (p < 0.001) were recorded in a range of morphological parameters including tree height, canopy diameter, mean stem circumference and number of stems. The incidence of resprouting also differed significantly between treatments, with burnt trees containing a high proportion of coppiced stems. The differences in tree size and architecture between the mid-dry season and early-wet season burning plots suggest that anthropogenic fires applied during July and August cannot substitute for a natural lightning fire regime. Anthropogenic fire yields a landscape that is shorter, more scrubby and populated by numerous coppiced shrubs than the landscape generated by natural lightning fire conditions.     

Vegetation responses in Alaskan arctic tundra after 8 years of a summer warming and winter snow manipulation experiment

We used snow fences and small (1 m²) open‐topped fiberglass chambers (OTCs) to study the effects of changes in winter snow cover and summer air temperatures on arctic tundra. In 1994, two 60 m long, 2.8 m high snow fences, one in moist and the other in dry tundra, were erected at Toolik Lake, Alaska. OTCs paired with unwarmed plots, were placed along each experimental snow gradient and in control areas adjacent to the snowdrifts. After 8 years, the vegetation of the two sites, including that in control plots, had changed significantly. At both sites, the cover of shrubs, live vegetation, and litter, together with canopy height, had all increased, while lichen cover and diversity had decreased. At the moist site, bryophytes decreased in cover, while an increase in graminoids was almost entirely because of the response of the sedge Eriophorum vaginatum. These community changes were consistent with results found in studies of responses to warming and increased nutrient availability in the Arctic. However, during the time period of the experiment, summer temperature did not increase, but summer precipitation increased by 28%. The snow addition treatment affected species abundance, canopy height, and diversity, whereas the summer warming treatment had few measurable effects on vegetation. The interannual temperature fluctuation was considerably larger than the temperature increases within OTCs (<2°C), however. Snow addition also had a greater effect on microclimate by insulating vegetation from winter wind and temperature extremes, modifying winter soil temperatures, and increasing spring run‐off. Most increases in shrub cover and canopy height occurred in the medium snow‐depth zone (0.5–2 m) of the moist site, and the medium to deep snow‐depth zone (2–3 m) of the dry site. At the moist tundra site, deciduous shrubs, particularly Betula nana, increased in cover, while evergreen shrubs decreased. These differential responses were likely because of the larger production to biomass ratio in deciduous shrubs, combined with their more flexible growth response under changing environmental conditions. At the dry site, where deciduous shrubs were a minor part of the vegetation, evergreen shrubs increased in both cover and canopy height. These changes in abundance of functional groups are expected to affect most ecological processes, particularly the rate of litter decomposition, nutrient cycling, and both soil carbon and nitrogen pools. Also, changes in canopy structure, associated with increases in shrub abundance, are expected to alter the summer energy balance by increasing net radiation and evapotranspiration, thus altering soil moisture regimes.     

Native herbivore exerts contrasting effects on fire regime and vegetation structure

Although native herbivores can alter fire regimes by consuming herbaceous vegetation that serves as fine fuel and, less commonly, accumulating fuel as nest material and other structures, simultaneous considerations of contrasting effects of herbivores on fire have scarcely been addressed. We proposed that a colonial rodent, vizcacha (Lagostomus maximus), reduces and increases fire intensity at different stages in its population cycle in the semiarid scrub of Argentina. Specifically, we hypothesized that, when colonies are active, vizcachas create natural fire-breaks through intense grazing, generating over time patches of large unburned shrubs in grazed zones. In contrast, when colonies are abandoned, recovery of fine fuels and previous accumulation of coarse wood on colonies during territorial displays increases fire intensity, creating patches of high shrub mortality. To test these hypotheses, we estimated stem age of the dominant shrub (Larrea divaricata) and measured aboveground biomass in zones actively grazed by vizcachas and in ungrazed zones, and compared densities of live and dead shrubs on abandoned colonies and adjacent zones following fire. In active colonies, age and biomass of shrubs were much greater in grazed than ungrazed zones. In abandoned colonies that had been burnt, density of dead, burned shrubs was higher and density of live shrubs was lower than in adjacent zones. These results support our hypotheses and reveal a new interaction between native herbivores and fire, in which herbivores augment fire intensity by gathering fuel. Our findings indicate that, through opposing effects on fire, native herbivores enhance the heterogeneity of vegetation in woody-dominated ecosystems.     

Effects of re‐introducing fire to a central Florida sandhill community

Abstract. A southern ridge sandhill site in central Florida, USA, was burned in 1989, 1991, and 1995 after 63 years of fire‐suppression to simulate a pre‐settlement fire regime. Fire changed species abundance and vegetation structure but caused only minimal changes in species turnover and diversity. There was a general trend for an increase in the cover of herbs following fire but this was a statistically significant effect for only one species, Liatris tenuifolia var. tenuifolia. Aristida beyrichiana increased, litter cover and litter depth were significantly reduced, and ground lichens were eliminated in response to burning. Scrub oaks and palmettos in the ground cover and small shrub layers (height ≤ 1 m) either increased or did not respond to burning, reflecting strong post‐fire resprouting. Diversity in the ground cover and small shrub layers were not affected by fire. Scrub oaks and palmettos in the large shrub and overstorey layers (height > 1 m) were reduced in density, basal area, and longest canopy measurements in response to fire. Species diversity also decreased within these layers following fire. Some Pinus elliottii var. densa survived fire, but their density was reduced. All Pinus clausa were eliminated by fire. Periodic burning can suppress the dominance of shrubs (Quercus spp.) while increasing the cover of grasses and herbs in southern ridge sandhill vegetation.     

Vegetation shift from deciduous to evergreen dwarf shrubs in response to selective herbivory offsets carbon losses: evidence from 19 years of warming and simulated herbivory in the subarctic tundra

Selective herbivory of palatable plant species provides a competitive advantage for unpalatable plant species, which often have slow growth rates and produce slowly decomposable litter. We hypothesized that through a shift in the vegetation community from palatable, deciduous dwarf shrubs to unpalatable, evergreen dwarf shrubs, selective herbivory may counteract the increased shrub abundance that is otherwise found in tundra ecosystems, in turn interacting with the responses of ecosystem carbon (C) stocks and CO₂ balance to climatic warming. We tested this hypothesis in a 19‐year field experiment with factorial treatments of warming and simulated herbivory on the dominant deciduous dwarf shrub Vaccinium myrtillus. Warming was associated with a significantly increased vegetation abundance, with the strongest effect on deciduous dwarf shrubs, resulting in greater rates of both gross ecosystem production (GEP) and ecosystem respiration (ER) as well as increased C stocks. Simulated herbivory increased the abundance of evergreen dwarf shrubs, most importantly Empetrum nigrum ssp. hermaphroditum, which led to a recent shift in the dominant vegetation from deciduous to evergreen dwarf shrubs. Simulated herbivory caused no effect on GEP and ER or the total ecosystem C stocks, indicating that the vegetation shift counteracted the herbivore‐induced C loss from the system. A larger proportion of the total ecosystem C stock was found aboveground, rather than belowground, in plots treated with simulated herbivory. We conclude that by providing a competitive advantage to unpalatable plant species with slow growth rates and long life spans, selective herbivory may promote aboveground C stocks in a warming tundra ecosystem and, through this mechanism, counteract C losses that result from plant biomass consumption.     

Dendroecological potential of shrubs for reconstructing fire history at landscape scale in Mediterranean-type climate grasslands: The case of Fabiana imbricata

Fire recurrently affects Mediterranean-type climate (MTC) regions causing major implications on the structure and dynamics of vegetation. In these regions, it is important to know the fire regime for which reliable fire records are needed. Dendroecology offers the possibility of obtaining fire occurrence data from woody species and has been widely used in forest ecosystems for fire research. Grasslands are regions with no trees where shrubs can provide dendroecological evidence for reconstructing fire history at landscape scale. We studied the dendroecological potential of the shrub Fabiana imbricata to reconstruct fire history at landscape scale in MTC grasslands of northwestern Patagonia. In order to accomplish this, we combined spatio-temporal information of recorded fires from the study area with the age structure of F. imbricata shrublands obtained from dendroecological methods. Shrubland age structure correctly described how often fires occurred in the past. In rocky outcrops, where fires cannot reach, individuals are long-lived and heterogeneous in age; while downhill, individuals are young and shrublands are even-aged. Five pulses of massive recruitment were found: three of these coincided with three known fires; the remaining two had not been recorded before. A bi-variated analysis showed that F. imbricata recruited mainly during two years after fire, and the spatial distribution of pulses coincided with the fire map. Information derived from shrubland age structure could be used to estimate fire regime parameters such as fire return interval at landscape or community scale. For instance, we estimated a fire return interval of nine years at landscape scale and ranging from 11 to 24 years at community scale (shrubland). Our results in northwestern Patagonia grasslands showed that the F. imbricata chronology can be used to complement other information sources such as remote sensing and operational databases improving the knowledge about fire regime. The present study demonstrates that is possible to utilize shrubs as a dendroecological data source to study fire history in regions where tree cover is absent.     

Small mammal responses to pyric successional changes in eucalypt forest

Sixteen 1-ha study plots covering five regeneration stages were simultaneously trapped five times over a 20-month period to provide data on small mammal response to vegetation changes following fire. Areas regenerating after fires from 9 years to 1 month before the investigation were sampled in a uniform open forest on a coastal sand plain. Two types of understorey were recognized: one dominated by true forest shrubs with which Rattus fuscipes, Antechinus stuartii and Sminthopsis murina were associated, and another dominated by heath elements where the addition of Pseudomys novaehollandiae and Mus musculus produced a significantly more diverse small mammal community. The two communities exhibited different responses to post-fire vegetation changes. Rattus fuscipes was the most abundant species and showed a logistic growth in biomass. No resident populations were established in the first 3 years, but a rapid increase in biomass occurred from 3 to 5 years to plateau after 8 years. Regeneration age had the greatest effect on R. fuscipes biomass mediated through the amount of accumulated leaf litter with additional variation being attributed to several vegetation structure variables and plant species diversity. A replacement sequence in time was observed for species reaching their maximum abundance (P. novaehollandiae and/or M. musculus → S. murina → A. stuartii → R. fuscipes) and was interpreted as species occupying stages in the succession when their optimal habitat requirements were fulfilled. These results have important implications for the design of management policies using fire or fire regimes as tools for habitat maintenance or alteration. A mosaic of forest patches of adequate size covering the entire range of seral stages is necessary to meet the optimum requirements of all the above species.     

Altered Fire Regimes Affect Landscape Patterns of Plant Succession in the Foothills and Mountains of Southern California

In the southern California foothills and mountains, pronounced and complex topographic gradients support fire regimes that vary over short distances. We used LANDIS, a spatially explicit landscape model of disturbance and plant succession, to examine the resilience of dominant plant species, representing different disturbance response strategies, to the effect of varying fire rotation intervals (FRI). The simulated fire regimes represented natural, current and very long FRIs for the foothill shrublands less than 1,400 m (90, 30 and 150 years) and montane forest greater than 1400 m (30, 150, 500 years). The 30-year FRI allowed obligate resprouting shrubs to dominate over obligate seeders, whereas the 90-year FRI resulted in a stable spatial distribution of both of these shrub functional types. This is consistent with the literature that suggests that shifts in shrubland composition are most likely to result from human-caused increases in fire frequency at the low-elevation urban-wildland interface. An ecotone conifer, Pinus coulteri, showed dramatic shifts in distribution under different FRIs, and retreated to the portion of the landscape representing its temporal regeneration niche. Both low and high frequency fire maintained the fire tolerant dominant pine (P. jeffreyi) in the montane zone. This contradicts the literature that suggests that a high frequency ground fire regime is required for the persistence of a pine-dominated forest, but is consistent with studies showing that conifer forests in the western U.S. have experienced, and are resilient to, a broad range of natural FRIs that include low frequency, high intensity crown fires.     

Effects of fire season and soil fertility on clonal growth in a pyrophilic forb,Pityopsis graminifolia (Asteraceae)

We experimentally manipulated fire season in a longleaf pine sandhill community in north Florida in 1990 and 1992. We determined the size and demography of shoots, rhizomes, and clones of Pityopsis graminifolia from late 1990 to early 1993. In addition, we examined shoot responses to seasonal variation in canopy/litter removal and soil fertility in 1992. May fires caused a higher short-term rate of increase in shoot density than did either January fires or August fires. The combination of canopy/litter removal and mineral nutrient addition as part of the clipping experiment revealed a similar effect on shoot density in May. Fire season influenced clone structure. May fires resulted in greater numbers of shoots/clone than did January or August fires. January-burned plots had clones and rhizomes that contained larger (but fewer) shoots than did May-burned or August-burned plots. We suggest that such variation in clone structure may indicate a greater capacity of clones to grow laterally following May fires than following either January or August fires. This capacity is positively associated with the likelihood that longleaf pine savannas will be burned by lightning fires at different times during the year.     

Interactive effects of grazing and shrubs on the annual plant community in semi‐arid Mediterranean shrublands

Question: We studied the interactive effects of grazing and dwarf shrub cover on the structure of a highly diverse annual plant community. Location: Mediterranean, semi‐arid shrubland in the Northern Negev desert, Israel. Methods: Variation in the biomass and plant density of annual species in the shrub and open patches was monitored during four years, inside and outside exclosures protected from sheep grazing, in two contrasting topographic sites: north and south‐facing slopes that differed in their dominant dwarf shrubs species: Sarcopoterium spinosus and Corydothymus capitatus, respectively. Results: Above‐ground biomass, density and richness of annual species were lower under the canopy of both shrub species compared to the adjacent open patches in the absence of grazing. Grazing reduced the biomass of annuals in open patches of both topographic sites, but not in the shrub patches. On the north‐facing slope, grazing also reduced plant density and richness in the open patches, but increased plant density in the shrub patches. At the species level, various response patterns to the combined effects of grazing and patch type were exhibited by different annuals. Protection against the direct impacts of grazing by shrub cover as well as species‐specific interactions between shrubs and annuals were observed. A conceptual mechanistic model explaining these interactions is proposed. Conclusion: In semi‐arid Mediterranean shrublands grazing and dwarf shrub cover interact in shaping the structure of the annual plant community through (1) direct impacts of grazing restricted to the open patches, (2) species‐specific facilitation/ interference occurring in the shrub patches and (3) subsequent further processes occurring among the interconnected shrub and open patches mediated through variation in seed flows between patches.     

Impacts of changed fire regimes on tropical riparian vegetation invaded by an exotic vine

Abstract Riparian habitats are highly important ecosystems for tropical biodiversity, and highly threatened ecosystems through changing disturbance regimes and weed invasion. An experimental study was conducted to assess the ecosystem impacts of fire regimes introduced for the removal of the exotic woody vine, Cryptostegia grandiflora, in tropical north‐eastern Australian woodlands. Experimental sites in subcatchments of the Burdekin River, northern Queensland, Australia, were subjected to combinations of early wet‐season and dry‐season fires, and single and repeated fires, with an unburnt control. Woody vegetation was sampled using permanent quadrats to record and monitor plants species, number and size‐class. Sampling was conducted pre‐fire in 1999 and post‐fire in 2002. All fire regimes were effective in reducing the number and biomass of C. grandiflora shrubs and vines. Few woodland or riparian species were found to be fire‐sensitive and community composition did not change markedly under any fire regime. The more intense dry‐season fires impacted the structure of non‐target vegetation, with large reductions in the number of sapling trees (<5 cm d.b.h.) and reductions in the largest tree size‐class and total tree basal area. Unexpectedly, medium‐sized canopy trees (10–30 cm d.b.h.) appear to have been significantly benefited by fires, with decreases in number of trees of this size‐class in the absence of fire. Although the presence of C. grandiflora as a vine in riparian forest canopies changed the nature and intensity of crown combustion patterns, this did not lead to the initiation of a self‐perpetuating weed–fire cycle, as invaders were unable to take advantage of gaps caused by fire. Low intensity, early wet‐season burning, or early dry‐season burning, is recommended for control of C. grandiflora in order to minimize the fire intensity and risk of the loss of large habitat trees in riparian habitats.