Fire survival of Cape Proteaceae-influence of fire season and seed predators

Many Cape Proteaceae store seed reserves in closed cones on the plant and rely entirely on these reserves for episodic recruitment after fires. Population size is sensitive to intervals between fires but also to fire season. Populations can be nearly eliminated by successive winter or spring fires. Three hypotheses explaining seasonal variation in recruitment were tested: (a) seeds germinate immediately after fire but seedlings die from summer drought; (b) seeds remain dormant over summer but the longer the delay between seed release after fire and germination 1) the greater the competition between seedlings and resprouts, or 2) the greater the seed losses to predators and/or decay before germination.Drought-avoiding dormancy occurred in 9 of 11 Cape Proteaceae studied, all of which delayed germination to autumn or winter. Seedling emergence and survival was not significantly increased after removal of competitors by methyl bromide poisoning. Seed predation, measured by exclosures, however, significantly reduced seed reserves before germination and also number of seedlings emerging. Post emergence seedling predation was negligible in the burn in contrast to adjacent mature vegetation where seedling predation was very heavy.The role of germination cues and rodent behaviour in controlling population recruitment is discussed and it is concluded that a knowledge of both is essential for predicting vegetation dynamics in this system.Acknowledgements: I am grateful to J. Vlok and R. America for field assistance and J. Breytenbach, F. Kruger, P. Slingsby and J. Yensch for observation and comment. This work was supported by the Directorate of Forestry, South Africa as part of their conservation research programme.     

Comparative seed biology and co-existence of two fynbos shrub species

Abstract. Fire-prone fynbos communities include numerous species which are difficult to distinguish in terms of conventional niche axes since they are morphologically similar and share the same broad life-history traits. These species compete for space after each fire and lottery theory predicts that those with dissimilar per capita regeneration would not co-exist in the long-term. We studied seed production, soil seed bank dynamics, germination and post-fire seedling establishment and mortality of two obligate reseeding, morphologically similar shrubs which co-occur in dune fynbos throughout the southwestern Cape. Passerina paleacea produced 30 x more plump seeds and maintained soil seed banks between 1.5 x (pre-dispersal) and 2.6 x (post-dispersal) times larger than Phylica ericoides. Seeds of both species showed a high degree of dormancy although germination was stimulated by direct fire-related cues for Phylica only. Approximately 4 x more Passerina seedlings than Phylica seedlings were counted at the end of the winter germination period after a mild autumn fire. However, seedling mortality in the first five months of the dry summer was more than twice as high for Passerina than for Phylica. We suggest that the long-term co-existence of these two species does not result from similar post-fire regeneration success but rather from population instabilities arising from differential regeneration in relation to fire regime. The results predict that hot fires followed by dry summers would favour Phylica over Passerina as the former has fire-stimulated germination and greater drought tolerance; cool fires followed by moist summers would favour Passerina.     

Stimulation of seed germination in South African species of Restionaceae by plant-derived smoke

The Restionaceae is one of the three major families defining fynbos, the characteristic vegetation type of the Cape Floristic Region. Periodic fires with a frequency of 5 to 40 years are a natural phenomenon in fynbos vegetation. Fire-stimulated seed germination has been reported for a variety of fynbos species, and species in the Asteraceae, Ericaceae, Proteaceae and Restionaceae have shown a germination response to smoke and/or aqueous smoke extracts. In the present study seed of 32 species was screened to obtain an indication of how important the smoke cue is for germination in the South African Restionaceae. The results of the present study represents the first occasion that comparative germination data for South African species in this family have ever been obtained. Twenty-five of the 32 species tested showed a statistically significant improvement in germination following smoke treatment. Untreated seeds of 18 of the species responding, showed a high degree of dormancy with only 0.1% to 2.0% germination. These results suggest that under natural conditions smoke from fynbos fires may provide an important cue for triggering seed germination in this family. The degree of improvement in germination following smoke treatment ranged from 147% in the case of Restio festuciformis to 25300% in the case of Rhodocoma capensis. It is suggested that the 16 species which showed a 1000% or more increase in germination following smoke treatment form a group in which smoke is likely to be the major cue for germination. In those species in which there is a lesser response, smoke may be one of a number of germination cues which include heat, and possibly alternating high and low incubation temperatures. The four species that did not germinate were all myrmecochorus, nut-fruited species. More information is needed concerning the fire survival strategy of South African Restionaceae species and many more will have to be investigated in the nursery and in the field, before the full pattern of response within the family and its significance can be elucidated.     

Fire season and intensity affect shrub recruitment in temperate sclerophyllous woodlands

The season in which a fire occurs may regulate plant seedling recruitment because of: (1) the interaction of season and intensity of fire and the temperature requirements for seed release, germination and growth; (2) post-fire rainfall and temperature patterns affecting germination; (3) the interaction of post-fire germination conditions and competition from surrounding vegetation; and (4) the interaction of post-fire germination conditions and seed predators and/or seedling herbivores. This study examined the effects of different fire intensities and fire seasons on the emergence and survival of shrubs representing a range of fire response syndromes from a summer rainfall cool climate region. Replicated experimental burns were conducted in two seasons (spring and autumn) in 2 consecutive years and fuel loads were increased to examine the effects of fire intensity (low intensity and moderate intensity). Post-fire watering treatments partitioned the effects of seasonal temperature from soil moisture. Higher intensity fires resulted in enhanced seedling emergence for hard-seeded species but rarely influenced survival. Spring fires enhanced seedling emergence across all functional groups. Reduced autumn recruitment was related to seasonal temperature inhibiting germination rather than a lack of soil moisture or competition. In Mediterranean-type climate regions, seedling emergence has been related to post-fire rainfall and exposure of seeds to seed predators. We think a similar model may operate in temperate summer rainfall regions where cold-induced dormancy over winter exposes seeds to predators for a longer time and subsequently results in recruitment failure. Our results support the theory that the effect of fire season is more predictable where there are strong seasonal patterns in climate. In this study seasonal temperature rather than rainfall appears to be more influential.     

Burning Influences on Wiregrass (Aristida beyrichiana) Restoration Plantings: Natural Seedling Recruitment and Survival

Regeneration and expansion of Aristida beyrichiana and Aristida stricta (wiregrass) populations in remaining fire‐maintained Pinus palustris (longleaf pine) stands of the southeastern United States has become an objective of land managers. Although growing‐season fire is required for successful wiregrass seed production, studies examining naturally occurring wiregrass seedling dynamics are few. This study investigates how seedling survivorship is affected by season of burn, seedling size, time since germination, and proximity to adult plants. Restoration at this research site was begun in 1992 with the planting of containerized longleaf pine and wiregrass seedlings. Study plots were established in November 1997 after a growing‐season prescribed fire (June 1996) that resulted in successful seed production and seedling recruitment. Burn treatment plots included (1) no burn (control), (2) fire in the dormant season of the first year after germination (March 1998), (3) fire in the growing season of the first year after germination (August 1998), and (4) fire in the growing season of the second year after germination (July 1999). Seedling mortality increased with growing season burning and close proximity to planted adults. Natural seedling recruitment continued into the second year after initial seed‐drop in all plots, which verifies that wiregrass seed banking occurs for a minimum of 2 years after seed drop. Where wiregrass management objectives include population expansion, seedling recruits should be allowed 1 to 2 years post‐germination without growing season fire for successful establishment.     

Fire season effects on the recruitment of non‐sprouting serotinous Proteaceae in the eastern (bimodal rainfall) fynbos biome,South Africa

Abstract Research in Mediterranean‐climate shrublands in both South Africa and Australia shows that recruitment of proteoid shrubs (non‐sprouting, serotinous Proteaceae) is best after warm‐season (summer and autumn) fires and worst after cool‐season (winter and spring) ones. This pattern has been attributed to post‐dispersal seed attrition as well as size of pre‐dispersal seed reserves. Here we investigate patterns of post‐fire recruitment for four proteoid species in the eastern part of South Africa's fynbos biome, which has a bimodal (spring and autumn) rainfall regime. Despite the lack of significant differences in recruitment between cool‐ and warm‐season burns, we find some evidence for favourable recruitment periods following fires in spring and autumn, immediately before, and coinciding with, the bimodal rainfall peaks. This suggests that enhanced recruitment is associated with conditions of high soil moisture immediately after the fire, and that rapid germination may minimize post‐dispersal seed attrition. In two of the species, we also find a shift from peak flowering in winter and spring in the Mediterranean‐climate part of the fynbos biome, to summer and autumn flowering in the eastern part. Because these two species are only weakly serotinous, warm‐season flowering would result in maximal seed banks in spring, which could explain the spring recruitment peak, but not the autumn one. We conclude that eastern recruitment patterns differ significantly from those observed in the western and central parts of the biome, and that fire management protocols for the east, which are currently based on data and experience from the winter‐rainfall fynbos biome, need to be adjusted accordingly. Fire managers in the eastern fynbos biome should be less constrained by requirements to burn within a narrow seasonal range, and should therefore be in a better position to apply the required management burns.     

Invasion and persistence of bird-dispersed,subtropical thicket and forest species in fire-prone coastal fynbos

Abstract. Seedling abundance at four microsites (open fynbos, beneath emergent fynbos shrubs, beneath thicket, and beneath forest) was determined at three coastal dune landscapes, located along a gradient of increasing summer rainfall and where fire-dependent fynbos was the predominant vegetation. At all sites thicket seedlings were most common beneath emergent fynbos shrubs and under thicket clumps; seedlings of forest species were most abundant at forest microsites although some individuals were recorded beneath thicket. Very few thicket seedlings were observed in open fynbos. Birds play a keystone role in facilitating establishment of the fleshy fruit-bearing thicket flora. Seedling abundance at microsites of different thicket and forest species was generally unrelated to fruit abundance. Germination success of most species was highest under shaded conditions; soil organic content had no effect on germination. Removal of pulp and birdingestion enhanced the germination, relative to untreated controls, of two out of three species tested. A simple Markov model predicted a gradual increase in cover of the thicket and forest component and a gradual decline in fynbos under a ‘normal’ (20-yr interval) fire regime simulated over 10 cycles. Although inter-fire seedling establishment under emergent fynbos shrubs is important in the initial colonisation of fynbos by obligate resprouting thicket shrubs, these species persist and expand by vegetative recruitment after and between fires, respectively. In the prolonged absence of fire, the endemic-rich and fire-dependent fynbos flora would be replaced by species-poor forest and thicket.     

Effect of fire on hard-coated Cistaceae seed banks and its influence on techniques for quantifying seed banks

The impact of fire on hard-coated Cistaceae (Halimium ocymoides, Cistus ladanifer, and C. salvifolius) soil seed banks in a Mediterranean 'maquis' shrubland, and its effect on seed germinability were studied. The study also contrasts the effectiveness of two widely used techniques for quantifying seed banks, the seedling emergence and the physical separation methods, in relation to fire. The null hypothesis that a massive enhancement of physically-dormant Cistaceae seed germination by fire would make use of the time-consuming physical separation technique unnecessary was tested. Fire reduced Cistaceae seed banks in the 0–2 cm deep soil layer by both seed fire-consumption and lethal temperatures, revealed by the significant decreasing of the seed bank density and by the increase of apparently-intact but soft-unviable seeds, respectively. In contrast, no damage was recorded in the 2–5 cm soil layer. A dramatic seed bank depletion (> 90%) in both soil layers was recorded one year after fire in the burnt area, coinciding with a significant increase of seedling density confined to the first post-fire year. The ecological consequences of this massive post-fire seed bank input are discussed. A germinability test revealed that germination of surviving Cistaceae seeds was significantly enhanced in all cases except for the C. salvifolious seed bank in the deeper soil layer. However, final germination levels (60–75%) did not correspond to the magnitude of seed bank depletion, especially for C. salvifolious, which suggests that other environmental factors not exclusively associated with fire may also be important in softening Cistaceae seeds. Germination enhancement by fire soil-heating was not high enough to reject the physical separation technique, at least in the deeper soil layer. The simultaneous use of both seedling emergence and physical separation is recommended for reliable seed bank estimates when a physically-dormant hard-seeded component can be expected in the soil, as in many Mediterranean ecosystems, regardless of fire occurrence.     

Aspects of the reproductive ecology of four australian Hakea species (Proteaceae) in South Africa

Summary Four shrub species of the Australian Proteaceae (Hakea sericea, H. gibbosa, H. suaveolens and H. salicifolia) were introduced to South African fynbos shrublands between 1840 and 1860. H. sericea is highly invasive, H. gibbosa and H. suaveolens are moderately invasive and H. salicifolia is not invasive. The allocation of reproductive energy, germinability, the ability to survive fires and to germinate in burnt and unburnt areas, and the nutrient content of seeds were assessed for the four species. The information was used to investigate whether the success of H. sericea relative to the other three species could be explained by the superior expression of any trait. The most important trait which separates H. sericea from the other species is its ability to produce a large seed bank in its adopted environment in the absence of seed predators. Seed production in H. sericea shrubs with an above-ground dry mass of 8 kg is four times greater than H. gibbosa and more than 16 times that of H. suaveolens. Although H. salicifolia also produces a large seed bank, its seeds are unable to survive fires due to inadequate insulation by the small follicles. The results are compared to dispersal and seed bank data for indigenous South African Proteaceae, which have low dispersal and suffer high pre-dispersal seed predation. We suggest that potential invasives in the fynbos can be identified as species that have: (i) a potentially high seed production that is limited by specialized predators; (ii) an ability to disperse over long distances; and (iii) are pre-adapted to frequent fires and low soil nutrients. The data also support the current strategy of combatting H. sericea using specialized insect seed predators.     

Rodent consumption and caching behaviour selects for specific seed traits

The seed characteristics selected for by scatter-hoarding rodents can have an impact on seed morphology, seedling establishment and ultimately on plant community structure. Using Leucadendron sessile (Proteaceae), it was recently discovered that rodents are seed dispersers in the fynbos biome of South Africa. However, little is known about the characteristics of rodent-dispersed seeds and the selective influence rodents have on seed morphology in this biome. We investigated the caching behaviour of rodents and asked whether variation in seed traits (size, hull thickness) influenced whether seeds were more likely to be consumed or cached. Rodents tended to disperse and bury, rather than consume, medium sized L. sessile seeds with medium hull thickness. In contrast, small or thin hulled seeds were preferentially eaten in situ and were seldom buried. Large seeds or seeds with thick hulls were often left untouched at depots. Our results suggest that rodents may impose stabilizing selective pressures on seed size and hull thickness, traits that may also have consequences for seedling mortality, dormancy, competitive interactions and the survival of fires.     

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.     

Impact of fire on Picea mariana and Pinus banksiana seedlings in subarctic lichen woodlands

Abstract. The demography of Picea mariana (black spruce) and Pinus banksiana (jack pine) seedlings was monitored through five censuses over 13 months in four different seed bed types after fire of four severity levels in lichen woodland. Most seeds germinated just before early frost in late summer 1990 or immediately after snow thaw in early spring 1991; the germination rate subsequently decreased. For both species, germination rate decreased along a gradient of fire severity. The proportion of Pinus seeds that produced a seedling surviving 13 months after sowing was 4.3 % and 0.4 % respectively in the intact lichen mat and in the mineral soil seed bed type. For Picea these values are 3.2 % and 0.2 % respectively. The low germination rate in a severely burned seed bed type appeared to be associated with the formation of a water-repellent crust at the soil surface following the fire. Seedlings were contagiously distributed and were more frequent in flat and hollow microsites, where there is probably more water available than on bumps or among pebbles. Properties of experimentally burned seed bed types may differ from those under natural fires where regeneration by seed generally occurs following dispersal. However, the high germination rate observed in the intact lichen mat suggests that scattered lichen woodland patches may respond to increased seed input by a higher frequency of seedling establishment.     

Seed depredation negates the benefits of midstory hardwood removal on longleaf pine seedling establishment

Midstory hardwoods are traditionally removed to restore longleaf pine on fire‐excluded savannas. However, recent evidence demonstrating midstory hardwood facilitation on longleaf pine seedling survival has brought this practice into question on xeric sites. Also, midstory hardwoods could facilitate longleaf pine seedling establishment, as hardwood litter may conceal seeds from seed predators or improve micro‐environmental conditions for seedling establishment. However, little is known about these potential mechanisms. In this study, we tracked longleaf pine seed depredation and germination in artificially seeded plots (11 seeds/m²) in a factorial design fully crossing hardwood retention or removal with vertebrate seed predator access or exclusion in the Sandhills Ecoregion of North Carolina, U.S.A. Seed depredation averaged 78% across treatments and was greatest in unexcluded plots. Hardwood retention did not affect seed depredation. Longleaf pine averaged 3.6 germinants/4 m² across treatments, and was six times more abundant where vertebrates had been excluded. Hardwood removal had a strong positive effect on seedling germination, likely due to the removal of litter, but only when vertebrates were excluded. Our results indicated midstory hardwoods are not facilitating longleaf pine seedling establishment. Nevertheless, our results indicated that hardwood removal may not increase longleaf pine seedling establishment, as seed depredation diminished the effectiveness of hardwood removal under mast seed availability. Collectively, these results demonstrate the underlying complexity of the longleaf pine ecosystem, and suggest that planting may need to be part of the restoration strategy on sites where seed depredation limits longleaf pine natural regeneration.     

The costs of leaving home: ants disperse myrmecochorous seeds to low nutrient sites

Summary Leucospermum conocarpodendron (L.) Buek (Proteaceae) seedlings were excavated several months after a fire in Cape fynbos. Seedlings under burnt parental skeletons had short hypocotyls (mean 25 mm) indicating passive dispersal whereas seedlings in the open were more deeply buried (mean 48 mm) by ants. Soil nutrient concentrations at the site of germination were negatively related to depth of burial and distance from parent. Ant dispersal resulted in seedlings emerging in soils with lower nutrient concentrations than passively dispersed seeds. Tissue analysis supported the soil results with lower P content in seedlings from open (ant dispersed) sites. Seedling survival in the first year of establishment was also lower in open sites, but not significantly so. However seedlings were slightly taller in the open. The results of this study, the first on naturally occurring intraspecific variation in myrmecochory, strongly contradict current explanations for the high incidence of myrmecochory in nutrient poor environments.     

Hydration status influences seed fire tolerance in temperate European herbaceous species

Prescribed burning is an important management tool in many parts of the world. While natural fires generally occur during the driest and warmest period of the year, prescribed burning is often timed out‐of‐season, when there is higher soil moisture and lower biomass combustibility. However, fire season may influence seedling recruitment after fire, e.g. through the effect of seed hydration status on fire tolerance. In non‐fire‐prone temperate regions, anthropogenic fire may occur exclusively in periods outside the growing season with higher soil moisture, which may have negative consequences on seedling recruitment. Fire tolerance of moist and dry seeds of 16 temperate European herbaceous species belonging to four families was assessed using heat treatment of 100 °C for 5 min and subsequent germination trials. Moist seeds of Asteraceae, Poaceae and Brassicaceae had a predominantly negative reaction to the heat treatment, while those of Fabaceae tolerated it or germination was even enhanced. The reaction of dry seeds was completely different, with positive responses in three species of the Fabaceae and fire tolerance in species of other families. Our results point out that hydration status may significantly influence the post‐fire germination of seeds. Dry seeds were found to tolerate high heat, while moist seeds were harmed in more than half of the species. This implies that if prescribed burning is applied in temperate grasslands of Europe, it should be timed to dry periods of the dormant season in order to protect seeds from negative effects of fire.     

Wide variation in post-emergence desiccation tolerance of seedlings of fynbos proteoid shrubs

Fynbos Proteaceae that are killed by fire and bear their seeds in serotinous cones (proteoids), are entirely dependent on seedling recruitment for persistence. Hence, the regeneration phase represents a vulnerable stage of the plant life cycle. In laboratory-based experiments we investigated the effect of desiccation on the survival of newly emerged seedlings of 23 proteoid species (Leucadendron and Protea) occurring in a wide variety of fynbos habitats. We tested the hypothesis that species of drier habitats would be more tolerant of desiccation than those from more moist areas. Results showed that with no desiccation treatment, or with desiccation prior to radicle emergence, all species germinated to high levels. However, with desiccation treatments imposed after radicle emergence, there were significant declines in seedling emergence after subsequent re-wetting. Furthermore, other than three species that grow in waterlogged habitats, germination responses could not be reliably modeled as a function of soil moisture variables. An important finding was that the species had highly individualistic responses to desiccation. In conclusion, early seedling emergence represents a species-specific stage that is highly sensitive to a decrease in soil moisture. Since species are killed by fire (non-sprouting), vulnerability to increasing aridity associated with anthropomorphic climate change would increase the odds of local and global extinction.     

Fire increases aboveground biomass, seed production and recruitment success of Molinia caerulea in dry heathland

During the last decades, the perennial tussock grass Molinia caerulea has shown an increased abundance in European heathlands, most likely as a result of increased nitrogen deposition and altered management schemes. Because of its deciduous nature, Molinia produces large amounts of litter each year, which may affect the intensity and frequency of accidental fires in heathlands. These fires may influence plant population dynamics and heathland community organization through their effects on plant vital attributes and competitive interactions. In this study, fire-induced changes in competitive ability and invasiveness of Molinia through changes in biomass production, seed set and seed germination under both natural and laboratory conditions were investigated. We found that fire significantly increased aboveground biomass, seed set and germination of Molinia. Seed set was twice as high in burned compared to unburned heathland. Two years after fire, seedling densities in natural conditions were on average six times higher in burned than in unburned heathland, which resulted in increased abundance of Molinia after burning. The seed germination experiment indicated that seeds harvested from plants in burned heathland showed higher germination rates than those from unburned heathland. Hence, our results clearly demonstrate increased invasive spread of Molinia after large and intense fires. Active management guidelines are required to prevent further encroachment of Molinia and to lower the probability of large fires altering the heathland community in the future.     

Fire increases aboveground biomass,seed production and recruitment success of Molinia caerulea in dry heathland

During the last decades, the perennial tussock grass Molinia caerulea has shown an increased abundance in European heathlands, most likely as a result of increased nitrogen deposition and altered management schemes. Because of its deciduous nature, Molinia produces large amounts of litter each year, which may affect the intensity and frequency of accidental fires in heathlands. These fires may influence plant population dynamics and heathland community organization through their effects on plant vital attributes and competitive interactions. In this study, fire-induced changes in competitive ability and invasiveness of Molinia through changes in biomass production, seed set and seed germination under both natural and laboratory conditions were investigated. We found that fire significantly increased aboveground biomass, seed set and germination of Molinia. Seed set was twice as high in burned compared to unburned heathland. Two years after fire, seedling densities in natural conditions were on average six times higher in burned than in unburned heathland, which resulted in increased abundance of Molinia after burning. The seed germination experiment indicated that seeds harvested from plants in burned heathland showed higher germination rates than those from unburned heathland. Hence, our results clearly demonstrate increased invasive spread of Molinia after large and intense fires. Active management guidelines are required to prevent further encroachment of Molinia and to lower the probability of large fires altering the heathland community in the future.     

Seed predation limits post-fire recruitment in the waratah (<Emphasis Type="Italic">Telopea speciosissima</Emphasis>)

Seed predation may reduce recruitment in populations that are limited by the availability of seeds rather than microsites. Fires increase the availability of both seeds and microsites, but in plants that lack a soil- or canopy-stored seed bank, post-fire recruitment is often delayed compared to the majority of species. Pyrogenic flowering species, such as Telopea speciosissima, release their non-dormant seeds more than 1 year after fire, by which time seed predation and the availability of microsites may differ from that experienced by plants recruiting soon after fire. I assessed the role of post-dispersal seed predation in limiting seedling establishment after fire in T. speciosissima, in southeastern Australia. Using a seed-planting experiment, I manipulated vertebrate access to seeds and the combined cover of litter and vegetation within experimental microsites in the 2 years of natural seed fall after a fire. Losses to vertebrate and invertebrate seed predators were rapid and substantial, with 50% of seeds consumed after 2 months in exposed locations and after 5 months when vertebrates were excluded. After 7 months, only 6% of seeds or seedlings survived, even where vertebrates were excluded. Removing litter and vegetation increased the likelihood of seed predation by vertebrates, but had little influence on losses due to invertebrates. Microsites with high-density vegetation and litter cover were more likely to have seed survival or germination than microsites with low-density cover. Recruitment in pyrogenic flowering species may depend upon the release of seeds into locations where dense cover may allow them to escape from vertebrate predators. Even here, conditions suitable for germination must occur soon after seed release for seeds to escape from invertebrate predators. Seed production will also affect recruitment after any one fire, while the ability of some juvenile and most adult plants to resprout after fire buffers populations against rapid declines when there is little successful recruitment.     

Patch to landscape patterns in post fire recruitment of a serotinous conifer

Obligate seeding species are highly specialized to fire disturbance and many conifers such as cypress, which are adapted to high intensity stand-replacing fires, have canopy seed banks stored in serotinous cones. Resilience of these trees to fire disturbance is a function of disturbance frequency and one focus of this study was to determine the effect of patch age on postfire recruitment. A second focus was to determine the extent to which fire induced a landscape level change in the location of the forest boundary. Prior to a fire in 1994, a large Cupressus sargentii forest was a mosaic landscape of different aged patches of nearly pure cypress bordered by chaparral. Patches less than 60 years of age were relatively dense with roughly one tree every 1–2 m² but older patches had thinned to one tree every 3–15 m². Older trees had substantially greater canopy cone crops but the stand level seed bank size was not significantly correlated with stand age. Fire-dependent obligate seeding species are sensitive to fire return interval because of potential changes in the size of seed banks – facing both a potential `immaturity risk' and a `senescence risk'. At our site, C. sargentii regeneration was substantial in stands as young as 20 years, suggesting that fire return interval would need to be shorter than this to pose any significant risk. Reduced seedling recruitment in stands nearly 100 years of age may indicate risk from senescence is greater, however, even the lowest density seedling recruitment was many times greater than the density of mature forests – thus this cypress would appear to be resilient to a wide range of fire return intervals. Changes in landscape patterning of forest and chaparral are unlikely except after fire. Factors that inhibit tree establishment within the shrubland, as well as factors that affect shrub establishment within the forest border likely affect the `permeability' of this ecotone. After the 1994 fire this boundary appeared to be stable in that cypress recruited best within the shadow of burned canopies and cypress were weak invaders of adjacent shrublands.