Effects of fire frequency on plant species composition of sandstone communities in the Sydney region: Inter-fire interval and time-since-fire

Abstract Fire frequency is the number of fires experienced by a particular community within a given time period. This concept can potentially be resolved into a number of interacting variables, including: time since the most recent fire, the length of the inter-fire intervals, and the variability of the length of the inter-fire intervals. We estimated the effects of these three variables on the floristic composition of 65 samples from dry sclerophyll vegetation with different fire histories in Brisbane Water, Ku-ring-gai Chase and Royal National Parks near Sydney. Our analyses suggest that fire frequency may account for about 60% of the floristic variation among our samples. They confirm the hypothesis that the recent (<30 years) fire frequency produces effects on floristic composition of fire-prone communities that can recognizably be attributed both to the time since the most recent fire and to the length of the intervals between fires. These effects are equal in magnitude but are different in the nature of the floristic variation they are associated with. Increasing time-since-fire is associated with a decline in the evenness of fire-tolerant species, indicating that fewer of these species come to dominate the community in the prolonged absence of fire. Herbs and small shrubs decrease in abundance, while larger shrubs increase in abundance. Inter-fire intervals of decreasing length are associated with a decrease in the evenness of the fire-sensitive species, particularly those large Proteaceae shrubs that often dominate the community biomass in dry sclerophyll shrublands of southeastern Australia. Furthermore, the variation associated with inter-fire intervals is not necessarily solely related to the shortest inter-fire interval, but is related to combinations of inter-fire intervals through time. Thus, increasing variability of the length of the inter-fire intervals is associated with an increase in the species richness of both fire-sensitive and fire-tolerant species, implying that it may be variation of the inter-fire intervals through time that is primarily responsible for maintaining the presence of a wide variety of plant species in a particular community. Our results also suggest that the floristic variation associated with different inter-fire intervals decreases with increasing time-since-fire.     

Effects of fire intensity on plant species composition of sandstone communities in the Sydney region

Abstract Fire intensity measures the heat output of a fire, and variation in fire intensity has been shown to have many effects on the demography of plant species, although the consequent effects on the floristic composition of communities have rarely been quantified. The effects of variation in fire intensity on the floristic composition of dry sclerophyll vegetation with different fire histories near Sydney was estimated. In particular, differences in species abundance of woodland and shrubland communities subjected to four fire‐intensity classes: unburnt, low intensity (<500 kW m^?1), medium intensity (500–2500 kW m^?1) and high intensity (>2500 kW m^?1) were examined. The samples had a standardized previous fire frequency and season, thus minimizing the effects of other aspects of the fire regime. There was a clear effect of fire intensity on the relative abundances of the vascular plant species, with increasing intensity of the fire producing vegetation that was increasingly different from the unburnt vegetation. This pattern was repeated in both the woodland and shrubland vegetation types, suggesting that it was not an artefact of the experimental conditions. However, the effects of fire intensity on floristic composition were no greater than were the differences between these two similar vegetation types, with variation in fire intensity accounting for only approximately 10% of the floristic variation. Nevertheless, the effects of fire intensity on the abundance of individual species were consistent across taxonomic groups, with the monocotyledon and Fabaceae species being more abundant at higher than lower intensities, the Proteaceae and Rutaceae more abundant at intermediate intensities, and the Epacridaceae more abundant at lower rather than higher intensities. The number of fire‐tolerant species increased with increasing fire intensity, and those fire‐tolerant species present were most abundant in the areas burnt with medium intensity. The number of fire‐sensitive species did not respond to fire intensity, and those species present were most abundant in the areas burnt with low intensity. This suggests that either fire‐sensitive species respond poorly to higher fire intensities or fire‐tolerant species respond poorly to lower fire intensities, perhaps because of differences in seed germination, seedling survival or competition among adults.     

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

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

The influence of fire frequency on arbuscular mycorrhizal colonization in the shrub Dillwynia retorta (Wendland) Druce (Fabaceae)

 Fire regimes have three inter-related components that can affect population dynamics: frequency, intensity and season. However, there has been little effort to study the effects of any of these components on arbuscular mycorrhizas (AM). In order to examine the long-term effects of fire frequency on AM colonization, the roots of Dillwynia retorta were examined at 32 sites supporting Hawkesbury Sandstone vegetation in the Sydney region of southeastern Australia. These sites were representative of the broad-scale variability in fire frequencies with respect to the length and timing of inter-fire intervals found in the Sydney region during the previous 30 years. The length of the shortest inter-fire interval was significantly correlated with total AM colonization, and the length of the longest inter-fire interval was related to the arbuscular colonization. The length of the most-recent inter-fire interval and the time since the shortest inter-fire interval were not related to AM colonization in D. retorta. Furthermore, AM colonization was directly related to the local abundance of the host plant, indicating that the effects of fire frequency on AM colonization are likely to occur indirectly via direct effects on the host plant. Canonical correspondence analysis demonstrated that the presence and abundance of alternate potential hosts had no influence on mycotrophy in D. retorta. Thus, the impact of fire on D. retorta was probably the main factor influencing its mycorrhizal status in relation to fire history. Accepted: 16 October 1998     

Carbohydrate storage in five resprouting Florida scrub plants across a fire chronosequence

Most research analyzing nonstructural carbohydrate (NSC) concentrations on resprouter species in fire-controlled ecosystems has concentrated on how NSC concentrations recover immediately after fire. However, we know little of the effect of long periods without fire on NSC concentrations. In order to assess the effect of different periods of time-since-fire on resprouter species, we studied carbohydrate concentrations (total [NSC], soluble sugars [SS] and nonsoluble sugars [NSS]) in five resprouting species with contrasting trends of abundance across a chronosequence of time-since-fire (0.5-40 yr) in Florida. Carbohydrate concentrations were highest in species with specialized reserve organs. [SS] was mainly explained by factors related to plant size, whereas time-since-fire was the main factor explaining [NSS]. Changes in [NSS] and [NSC] were correlated with the time-since-fire abundance patterns. Variation in [NSS] carbohydrates can be related to the structural development of vegetation, with only those species capable of accessing full light able to accumulate carbohydrates, whereas subordinate plants show reductions in the [NSS] carbohydrate fractions. In areas with long intervals between fires, this carbohydrate reduction could affect subsequent postfire resprouting vigour, although this remains to be confirmed.     

The size,distribution and germination requirements of the soil-stored seed-bank of Grevillea barklyana (Proteaceae)

Abstract Fire-triggered release from seed dormancy is a characteristic of many Australian plant species. We investigated aspects of the seed-bank dynamics and dormancy characteristics in seeds of Grevillea barklyana, an understorey shrub of coastal sclerophyll vegetation in the Jervis Bay Region on the south coast of New South Wales. We used two soil core sizes to compare the number and distribution of stored seeds in soil cores taken from underneath and outside the limits of plant canopies at three study sites. Core size did not affect the estimate of seed density. No seeds were found outside the existing canopies. Even in under-canopy samples, seed numbers were small. Mean seed densities were estimated as 10. 9, 14. 1 and 4. 3 seeds per for the three sites. A hierarchical series of laboratory experiments was used to test the germination response of both fresh and soil-stored seeds. Likewise, we attempted to simulate ‘natural’ disturbance conditions in a glasshouse experiment. Both laboratory and glasshouse experiments indicated polymorphism in germination behaviour. A constant proportion of seeds exhibited enforced dormancy, when moist at room temperature, while a smaller number of seeds showed either an induced dormancy or a non-seed-coat linked innate dormancy. The majority (75%) of seeds were innately dormant due to a hard seed-coat. This dormancy was broken when the seed-coat was damaged, for instance, by heat. The level of polymorphic germination behaviour will be dependent on the length of the inter-fire periods. We conclude that the expression of polymorphism within and between species across a range of environments and fire regimes is an important consideration for any further study attempting to assess the role of the seed-bank.     

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.     

Fire, soil fertility and delayed seed release: a community analysis of the degree of serotiny

Delayed seed release (serotiny) is a convergent plant trait in fire-prone regions of the world but explaining the degree of serotiny has remained elusive because of the paucity of community data. Selective forces involving seed predators, fire and soil nutrients have been suggested as factors influencing serotiny. We tested whether protection of seeds and/or synchronized dispersal were associated with different levels of serotiny and if resprouting ability influences selection for strong serotiny. We compared the numbers and abundance of 146 woody species with delayed dispersal among five community types varying in combinations of fire severity, fire frequency, soil fertility and seed predators. The strength of the relationship between levels of serotiny and environmental factors was tested among community types ranging from rainforests to heathlands. Highest levels of serotiny were recorded in low nutrient shrublands with intermediate fire return intervals that burn at high severity, while the lowest were recorded in high nutrient, low flammability forests. Both protection of seeds and synchronized seed release were related to fire effects in nutrient-limited environments. Strong serotiny is prominent in species killed by fire whereas weak serotiny is more common in resprouting species. Recruitment failure in the inter-fire interval appears to drive selection for strong maternal care of seeds and synchronized seed dispersal in fire-prone environments. Weak serotiny is proposed as a bet-hedging strategy that relies on resprouting after fire for population persistence and higher probability of inter-fire recruitment. The spectrum of serotiny (weak to strong) in these communities is proposed to be driven by the interactive effect of both fire and soil nutrients on the selection for delayed seed dispersal.     

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.     

Vertebrate-dispersed species in a fire-prone environment

Abstract It is hypothesized that plant species that produce vertebrate-dispersed seeds (fleshy fruits and brightly coloured arillate seeds) would not be common in fire-prone vegetation because seeds are deposited on the soil surface and are unlikely to survive fires. They have not previously been known to have any means of burial that would enable them to await the next fire as a buried soil seed-bank. We studied vertebrate-dispersed species in fire-prone sclerophyll vegetation to look at mechanisms used by vertebrate-dispersed plants for persisting in these environments. Seeds do not survive heating to 150°C, although species from fire-prone environments are more likely to tolerate moderate heating (80°C) without affecting viability than vertebrate-dispersed species from environments where fire is rare. Most vertebrate-dispersed species have the capacity to regenerate vegetatively, although they are not more likely to have this strategy than ant-dispersed species. Finally, ants do take vertebrate-dispersed seeds and fruits (particularly small ones) indicating the potential for the formation of a soil seed-bank. Some species only persist in fire-prone environments by recolonizing from unburnt areas.     

Fire frequency and time‐since‐fire effects on the open‐forest and woodland flora of Girraween National Park,south‐east Queensland,Australia

Abstract The effects of recent fire frequency and time‐since‐fire on plant community composition and species abundance in open‐forest and woodland vegetation in Girraween National Park, south‐east Queensland, Australia, were examined. Cover‐abundance data were collected for shrub and vine species in at least 10 400‐m² plots in each of four study areas. Study areas were within one community type and had burnt most recently either 4 or 9 years previously. Variations in fire frequency allowed us to compare areas that had burnt at least three times in the previous 25 years with less frequently burnt areas, and also woodlands that had experienced a 28‐year interfire interval with more frequently burnt areas. Although species richness did not differ significantly with either time‐since‐fire or fire frequency, both these factors affected community composition, fire frequency being the more powerful. Moisture availability also influenced floristics. Of the 67 species found in five or more plots, six were significantly associated with time‐since‐fire, whereas 11 showed a significant difference between more and less frequently burnt plots in each of the two fire‐frequency variables. Most species, however, did not vary in cover‐abundance with the fire regime parameters examined. Even those species that showed a marked drop in cover‐abundance when exposed to a particular fire regime generally maintained some presence in the community. Five species with the capacity to resprout after fire were considered potentially at risk of local extinction under regimes of frequent fire, whereas two species were relatively uncommon in long‐unburnt areas. Variable fire regimes, which include interfire intervals of at least 15 years, could be necessary for the continuity of all species in the community.     

Variation in bradyspory and seedling recruitment without fire among populations of Banksia serrata (Proteaceae)

Abstract Long-term canopy storage of seeds in fruits (bradyspory) varies among species and is thought to be related to the fire regime experienced by the plant community. This study sought to quantify intraspecific variation in bradyspory among nine populations of Banksia serrata (Proteaceae) on the south coast of New South Wales. Marked differences were found between populations, ranging from more than 30% of follicles open at two coastal sites, to less than 5% of follicles open at two inland heath sites. Low levels of bradyspory occurred particularly in coastal sites which also had Banksia integrifolia, a non-bradysporous species, present. In addition to spontaneous follicle opening without fire, fall of infructes-cences to the ground after breaking of branches was observed to be a major contributor to seed release. At three sites where fire had not occurred for many years, the amount of seedling establishment since the last fire corresponded well with levels of bradyspory; there were no inter-fire recruits at the site with the strongest bradyspory, while at the site with the weakest bradyspory there were numerous inter-fire plants, some with lignotubers large enough to be able to tolerate the next fire. It is argued that populations of B. serrata exhibit variable patterns of recruitment, depending on the site characteristics, ranging from complete dependence on fire for population maintenance to bet-hedging (spreading potential recruitment between fire events and inter-fire periods). Variations in populations and reproductive characteristics among sites make the application of demographic models to management decisions habitat-specific.     

Contrasting changes in vegetation structure and diversity with time since fire in two Australian Mediterranean‐climate plant communities

Mallee‐heath and mallee communities occur in a mosaic across large areas of south‐western Australia, in topographically subdued and fire‐prone landscapes. Consequently, it could be expected that these communities would have historically experienced similar fire regimes, and would respond similarly to variation in aspects of the fire regime. We studied the response of mallee‐heath and mallee to time since the last fire, measuring species density, species–area relationships, diversity indices and vegetation structure. Floristic responses to time since fire accorded with the initial floristic composition model of plant succession, with declining species density and Shannon diversity with age in mallee‐heath. Mallee‐heath exhibited structural senescence when > approximately 45–55 years since fire, with increasing standing dead vegetation, bare ground and stagnating or declining size in sprouting Eucalyptus spp. Mallee showed no such evidence of senescence, and indeed continued to increase in stature beyond the mean fire interval but without the compositional change required to provide support for the relay floristic model of plant succession. These results indicate that mallee‐heath is a fire maintained community and as such is reliant upon periodic burning to maintain diversity and vigour. Mallee, in contrast, is modified but not maintained by fire (at least over the period of time since fire examined) and hence is less susceptible to fire interval effects. Indeed, structural attributes likely to be significant for fauna habitat and carbon sequestration continue to develop in mallee unburnt for 55 years or more. Different responses to time since fire will create challenges for management, particularly in fragmented landscapes where fire potentially interacts with other threatening processes.     

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

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

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.     

Effects of fire regime on plant species richness and composition differ among forest,woodland and heath vegetation

Question

Do the effects of fire regimes on plant species richness and composition differ among floristically similar vegetation types?

Location

Booderee National Park, south‐eastern Australia.

Methods

We completed floristic surveys of 87 sites in Sydney Coastal dry sclerophyll vegetation, where fire history records have been maintained for over 55 years. We tested for associations between different aspects of the recent fire history and plant species richness and composition, and whether these relationships were consistent among structurally defined forest, woodland and heath vegetation types.

Results

The relationship between fire regime variables and plant species richness and composition differed among vegetation types, despite the three vegetation types having similar species pools. Fire frequency was positively related to species richness in woodland, negatively related to species richness in heath, and unrelated to species richness in forest. These different relationships were explained by differences in the associations between fire history and species traits among vegetation types. The negative relationship between fire frequency and species richness in heath vegetation was underpinned by reduced occurrence of resprouting species at high fire frequency sites (more than four fires in 55 years). However, in forest and woodland vegetation, resprouting species were not negatively associated with fire frequency.

Conclusions

We hypothesize that differing relationships among vegetation types were underpinned by differences in fire behaviour, and/or biotic and abiotic conditions, leading to differences in plant species mortality and post‐fire recovery among vegetation types. Our findings suggest that even when there is a high proportion of shared species between vegetation types, fires can have very different effects on vegetation communities, depending on the structural vegetation type. Both research and management of fire regimes may therefore benefit from considering vegetation types as separate management units.     

Demography of woody plants in relation to fire: Banksia serrata Lf. and Isopogon anemonifolius (Salisb.) Knight

Banksia serrata and Isopogon anemonifolius are serotinous resprouters (single-stemmed tree, multi-stemmed shrub, respectively) found in forests within the Sydney region. Studies were conducted to predict the population dynamics of these species. Seed production and survival and the accumulation of seed-bank within cones were estimated in relation to time since fire. Emergence, survival and development of lignotubers were measured in young juveniles and the time taken to reach adulthood was estimated. This information and published data on survival were used to estimate the amount of recruitment of adults and juveniles necessary for stands to remain in a stable state under frequent (<16 years), high and low intensity fire regimes. The effects of longer intervals (up to 80 years) between fires were also estimated. It was predicted that B. serrata populations will decline in numbers when the interval between high intensity fires is <9 years, while under low intensity fires the critical interval was 12–13 years. In I. anemonifolius the predicted intervals were 14 and 16 years, respectively. When fires are timed so that maximum seed-bank is available (about 30 year interval), it is unlikely that resprouters will dominate communities because the seed-banks and rates of growth of seedlings of obligate seeder shrubs are greater than these resprouters. Populations of these resprouters may be more able to persist than obligate seeders when the fire frequency is either very high (<6 years) or low (>50years), though the density of resprouter populations may slowly decline under such fire regimes.     

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.     

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

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

Patterns of seed longevity and dormancy in obligate seeding legumes of box‐ironbark forests,south‐eastern Australia

Current fuel loads and distribution suggest that fire events are infrequent and of a low intensity in the regenerated dry sclerophyll forests of the Victorian box‐ironbark ecosystem. However, many box‐ironbark species possess traits consistent with fire‐cued regeneration. It is unclear the degree to which human disturbance may have altered fire regimes in these forests. The infrequent and low‐intensity fire regime suggested by current fuel dynamics may pose a threat to the persistence of fire‐cued species. Obligate seeders such as those of the Fabaceae and Mimosaceae, common in box‐ironbark understoreys, may be particularly vulnerable if inter‐fire intervals exceed seed longevity. This study used seed burial trials to examine seed dormancy and longevity in five legume species to explore their capacity to regenerate under an infrequent, low‐intensity fire regime. All species displayed dormancy and longevity patterns consistent with other south‐east Australian legumes. Before burial, dormancy levels were high for all species (98–100%). After 3 years, storage under in situ and ex situ conditions, dormancy in Pultenaea prostrata remained at pre‐burial levels with virtually no seed becoming non‐dormant. Over time, some Acacia seed became non‐dormant under both in situ and ex situ storage, with the pattern varying among species. Longevity also varied between species. Variation in the dormancy and longevity patterns observed in these obligate seeder legumes suggests two strategies: (i) releasing a portion of soil‐stored seed from dormancy during the inter‐fire period to permit inter‐fire recruitment; and (ii) retaining most soil‐stored seed as dormant during the inter‐fire interval. Both strategies represent potential weaknesses under a long fire interval regime. The first relies on dormancy release translating to successful recruitment and requires ongoing inter‐fire input into the soil seed bank. The second relies on seed longevity exceeding the inter‐fire interval. Whether either is more suitable to coping with long‐term infrequent fire requires long‐term monitoring.