Survival and life expectancy of the tree Protea roupelliae subsp. roupelliae in a montane grassland savanna: Effects of fire regime and plant structure

Survival and life expectancy are key demographic determinants of population dynamics. Using data collected in a field experiment monitored over 14 years in montane grassland of the Ukhahlamba‐Drakensberg Park, South Africa, we determined the effects of components of fire regime and plant structure on the survival and life expectancy of the tree Protea roupelliae subsp. roupelliae (Proteaceae). The field experiment comprised six plots (0.2–0.5 ha in area) from which the survival and life expectancies of 1567 juveniles (non‐reproductives) and 329 adults (reproductives) were estimated in response to differences in fire frequency, biennial seasonal fire, flame height, juvenile height, adult height, basal area and canopy vigour. Juvenile survival and life expectancies were highest when fires were excluded for 8 years. However, a fire after 12 years of fire exclusion and another fire 2 years later eliminated all juveniles. Over the same 14‐year period of biennial fires, juvenile survival was 5%. Juvenile survival and life expectancy were higher after biennial, winter fires than after annual, winter fires. Flame height had no effect on juvenile survival and life expectancy. Both survival and life expectancy of juveniles increased as plants got older and grew taller. Adult survival was unaffected by fire frequency, flame height or tree size, but the survival of adults in response to fire seasonality was inconclusive. Adults with low canopy vigour (<25%) were negatively affected by fire. Juvenile survival and life expectancy are critical bottlenecks in the demography of P. roupelliae. This species is neither a reseeder nor a resprouter. It avoids lethal fire damage by being restricted to rocky habitats with low fire intensities. Biennial winter fires least threaten the survival and life expectancy of P. roupelliae and impact least on its role in the summer feeding and breeding of Gurney's sugarbird.     

Fire interval effects on persistence of resprouter species in Mediterranean-type shrublands

The capacity of many plant species to resprout in fire-prone shrublands is thought to engender persistence, yet management concerns exist for the long-term persistence of some resprouting species given anthropogenic impacts including shortened fire intervals, long periods of fire exclusion, and/or fires of increasingly high severity. We explored the potential demographic effects of different fire interval regimes on lignotuberous resprouter species using the last fire interval for 36 sites (33 experimental fires, 3 wildfires) in biodiverse SW Australian shrublands, spanning an interval range of 3–42 years. Mortality and regrowth 1 year following the last fire was assessed for >7,000 tagged individuals from 20 shrub and sub-shrub species. Using generalized linear mixed effect models, we estimated the influence of fire interval (and selected fire and environmental covariates) on mortality and regrowth rates across all species, and individually for the four most common species. The overall model, as well as the models for three of the four most common species (Banksia attenuata, Melaleuca leuropoma, and M. systena, but not Hibbertia hypericoides) supported the hypothesis of increased mortality at short and long fire intervals, most likely due to total non-structural carbohydrate (TNC) and bud-bank limitation, respectively. However, no relationship between regrowth rate and fire interval was detected, suggesting that increased mortality at short (3–5 year) fire intervals may not be due solely to resource (TNC) limitation. Results show that lignotuberous resprouters are potentially vulnerable to population decline through attrition of mature plants under both shortened and lengthened fire interval regimes.     

Demography of woody plants in relation to fire: Banksia ericifolia L.f. and Petrophile pulchella (Schrad) R.Br.

Plants of Banksia ericifolia and Petrophile pulchella are sensitive to fire. Changes in population size under different fire regimes were estimated, based on measurements of post-fire seedling emergence, seedling survival, survival and seed production in established plants of differing ages, survival of seeds held in serotinous cones and seed-release in the periods between fire. Seeds were first available at 5 years in P. pulchella and 6 years in B. ericifolia. Exact replacement would be possible when burnt at these ages, if seedling establishment were very high. Low establishment would delay replacement to 13 years of age in both species. Late summer/autumn fires of high intensity favour high establishment. Such fires at 8–10 year intervals would be tolerated without any sustained decline in numbers. Fires at 10–15 year intervals could occur regardless of season or intensity with little risk of a population decline. Large increases in numbers and density would follow fires spaced at 15–30 years. Enough seeds would be available for replacement up to about 50 years in both species. Viable seed-release in unburnt conditions was sufficient to compensate for deaths in stands over 20 years old, even with very low levels of establishment. Two variables accounted for the biggest changes in numbers and density between generations interspersed by fires; namely the age at which a stand is burned and the proportion of seeds which emerge as seedlings. A comparison with other similar species showed similarities in controls on emergence and establishment, lengths of primary juvenile periods and life spans.     

The impact of experimental fire regimes on seed production in two tropical eucalypt species in northern Australia

Abstract This study investigated the effect of three experimental fire regimes on the fecundity, ovule development and seedfall of two common wet-dry tropical savanna eucalypts, Eucalyptus minima and Eucalyptus tetrodonta, in northern Australia. Both species flower early in the dry season and ovule development occurs during the dry season. This coincides with a period of frequent fires. The three fire regimes considered were applied for four years between 1990 and 1994. These regimes were (i) Unburnt, (ii) Early, fires lit early in the dry season, and (iii) Late, fires lit late in the dry season. The treatments were applied to nine catchments (15–20 km²) with each fire regime replicated three times. Fire intensity typically increases as the dry season proceeds. Therefore, early dry season fires generally differ from late dry season fires in both their intensity and their timing in relation to the reproductive phenology of the eucalypts. Late dry season burning significantly reduced the fecundity of both species, whereas Early burning had no significant effect. Ovule success was significantly reduced by the Early burning for both species. The Late burning significantly reduced ovule success in E. tetrodonta, but not in E. miniata. The results suggest that fire intensity and fire timing may both be important determinants of seed supply. Fire intensity may be a determinant of fecundity, whereas fire timing in relation to the reproduction phenology may have a significant impact on ovule survival. Both fire regimes resulted in a substantial reduction in seed supply compared with the Unburnt treatment. This may have a significant impact on seedling regeneration of these tropical savanna eucalypts.     

The effect of fire and ants on the seed-bank of a shrub in a semi-arid grassland

Abstract Invasion of grasslands by woody plants following the introduction of domestic stock is a worldwide phenomenon. Burning is frequently recommended as a remedial measure but for a pastoral enterprise it is costly and the frequency of the fires required is of critical economic importance. The size and longevity of the soil seed-bank is an essential part of the response of shrub populations to prescribed fire regimes. In this study the seed-bank of the semi-arid zone shrub Dodonaea attenuata in Eragrostis eriopoda tussock grassland was examined in relation to harvesting by ants and the burning history of the sites. On unburnt sites, more than 3500 seeds per m2 entered the seed-bank in the summer 1984–85 but sites burnt 5 years previously produced less than one-third of that number. Burnt shrubs did not flower for 5 years and no seeds survived in the soil through to 1985. Burning immediately prior to seed ripening destroyed the seed crop but burning after seed-fall stimulated a greater germination in the following spring than on unburnt treatments. Ants rapidly harvested most of the seeds produced and after 20 months the combined effects of ant harvesting and germination had reduced the seed-bank at unburnt sites to 8–21 seeds per m2 and at sites burnt 7 years previously to less than two seeds per m2. Seeds were initially harvested in summer by Pheidole spp. of ants for their elaiosome and then discarded in middens outside the entrance to the ant nest. During the subsequent cool season the seeds were taken back into the nest and stored at depths ranging from 2 to 30cm. It was concluded that ants provided short-range dispersal (< 10 m) and promoted the contagious distribution of D. attenuata, which is advantageous for a fire-susceptible, arid-zone shrub invading a grassland liable to be burnt: seedlings derived from seeds in ant storage chambers near the soil surface and in sparse grass situations caused by competition from shrubs may obtain some survival advantage. Prescribed fire has potential as a management tool for controlling population density of D. attenuata because, depending upon season and frequency, it reduces seed rain by killing shrubs, suppresses flowering activity and destroys seed crops on the plant. Under the influence of a regime of regular burning, such as prevailed prior to the European pastoral industry, the limited soil seed-bank would have been a major constraint on D. attenuata populations.     

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.     

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.     

Seeds of the annual grasses Schizachyrium spp. as a food resource for tropical granivorous birds

Seed dynamics of the annual tropical grasses Schizachyrium fragile (R. Br.) A. Camus, S. pachyarthron C. Gardner and S. pseudeulalia (Hosok.) S.T. Blake were studied with the aim of documenting fluxes in granivore food resources. In S. fragile, seed production began in the early dry season, and seed output was primarily influenced by seedling survival. Following seed-fall, there were 651 S. fragile seeds/m² (393 kernels/m²⁾ and 1014 S. pachyarthron seeds/m² (593 kernels/m²) across the study area, with a combined kernel biomass of 14.1 × 10³ g/ha. Seed densities remained stable through the dry season, then declined rapidly after wet season rain. Initial wet season rains of up to 25 mm, 40 mm and 50 mm resulted in seed-bank germination of < 5%, 57%, and 93%, respectively. Some seeds were buried by rain and resurfaced or germinated later, but about 30% was lost. Widespread and abrupt depletion of the seed-bank is likely to occur at the start of about 8% of wet seasons on central Cape York Peninsula, leaving little seed, either for subsequent germination or as food for granivores. Burning early in the dry season, when most seeds were still on the plants, reduced seed densities by 85%. The proportion of seeds with sound kernels was reduced in areas burnt by dry season fires, both directly after the fires and, subsequently, as a result of preferential granivore activity. Areas burnt in the dry season were thus depleted of seed earlier than were unburnt areas. Food availability for granivores should therefore be optimized by fire regimes that include a range of burning histories, including fires in both early dry and early wet seasons as well as keeping other areas unburnt.     

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.     

Relationship between leaf traits and fire-response strategies in shrub species of a mountainous region of south-eastern Australia

Background and Aims

Resprouting and seed recruitment are important ways in which plants respond to fire. However, the investments a plant makes into ensuring the success of post-fire resprouting or seedling recruitment can result in trade-offs that are manifested in a range of co-occurring morphological, life history and physiological traits. Relationships between fire-response strategies and other traits have been widely examined in fire-prone Mediterranean-type climates. In this paper, we aim to determine whether shrubs growing in a non-Mediterranean climate region exhibit relationships between their fire-response strategy and leaf traits.

Methods

Field surveys were used to classify species into fire-response types. We then compared specific leaf area, leaf dry-matter content, leaf width, leaf nitrogen and carbon to nitrogen ratios between (a) obligate seeders and all other resprouters, and (b) obligate seeders, facultative resprouters and obligate resprouters.

Key Results

Leaf traits only varied between fire-response types when we considered facultative resprouters as a separate group to obligate resprouters, as observed after a large landscape-scale fire. We found no differences between obligate seeders and obligate resprouters, nor between obligate seeders and resprouters considered as one group.

Conclusions

The results suggest that facultative resprouters may require a strategy of rapid resource acquisition and fast growth in order to compete with species that either resprout, or recruit from seed. However, the overall lack of difference between obligate seeders and obligate resprouters suggests that environmental factors are exerting similar effects on species'' ecological strategies, irrespective of the constraints and trade-offs that may be associated with obligate seeding and obligate resprouting. These results highlight the limits to trait co-occurrences across different ecosystems and the difficulty in identifying global-scale relationships amongst traits.     

Fire resilience of a rare,freshwater crustacean in a fire‐prone ecosystem and the implications for fire management

Understanding species’ responses to fire regimes, particularly rare or threatened species, is important for land managers tasked with managing for biodiversity. Hickman's Allanaspides (Allanaspides hickmani, Anaspidesidae) is a rare, primitive, shrimp‐like crustacean, with high conservation value. It is restricted to a single catchment in the island state of Tasmania, Australia, where it occurs within moorland pools typically containing crayfish (Ombrastacoides spp.) burrows. Although its moorland habitat has a long history of firing, adverse fire regimes are a potential threat to the species. A large part of its range is subject to planned burning to help manage the detrimental effects of high‐intensity wildfires. The resilience of A. hickmani to low–moderate‐intensity fires was investigated over 13 years using a replicated before‐after‐control‐impact design. The fires resulted in an initial reduction in vegetation cover and surface water and an increase in water temperature. There was no effect of fire on A. hickmani captures 4 months after small‐scale, low‐intensity autumn burns. However, 5 months later, following an unintended larger‐scale, medium‐intensity spring burn, there was an 80–90% reduction in A. hickmani captures and their numbers did not recover until 6–9 years post‐fire. It is not known whether the reduced catch was due to a reduction in the number of A. hickmani or their movement from pools into crayfish burrows. These findings together with evidence of a varied fire history, including high‐intensity wildfires, within their range suggests that A. hickmani and its habitat are resilient to a range of fire frequencies and intensities provided that the fire regime does not degrade or lead to a complete loss of peat. Climate change predictions for warmer and drier summers in western Tasmania will increase the risk of peat loss. Planned burning is likely to be important for the protection of A. hickmani habitat from predicted adverse fire regimes.     

Tropical and north Pacific teleconnections influence fire regimes in pine-dominated forests of north-eastern California, USA

Aim To assess the importance of drought and teleconnections from the tropical and north Pacific Ocean on historical fire regimes and vegetation dynamics in north‐eastern California. Location The 700 km² study area was on the leeward slope of the southern Cascade Mountains in north‐eastern California. Open forests of ponderosa pine (Pinus ponderosa var. ponderosa Laws.) and Jeffrey pine (P. jeffreyi Grev. & Balf) surround a network of grass and shrub‐dominated meadows that range in elevation from 1650 to 1750 m. Methods Fire regime characteristics (return interval, season and extent) were determined from crossdated fire scars and were compared with tree‐ring based reconstructions of precipitation and temperature and teleconnections for the period 1700–1849. The effect of drought on fire regimes was determined using a tree‐ring based proxy of climate from five published chronologies. The number of forest‐meadow units that burned was compared with published reconstructions of the El Niño/Southern Oscillation (ENSO) and the Pacific Decadal Oscillation (PDO). Results Landscape scale fires burned every 7–49 years in meadow‐edge forests and were influenced by variation in drought, the PDO and ENSO. These widespread fires burned during years that were dryer and warmer than normal that followed wetter and cooler years. Less widespread fires were not associated with this wet, then dry climate pattern. Widespread fires occurred during El Niño years, but fire extent was mediated by the phase of the PDO. Fires were most widespread when the PDO was in a warm or normal phase. Fire return intervals, season and extent varied at decadal to multi‐decadal time scales. In particular, an anomalously cool, wet period during the early 1800s resulted in widespread fires that occurred earlier in the year than fires before or after. Main conclusions Fire regimes in north‐eastern California were strongly influenced by regional and hemispheric‐scale climate variation. Fire regimes responded to variation that occurred in both the north and tropical Pacific. Near normal modes of the PDO may influence fire regimes more than extreme conditions. The prevalence of widespread teleconnection‐driven fires in the historic record suggests that variation in the Pacific Ocean was a key regulator of fire regimes through its influence on local fuel production and successional dynamics in north‐eastern California.     

Fire frequency has a contrasting effect on vegetation and topsoil in subcoastal heathland,woodland and forest ecosystems,south-east Queensland,Australia

Ecosystems managed with contrasting fire regimes provide insight into the responses of vegetation and soil. Heathland, woodland and forest ecosystems along a gradient of resource availability were burnt over four decades in approximately 3- or 5-year intervals or were unburnt for 45–47 years (heathland, woodland), or experienced infrequent wildfires (forest: 14 years since the last fire). We hypothesized that, relative to unburnt or infrequent fires, frequent burning would favour herbaceous species over woody species and resprouting over obligate seeder species, and reduce understorey vegetation height, and topsoil carbon and nitrogen content. Our hypothesis was partially supported in that herbaceous plant density was higher in frequently burnt vegetation; however, woody plant density was also higher in frequently burnt areas relative to unburnt/infrequently burnt areas, across all ecosystems. In heathland, omission of frequent fire resulted in the dominance of fern Gleichenia dicarpa and subsequent competitive exclusion of understorey species and lower species diversity. As hypothesized, frequent burning in woodland and forest increased the density of facultative resprouters and significantly reduced soil organic carbon levels relative to unburnt sites. Our findings confirm that regular burning conserves understorey diversity and maintains an understorey of lower statured herbaceous plants, although demonstrates the potential trade-off of frequent burning with lower topsoil carbon levels in the woodland and forest. Some ecosystem specific responses to varied fire frequencies were observed, reflecting differences in species composition and fire response traits between ecosystems. Overall, unburnt vegetation resulted in the dominance of some species over others and the different vegetation types were able to withstand relatively high-frequency fire without the loss of biodiversity, mainly due to high environmental productivity and short juvenile periods.     

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.     

Population dynamics of the shrub Acacia suaveolens (Sm.) Willd.: Survivorship throughout the life cycle,a synthesis

Survivorship in Acacia suaveolens was assessed through seedling and adult stages. Moisture stress was found to be the critical factor limiting early seedling survival. Both seedling and adult populations were characterized by periods of low mortality interspersed with pulses of high mortality. A composite survivorship curve for A. suaveolens based on nine sites predicts that some 20–25 years after afire, established plants should disappear from the above-ground flora if another fire does not occur. Fecundity and survivorship data were used to estimate the flux of seed in the soil over time in a hypothetical A. suaveolens population. From this it was predicted that, following establishment of plants after a fire, the seed-bank would rapidly reach a maximum after 6 years and thereafter slowly decline, until after 60 years there would be only as many seeds as there were original parental plants. The situation would vary with predispersal seed predation, seed predation on the soil surface, seed dispersal by ants to ‘unsafe sites’ and the size of the initial seed-bank prior to establishment. Only after a very long inter-fire period would A. suaveolens be eliminated from a site. Elimination of the species is also possible under very frequent fires. A 2–5 year fire-free period is needed for plants to reach maturity and another 6 years are needed to maximize seed input into the soil seed-bank. In addition, seedling recruitment following cool burns is low to non-existent as dormancy is not broken for most seeds in the soil during such burns.     

Determinants for the conservation of a vulnerable fire-dependent species at its marginal range

We investigated the role of fire and other potential biotic and abiotic determinants for the occurrence of an obligate seeder and active pyrophyte, Cistus salviifolius, a vulnerable species in the Swiss Red List. Populations of this species are threatened by the change of fire regime that has occurred during the last ten years. Fires and burnt areas are today less frequent and it is not clear whether other factors can compensate for such fire-free periods should this trend continue. We used a stratified random design to collect data on the cover of three growth stages, i.e., juvenile, chamaephytic and nanophanerophytic (oldest), of C. salviifolius, on plant communities, and on environmental factors. A tree-based discriminant analysis showed that the time elapsed since last fire was the best predictor of Cistus occurrence, but plants could survive on rocky and sunny sites if no fire had occurred for more than 32 years. Contrary to our expectations, the number of fires and time elapsed since the last event was correlated with the oldest stages only, and not with that of seedlings or juveniles. Rank correlations showed that bare soil was a good predictor of young stages, whereas rocky outcrops and shallow soils were important for the chamaephytic stage. Our results confirm the role of disturbances and competition-free habitats as determinants of the survival of this vulnerable pyrophyte during long periods without fires and the existence of ontogenetic niche segregation of this species. In order to maintain viable C. salviifolius populations in the Alps, managers should undertake conservation actions according to the requirements of the different growth stages of this species.     

Effects of fire and drought in a tropical eucalypt savanna colonized by rain forest

Aim This study documents the effects of multiple fires and drought on the woody structure of a north Australian savanna never grazed by domestic stock. Location The study was conducted in a 500 ha pocket of Eucalyptus‐dominated savanna surrounded by a late Quaternary lava flow. The flow is known as the Great Basalt Wall, located c. 50 km northeast of Charters Towers in semi‐arid north‐eastern Australia. This region was exposed to the largest 5‐year rainfall deficit on record between 1992 and 1996. Methods All individual woody plants were tagged within a 1.56 ha plot. Species were segregated into their habitat affinities (rain forest, ecotone, savanna) and regeneration strategy (resprouter, seeder). The survivorship of plants within these categories was analysed in relation to fire intensity from the first fire, and to each of four fires lit between 1996 and 2001. Results Before the first fire, the plot contained thirty‐one tree species including twenty‐one typical of the surrounding dry rain forest. These rain forest species were represented by small individuals and constituted <1% of the total basal area of woody plants. The basal area of savanna trees was 7.5 m² ha^?1 at the commencement of monitoring, although 31% had recently died and others had major crown damage. Further death of the drought debilitated savanna trees was substantial during the first year of monitoring and the basal area of live savanna trees declined to 1.1 m² ha^?1 after 5 years. Most species from both rain forest and savanna were classified as resprouters and are capable of regenerating from underground organs after fire. Species without this ability (rain forest seeders and ecotone seeders) were mostly eliminated after the first two consecutive fires. Among resprouters, survivorship declined as fire intensity increased and this was more pronounced for rain forest than for savanna species. Repeated burning produced a cumulative effect of decreasing survivorship for rain forest resprouters relative to savanna resprouters. Main conclusions The study provides evidence that savanna and rain forest trees differ in fire susceptibility and that recurrent fire can explain the restricted distribution of rain forest in the seasonally arid Australian tropics. The time of death of the savanna trees is consistent with the regional pattern after severe drought, and highlights the importance of medium term climate cycles for the population dynamics of savanna tree species and structure of Australian savannas.     

Influence of canopy openness,ungulate exclosure,and low‐intensity fire for improved oak regeneration in temperate Europe

Failed oak regeneration is widely reported in temperate forests and has been linked in part to changed disturbance regimes and land‐use. We investigated if the North American fire–oak hypothesis could be applicable to temperate European oaks (Quercus robur, Quercus petraea) using a replicated field experiment with contrasting canopy openness, protection against ungulate browsing (fencing/no fencing), and low‐intensity surface fire (burn/no burn). Survival, relative height growth (RGR_H), browsing damage on naturally regenerated oaks (≤300 cm tall), and changes in competing woody vegetation were monitored over three years. Greater light availability in canopy gaps increased oak RGR_H (p = .034) and tended to increase survival (p = .092). There was also a trend that protection from browsing positively affected RGR_H (p = .058) and survival (p = .059). Burning reduced survival (p < .001), nonetheless, survival rates were relatively high across treatment combinations at the end of the experiment (54%–92%). Most oaks receiving fire were top‐killed and survived by producing new sprouts; therefore, RGR_H in burned plots became strongly negative the first year. Thereafter, RGR_H was greater in burned plots (p = .002). Burning altered the patterns of ungulate browsing frequency on oaks. Overall, browsing frequency was greater during winter; however, in recently burned plots summer browsing was prominent. Burning did not change relative density of oaks, but it had a clear effect on competing woody vegetation as it reduced the number of individuals (p < .001) and their heights (p < .001). Our results suggest that young, temperate European oaks may respond similarly to fire as their North American congeners. However, disturbance from a single low‐intensity fire may not be sufficient to ensure a persistent competitive advantage—multiple fires and canopy thinning to increase light availability may be needed. Further research investigating long‐term fire effects on oaks of various ages, species‐specific response of competitors and implications for biodiversity conservation is needed.     

Robustness of demographic estimates in studies of plant responses to fire

Abstract For demographic models of resprouting species in relation to fire regimes, spatially separated populations of different ages are usually treated as a chronosequence, and the average behaviour of these populations is used as the estimate of the generalized behaviour of a single population through time. Using the data of Bradstock (1990), we examine the way in which the between-population variability affects the demographic parameters for Banksia serrata (Proteaceae) and thus assess whether the estimates of the shortest inter-fire interval necessary to maintain a stable population size under low and high intensity fires are robust to variation in these parameters. The range of our estimates compares well to the average estimate of Bradstock (1990) for high intensity fires, but not for low intensity fires. However the relevant demographic parameters appear to vary more between populations than they do as a result of the length of the inter-fire intervals, and the specific demographic model discussed here may have only local application.