Interactive effects of heat shock and smoke on germination of nine species forming soil seed banks within the Sydney region

Abstract The germination response of seeds from fire‐prone vegetation to fire‐related cues such as heat shock and smoke has usually been studied by applying the cues singly. The few studies that have applied the cues in combination have shown that interactions between the cues are possible. Here, the response of seeds from a number of species to combined heat shock and smoke is reported. Heat shock (25, 50, 75 and 100°C) and aerosol smoke (0, 5, 10 and 20 min) were applied factorially to nine species that form soil seed banks in the Sydney region of south‐eastern Australia. These species were from Epacridaceae (four species), Myrtaceae (four species) and Cyperaceae (one species) and ranged from fire‐sensitive obligate seeders to fire‐tolerant facultative resprouters. Germination of Dracophyllum secundum R. Br and Sprengelia monticola (A. Cunn. ex DC.) Druce was low and did not respond to the germination cues. The positive response of Gahnia sieberiana Kunth and Kunzea ambigua (Sm.) Druce to heat shock and smoke was independent and additive. The positive response of Kunzea capitata Rchb. to the interaction between heat shock and smoke was synergistic, and the response of Baeckea diosmifolia Rudge and Baeckea imbricata (Gaertn.) Druce was unitive, with germination increase only occurring following combined heat and smoke application. Epacris coriacea A. Cunn. ex DC. and Epacris obtusifolia Sm. had low levels of dormancy and hence it was not possible to find a fire response. Gahnia sieberiana and K. capitata responded differently to the combination of heat shock and smoke than has previously been reported. Germination of species from habitats that are infrequently burnt was not affected by heat shock or smoke. Low‐intensity fire or patches within fire may be important for seedling recruitment as the 50°C heat shock stimulated germination in four of the five species that responded to the heat cue, and germination of Baeckea imbricata declined within the 100°C heat shock treatment. Germination of one species, Baeckea imbricata, was only stimulated by a specific combination of cues, indicating that regeneration niches may be narrow for some species and that the application of a range of heat and smoke doses is required to find such responses. Of the species positively responding to heat shock and smoke, a requirement for both cues was prevalent, therefore the response to these cues in isolation cannot be relied upon to give a true indication of the fire response of a species.     

Fire‐related cues and germination from the soil seed bank of senescent remnants of mallee vegetation on Eastern Kangaroo Island

Plant communities dominated by narrow‐leaved mallee (Eucalyptus cneorifolia) are almost entirely confined to north‐eastern Kangaroo Island, South Australia, an area which has been extensively cleared for agriculture. Consequently, surviving examples consist mostly of small remnants which are thought to be senescent due to the exclusion of fire. This senescence is associated with the loss of many native understory species. Prescribed burns have been suggested as a management tool to stimulate the restoration of native plants from the soil seed bank; however, no seed bank studies have previously been conducted on Kangaroo Island and the seed bank literature usually focuses on particular species rather than on plant communities. We conducted an experiment to investigate the effects of the fire‐related cues heat and smoke on the germination of plants from the seed bank in soil sampled from 10 long‐ungrazed narrow‐leaved mallee sites on Kangaroo Island. Eighty trays of soil were monitored in a controlled glasshouse for five months after being subjected to heat and/or smoke treatments. The overall number of native, but not exotic, plant species germinating from the soil seed bank was significantly increased by all three fire‐related treatments (heat, smoke and heat plus smoke) compared with the control (no fire‐related treatment). Different plant life forms exhibited varying responses to heat and smoke treatments. The results of this study illustrate that the application of fire‐related treatments to soil seed banks in controlled glasshouse conditions can stimulate the recruitment of native species, including several species of conservation concern. These findings also indicate the potential of using these treatments for the ex situ germination of fire dependent species for revegetation purposes and indicate aspects of prescribed burns that may be important for restoring different components of native vegetation.     

Germination response of common annual and perennial forbs to heat shock and smoke treatments in the Chaco Serrano,central Argentina

Fire is a key ecological factor affecting plant dynamics. In the last few decades, fire occurrence in the Chaco region has increased noticeably, challenging the adaptive capacity of plants to regenerate after a fire. Broad‐leaved forb species have been much less studied than woody and graminoids, although they are an important component of fire dynamics. Here we analysed the germination response to heat shock of 70 and 110°C, smoke and their combination in 10 broad‐leaved herbaceous species frequently occurring in the Chaco Serrano of Córdoba province, central Argentina, including five annual (Bidens subalternans, Conyza bonariensis, Schkuhria pinnata, Tagetes minuta and Zinnia peruviana) and five perennial species (Borreria eryngioides, Sida rhombifolia, Solidago chilensis, Taraxacum officinale and Verbena litoralis). We also compared the response of annual versus perennial species. Six species had highest germination when treated with heat and smoke combined, whereas two had lowest germination under this treatment, indicating synergistic and antagonistic interaction of these factors respectively. Most of the species tolerated heat shock (i.e. germination was similar to that in control treatment), whereas others had higher germination in response to heat shock, especially under the moderate 70°C treatment. Germination was higher than control (i.e. no heat and no smoke) after smoke treatment in four species. Perennial species showed higher average germination than annuals in both heat treatments and in the control. Annual species had higher average germination for all treatments involving smoke. The high variability observed at the species level, and the limited number of species studied calls for precaution in interpreting and extrapolating results. Nevertheless, our study shows a general positive response of both perennial and annual species to fire cues, suggesting an advantage of these species for colonizing post‐fire environments, and being favoured under scenarios of increasingly frequent low‐to‐medium intensity fires.     

Are germination cues for soil‐stored seed banks different in structurally different fire‐prone communities?

Many plant species are dependent on soil‐stored seeds for their persistence in fire‐prone systems. Seed germination is often stimulated by fire‐related cues including heat and smoke, but the way these cues promote germination may differ between structurally distinct plant communities with historically different fire regimes. In this study, we examined the effects of heat, smoke and their interactions on the germination of soil‐stored seeds from shrubby woodlands and herbaceous forests in south‐east Australia. The effect of these treatments on species richness, diversity and composition, and species richness and density of germinants within life‐forms (grass, forb and shrub) were assessed. Soils from each community were subjected to low heat (40°C), low heat with smoke, smoke, high heat (80°C), high heat with smoke and untreated (control) before being placed in a glasshouse, where the germinants were identified and counted. Greater species richness was stimulated by high heat treatments and smoke in both communities, a trend driven by shrubs and forbs, rather than grasses. Greater species diversity was stimulated by high heat with smoke in both communities. Greater densities of grass germinants were stimulated by all treatments, except low heat, in both communities. For forbs and shrubs, the effect of treatment depended on community. Compared to the control, low heat with smoke (forbs) and both low heat and low heat with smoke (shrubs) increased densities in the woodland but not in the forest. There were unique species compositions, different from the control, in all treatments in the forest but not in the woodland, where composition in low heat was not different from the control. These results indicate the importance of high soil temperatures and smoke in both communities. In the absence of wildfires, recurring prescribed burns that heat the soil to low temperatures are likely to reduce plant richness, diversity, and density resulting in a change in understorey species composition and structure.     

Seed Dormancy and Germination Responses of Nine Australian Fire Ephemerals

Fire ephemerals are short-lived plants with seeds that persist in the soil and germinate after a fire or physical soil disturbance. Ex situ germination of many Australian fire ephemerals has previously been difficult. Dormancy was present in most of the nine fire ephemerals examined. Alyogyne hakeifolia (Giord.) Alef. and Alyogyne huegelii (Endl.) Fryxell (Malvaceae) seeds had physical and possibly also physiological dormancy, Actinotus leucocephalus Benth. (Apiaceae) seeds had morphophysiological dormancy, Austrostipa compressa (R.Br.) S.W.L. Jacobs & J. Everett and Austrostipa macalpinei (Reader) S.W.L. Jacobs & J. Everett (Poaceae) seeds were either non-dormant or possessed physiological dormancy, and seeds of all remaining species possessed physiological dormancy. A proportion of the Alyogyne hakeifolia, Alyogyne huegelii, Austrostipa compressa and Austrostipa macalpinei seed populations were non-dormant because some seeds could germinate at the various incubation temperatures without further treatment. At 20 °C, artificial methods of inducing germination such as manual or acid scarification were among the optimal treatments for Austrostipa compressa, Austrostipa macalpinei, Alyogyne huegelii, Actinotus leucocephalus and Grevillea scapigera A.S. George (Proteaceae), and gibberellic acid induced maximum germination of Tersonia cyathiflora (Fenzl) J.W. Green (Gyrostemonaceae) seeds. Heat (70 °C for 1 h) and smoke water was one of the most effective treatments for germinating Actinotus leucocephalus and Codonocarpus cotinifolius (Desf.) F. Muell. (Gyrostemonaceae) seeds. Germination of Grevillea scapigera, Codonocarpus cotinifolius, Gyrostemon racemiger H. Walter (Gyrostemonaceae) and Tersonia cyathiflora did not exceed 40% and may require other treatments to overcome dormancy. Although the nine fire ephemerals examined require fire to germinate under natural conditions, a range of germination responses and dormancy types was observed.     

Evidence for direct effects of fire‐cues on germination of some perennial forbs common in grassy ecosystems

In grassy ecosystems of south‐eastern Australia, fire maintains richness of native forbs. It is commonly thought that fire promotes regeneration indirectly by reducing competition for light and providing gaps for recruitment, rather than directly stimulating germination. However, physiological dormancy and morphophysiological dormancy are common, and few studies have explored responses to fire‐cues among dormant or hard‐to‐germinate forbs. Recent studies from other fire‐prone ecosystems suggest that in some cases, fire‐cues may not alleviate physiological or morphophysiological dormancy, but instead promote germination in combination with treatments which alleviate dormancy. We experimentally tested the prevailing hypothesis that perennial forbs common in south‐eastern Australian grassy ecosystems do not germinate in direct response to fire. Responses to fire‐cues both inherently and in combination with treatments which alleviate dormancy were investigated for seven species. Two fire‐cues (smoke and heat) plus a treatment of both heat + smoke were applied to fresh seed at three temperatures (35/25°C, 30/20°C and 25/15°C). Following this, the effect of fire‐cues on seed that had undergone warm stratification, cold stratification and dry‐after‐ripening was investigated. Three species — Arthropodium strictum, Cheiranthera cyanea and Dianella revoluta — responded to fire‐cues inherently, although germination in C. cyanea was low. High germination of D. revoluta was found when fire‐cues were combined with warm stratification. Fire‐cues had no effect on germination of Brunonia australis, Burchardia umbellata and Eryngium ovinum. Germination of Stypandra glauca was zero following all treatment combinations. Our finding that fire‐cues promote germination of three of the seven study species did not provide sufficient evidence to reject the current hypothesis that germination of perennial forbs is not typically promoted by fire‐cues. However, this study highlights the important direct role fire‐cues can play in promoting germination of some perennial forbs both inherently and in combination with treatments used to alleviate physiological dormancy.     

Comparative seed ecology of the endangered shrub, Pimelea spicata and a threatening weed, Bridal Creeper: Smoke, heat and other fire-related germination cues

Summary Pimelea spicata R. Br. is a nationally listed endangered Australian shrub threatened with extinction by habitat fragmentation and environmental weed invasion. Bridal Creeper (Asparagus asparagoides L. W. Wight) is the primary weed threat to the largest remaining populations of P. spicata in the Cumberland Plain. Fire, as part of an integrated pest management program, offers the potential to stimulate P. spicata populations while controlling Bridal Creeper. It is important, therefore, to understand how the components of fire affect the germination and growth of both species. Using laboratory experiments we investigated the effects of smoke, heat, ash and/or light on the germination of P. spicata and Bridal Creeper. We found a significant promotive effect of smoke and indication of an inhibitory heat shock (90°C for 10 min) effect on the germination of P. spicata seeds. The response of Bridal Creeper seeds to the same factors was complex; while the results of one experiment suggested an inhibitory effect of smoke and a promotive effect of heat, subsequent trials were contradictory, implying that Bridal Creeper, like many weeds, is able to germinate under a wide range of environmental conditions. Other experiments investigated the optimal germination temperature and innate dormancy of P. spicata in the absence of fire‐related germination cues. Of the incubation temperatures investigated, the optimal diurnally fluctuating regime for P. spicata germinations was 10°C and 20°C in the night and day, respectively. The innate dormancy of freshly produced seeds disappeared after 3 months. In contrast to Bridal Creeper, we found a persistent germinable seed bank of about 97 P. spicata seeds/m² located in the top 5 cm of the soil profile. While fire alone is unlikely to kill Bridal Creeper plants, fire may help to manage local infestations of the weed by limiting germination and providing opportunity for herbicide treatment of regrowth.     

Effects of smoke,heat and charred wood on the germination of dormant soil‐stored seeds from a Eucalyptus baxteri heathy‐woodland in Victoria,SE Australia

The effects of dry heat, wet heat, charred wood and smoke on the germination of dormant soil‐stored seeds from a Eucalyptus woodland in western Victoria were tested by using a glasshouse seed‐bank germination experiment. Seedling density, species richness and species composition were compared between replicated treated and control samples. A total of 5922 seedlings, comprising 59 plant species, was recorded from the soil samples over a period of 150 days. While a few species dominated (including Centrolepis strigosa, Wahlenbergia gracilenta and Ixodia achillaeoides), 26 species were represented by fewer than five seedlings and 18 species were restricted to single treatment types. With the exception of charred wood, all treatments led to a significant increase in seed germination relative to the control. The highest number of germinants was obtained for the smoke treatment, with a mean (± SE) of 12 547 ± 449 seedlings m^–2. Heat treatments yielded intermediate densities, with means (± SE) varying between 7445 ± 234 and 9133 ± 445 seedlings m^–2. In comparison with the estimates of seed‐bank sizes from other fire‐prone ecosystems, these densities are high. Species richness differed significantly among treatments. Highest mean richness was recorded in the smoke treatment and lowest for the control and charred wood treatments. There were significant differences in seed‐bank species composition between treatment types based on analysis of similarity (Anosim) using Bray–Curtis similarity. While heat was a specific requirement for triggering germination in hard‐seeded species (e.g. Fabaceae), smoke was the most effective trigger for species from a broad range of other families. The potentially confounding effect of physical and chemical mechanisms of germination stimulation in heated bulk soil samples is raised as an issue requiring further investigation in relation to the role of smoke as a germination trigger.     

Fire‐related cues break seed dormancy of six legumes of tropical eucalypt savannas in north‐eastern Australia

Abstract This paper describes an assessment of the effect of exposure to fire‐related cues (heat shock, smoke and nitrate) and the interactions between the cues on seed dormancy release of tropical savanna legumes in north‐eastern Australia. Ten legume species were tested, comprising both native and exotic species. The ten species responded variously to the treatments. Brief exposure to temperatures between 80 and 100°C was found to break the seed dormancy of the native ephemeral herbs Chamaecrista mimosoides, Crotalaria calycina, Crotalaria montana, Indigofera hirsuta and Tephrosia juncea, as well as the exotic ephemeral herb Crotalaria lanceolata. Exposure to 80°C combined with treatment with a nitrate solution produced an additive effect on the germination of Chamaecrista mimosoides and Crotalaria lanceolata. However, the four species with the heaviest seeds, two exotic ephemeral herbs (Chamaecrista absus and Crotalaria pallida) and two native perennials (Galactia tenuiflora and Glycine tomentella) displayed no significant increase in germination with exposure to fire‐related cues. Exposure to 120°C for 5 min produced seed mortality in all species tested. Two of the largest seeded species, Crotalaria pallida and Galactia tenuiflora, displayed the lowest tolerance to heat shock, with seed mortality after exposure to 100°C for 5 min. These data indicate that fire can promote the germination of some tropical savanna legumes. As a proportion of seeds of each species displayed no innate dormancy, some germination may occur in the absence of fire, especially of exotic species.     

Fire and flood: Soil‐stored seed bank and germination ecology in the endangered Carrington Falls Grevillea (Grevillea rivularis,Proteaceae)

Abstract Seed germination is dependent on the interaction between the dormancy state of a seed and the presence of favourable environmental conditions. Thus, the spectacular pulse of seedling recruitment in many Australian vegetation communities following disturbances such as fire can be attributed to changes in microsite conditions and/or the dormancy‐breaking effect of the disturbance on accumulated seed banks. Grevillea rivularis is a threatened species endemic to the area immediately above Carrington Falls in the NSW Southern Highlands. Most of the population is confined to the riparian vegetation zone in woodland and heath, and is therefore subject to periodic disturbance from fire and flood. For this species, a pulse of seedling recruitment has been recorded after fire, flood and mechanical soil disturbance. The aims of this study were to examine the density and vertical distribution of the soil‐stored seed bank and to investigate the role of heat and scarification as cues for germination of fresh and soil‐stored seed. There was a large seed bank under the canopies of established individuals (194 ± 73 seeds m^?2) and most seeds were found in the 0–2 cm and leaf‐litter layers of the soil profile. The germination response of soil‐stored and fresh seed was examined using a hierarchical series of laboratory experiments. Seeds of G. rivularis showed marked dormancy polymorphism. Thirty‐six percent of soil‐stored seed germinated without treatment, whereas no untreated fresh seeds germinated. Scarification or heating caused significant germination of dormant soil‐stored seed, but only scarification resulted in germination of dormant fresh seeds. These results highlight important differences in the dormancy state of soil‐stored and fresh seed. Thus, being a riparian species in a fire‐prone environment, the dormancy mechanisms in seeds of G. rivularis suit this species to disturbance by both fire and flood.     

Germinable soil seed banks in a tropical savanna: seasonal dynamics and effects of fire.

Abstract The germinable soil seed bank of a tropical eucalypt savanna of north‐eastern Australia was found to be dominated by grasses and forbs, with seed bank density ranging from 58 to 792 seeds per square metre, from a total of 53 species. Late dry season fires and the fire‐related cues, heat shock and smoke, broke the seed dormancy of a range of tropical savanna species. Heat shock promoted the germination of the species groups natives, exotics, subshrubs, ephemeral and twining perennial forbs, and the common species Indigofera hirsuta, Pycnospora lutescens and Triumfetta rhomboidea. Exposure to smoke at ambient temperature promoted germination from the soil seed bank of the species groups combined natives, upright perennial forbs and grasses, as well as the common grasses Digitaria breviglumis and Heteropogon triticeus. The germinable soil seed bank varied seasonally, increasing from the mid wet season (February) and early dry season (May) to a maximum in the late dry season (October). The effect of recent fire history on soil seed bank dynamics was limited to the immediate release of some seed from dormancy; a reduction in seed densities of subshrubs and monocots, other than grasses, in recently burnt savanna; and enhanced seed density of the ephemeral I. hirsuta in the year following fire. The seed banks of most savanna species were replenished in the year following burning.     

Germination behaviour of 14 Mediterranean species in relation to fire factors: smoke and heat

Fire is an ecological factor that has been present in the ecosystems of the Mediterranean region for thousands of years. Our study was undertaken to acquire knowledge of the effect of fire on the germination of Mediterranean species. We used high temperatures (up to 60°C) and smoke to determine the effect of these factors on the germination of species from the Mediterranean region. The species selected are characteristic of the central Mediterranean basin and are representative of both woody and herbaceous species: Rhamnus alaternus L., Cistus albidus L., Cistus monspeliensis L., Fumana ericoides (Cav.) Gand., Rosmarinus officinalis L., Melica ciliata L., Avena sterilis L., Bituminaria bituminosa (L.) C.H. Stirt., Anthyllis vulneraria L., Coronilla glauca L., Argyrolobium zanonii (Turra) P.W. Balland, Emerus major Mill., Genista scorpius (L.) D.C. and Spartium junceum L. The seeds were collected in Mediterranean shrubland (8) and woodland (6) ecosystems, around Montpellier, France (24°45′N and 3°50′E). Ten treatments were tested: a control, three smoke treatments and six heat treatments. The average germination level (germination percentage) and the average T₅₀ rates (time taken to reach 50% of germination) were calculated. The smoke and heat act in a different way on each of the species. The smoke enhanced the germination of two species, whilst moderate heat increased germination in all of the species excepting R. officinalis, F. ericoides, A. sterilis, A. vulneraria, and G. scorpius. Germination was fastest in M. ciliata and S. junceum and slowest in A. sterilis, E. major and C. albidus. The cues did not significantly affect the rate of germination. Fire modified the germination response of 12 of the 14 species studied.     

Does Fire Trigger Seed Germination in the Neotropical Savannas? Experimental Tests with Six Cerrado Species

The Cerrado (Brazilian savanna) is a biodiversity hotspot with a history of fire that goes back as far as 10 million years. Fire has influenced the evolution of several aspects of the vegetation, including reproduction and life cycles. This study tested how fire by‐products such as heat and smoke affect the germination of six species common to two Cerrado open physiognomies: wet grasslands and the campo sujo (grassland with scattered shrubs and dwarf trees). We subjected seeds collected in northern Brazil to heat shock and smoke treatments, both separately and combined, using different temperatures, exposure times, and smoke concentrations in aqueous solutions. High temperatures and smoke did not break seed dormancy nor stimulate germination of the Cerrado study species. However, seeds were not killed by high temperatures, indicating that they are fire‐tolerant. Our findings differed from those of other fire‐prone ecosystems (mostly of Mediterranean vegetation), where fire stimulates germination. Moreover, we provide important information regarding germination strategies of non‐woody Cerrado plants, showing the importance of considering the tolerance of seeds to high temperatures when evaluating fire‐related traits in fire‐prone ecosystems.     

Response surfaces for the combined effects of heat shock and smoke on germination of 16 species forming soil seed banks in south‐east Australia

Abstract There is limited understanding of how fire‐related cues such as heat shock and smoke can combine to affect the germination response of seeds from fire‐prone vegetation because combinations of multiple levels of both cues have rarely been investigated. Germination response surfaces were determined for the combination of heat shock and smoke by applying factorial combinations of temperature (up to 100°C) and aerosol smoke (0–20 min) to 16 species that form soil seed banks in the Sydney region of south‐eastern Australia. Duplicate populations of three species were also examined to assess the constancy of a species response surface. Of the 19 populations examined, 16 showed a germination response to both the fire cues, which combined interactively in 14 populations, and independently in two. No population responded only to a single cue; however, seeds of 11 populations responded to heat in the absence of smoke, and nine responded to smoke in the absence of heat. Heat applied in the absence of smoke negatively affected germination in seven populations, either progressively as temperature increased, or above a set temperature. Negative germination responses over part of the temperature range were fully reversed at higher temperatures for unsmoked seeds of four populations (curvilinear heat response). Smoke effects were most frequently positive over all or part of the range of durations used, and when combined with heat frequently fully or partially reversed negative heat effects. Three populations required the obligatory combination of smoke and heat. A novel response to the cues was observed for three species, with smoke reversing negative heat effects at 75°C, being supplanted by a positive heat response of unsmoked seed at 100°C. The response surface for duplicate populations of two of the three species examined was variable. Heat shock and smoke frequently combined to affect germination, in both positive and negative ways. Consequently, to gain an accurate assessment of the response of seeds to fires, an experimental design that samples within the potential response zones of germination cues is essential.     

Seed size and the regeneration niches of one rare (Melaleuca deanei) and three common (Melaleuca styphelioides,Melaleuca thymifolia and Melaleuca nodosa) Melaleuca (Myrtaceae) species of the Sydney region

Melaleuca species occupy varied habitats across Australia with 16 members of the genus occurring in the Sydney district including the rare species Melaleuca deanei F. Muell. Little is known of their germination and recruitment requirements. This paper reports on experiments assessing the effects of temperature, water potential, fire cues, light and shade levels on germination in four species of Melaleuca native to the Sydney region. For the shade experiment, seedling survival at 12 months is also reported. The experiments tested the hypothesis that M. deanei, which exhibits few seedlings in the field, has limited seedling recruitment because of its particular requirements for germination and establishment. Further, that it differs in these requirements from three common congenerics: M. nodosa (Sol ex Gaertn.) Sm., M. thymifolia Sm. and M. styphelioides Sm. Results indicate that M. deanei has a substantially similar temperature and water potential range for germination to the common congeners from 15 to 35°C, and 0 to ?0.65 ψ. Melaleuca nodosa displayed the broadest regeneration niche on all factors assessed. Germination of M. styphelioides was significantly reduced in the dark and M. styphelioides and M. deanei were most sensitive to shade in the seedling establishment/post‐germination phase. Melaleuca deanei had significantly larger seed that was slower to germinate in all experiments. Germination of the four species was unaffected by the application of heat and smoke. The substantially similar germination parameters exhibited by the four species, despite differences in habitat, may reflect their close phylogenetic affinities. The effects of fire on stimulating seed release from the canopy, the high‐light environment post‐fire together with adequate follow‐up rainfall may all be critical in seedling establishment for M. deanei.     

Seed moisture content affects afterripening and smoke responsiveness in three sympatric Australian native species from fire‐prone environments

Germination of freshly collected seeds of three sympatric herbaceous species native to fire‐prone environments in south‐western Australia was significantly improved through the application of novel combinations of dry heat, gibberellic acid, smoke water and dry afterripening. For fresh seeds, combinations of dry heat, gibberellic acid and/or smoke water resulted in >80% germination in Austrostipa elegantissima (Poaceae) and Stylidium affine (Stylidaceae) seeds and >60% germination in Conostylis candicans (Haemodoraceae) seeds, compared with <10% germination of control seeds. For fresh seeds, two broad germination patterns were observed in response to smoke water: nil – low germination for both control and smoke water‐treated seeds (A. elegantissima and S. affine); and a significant smoke response (35%) compared with control seeds (1%) (C. candicans). During afterripening, high germination for A. elegantissima seeds was achieved following 3 months storage of seeds at equilibrium relative humidities of 23–75%, but seeds stored at 5–13% equilibrium relative humidities took 6–36 months to achieve similar levels of germination. Germination of C. candicans seeds also increased after 3 months storage, to >60% at each equilibrium relative humidity and further increases over time were slight. For S. affine seeds >60% germination was achieved only after 36 months storage at 50% equilibrium relative humidity. Seeds from all three species were smoke‐responsive at some point, but the interaction/effects of afterripening on the smoke response varied significantly between species. This study highlights an apparent effect of seed dormancy status on response to smoke and a surprisingly high level of ecological variation in pre‐germination requirements (cues) for these co‐occurring species that may relate to variation(s) in microsite selection forces operating on the soil seed bank of the different species.     

Impacts of fire cues on germination of Brassica napus L. seeds with high and low secondary dormancy

• Agricultural burning is used in farm management operations; however, information about the impact of fire cues on the release and/or induction of secondary dormancy in crop seeds is scarce.
• Seeds from two oilseed rape cultivars were induced for high (HD) or low (LD) secondary dormancy using polyethyleneglycol (PEG) pre‐treatment, and their germination after exposure to various fire cues was compared to control PEG pre‐treated and non‐dormant seeds.
• Non‐dormant seed germination was unaffected by various fire cues. Low doses of aerosol smoke released secondary dormancy in HD seeds, while higher doses increased dormancy of LD seeds. Dilute smoke water also released HD seed secondary dormancy, but concentrated smke water enhanced dormancy in both LD and HD seeds. The concentrated aqueous extracts from charred oilseed rape straw only promoted germination of HD seeds, while dilution inhibited LD seed germination. Heat shock (80 °C, 5 min) released secondary dormancy in HD seeds; however, higher temperatures and/or increased exposure time was associated with seed death. GC‐MS analyses of smoke water revealed two butenolides and an array of monoaromatic hydroxybenzene compounds with potential germination inhibitor or promoter activity.
• The extent of secondary dormancy induction in seeds affects their subsequent responses to fire cues. Both aerosol smoke and smoke water have both germination promoter and inhibitor activity. Lacking any butenolides, aqueous extracts of charred straw contain a potential germination stimulating steroid, i.e. ergosterol. The significance of fire‐derived cues on behaviour of oilseed rape seeds in the soil seed bank is discussed.

     

Disentangling the role of heat and smoke as germination cues in Mediterranean Basin flora

Background and Aims

The role of fire as a germination cue for Mediterranean Basin (MB) plants is still unclear. The current idea is that heat stimulates germination mainly in Cistaceae and Fabaceae and that smoke has a limited role as a post-fire germination cue, in comparison with other Mediterranean-type ecosystems (MTEs), suggesting that fire-stimulated germination is less relevant in the MB than in other MTEs. However, recent studies showed that the assembly of Mediterranean plant communities is strongly driven by post-fire germination, suggesting an important role for fire as a germination cue. We hypothesize that both heat and smoke have important effects on the different post-fire recruitment processes of MB species (e.g. level and rate of germination and initial seedling growth).

Methods

To ascertain the role of heat and smoke in the post-fire germination response of MB woody plants, a germination experiment was performed with seven heat and two smoke treatments on 30 MB woody species from seven different families, including species with water-permeable seeds and species with water-impermeable seeds.

Key Results

Heat stimulated the germination (probability and rate) of 21 species and smoke in eight species, out of the 30 species studied. In addition, six species showed enhanced initial seedling growth after the smoke treatments.

Conclusions

The results suggest that both heat and smoke are important germination cues in a wide range of MB woody species and that fire-cued germination in woody plants of the MB may be as important as in other MTEs.     

How much seed remains in the soil after a fire?

Soil seed banks that persist after a fire are important in fire-prone habitats as they minimise the risk of decline or local extinction in plants, should the fire-free interval be less than the primary juvenile periods of the species. In two common woody plant genera (Acacia and Grevillea) in southeastern Australia, we examined the size and location of the residual seed bank after fire across areas of varying seedling densities at three locations in comparison to the distribution of seeds in the soil at an unburnt site. We found viable dormant seeds remaining in the soil after fire (evidence of residual soil seed bank). A significantly lower proportion of seeds remained in the top 5 cm of soil than at 5–10 cm or 10–15 cm soil depths, independent of seedling density or plant genus. This was due to greater germination, and possibly some seed mortality, near the soil surface. Reduced germination below 5 cm was probably due to the reduced efficacy of the fire cues that break seed dormancy, a declining ability of seeds to emerge successfully from such depths, and the lower abundance of seeds in the soil at such depths. The magnitude of the residual seed bank was similar across 0–5, 5–10 and 10–15 cm soil depths in Acacia suaveolens. For two Grevillea species, most residual seeds were at 0–5 and 5–10 cm. The residual soil seed bank in the top 10 cm of soil after fire varied across sites with estimates of 0, 19 and 27% in G. speciosa and 23, 35, and 55% in A. suaveolens. At two sites, both species had similar residual seed bank sizes, while at a third, there were large differences between the species (0–55%). The observed patterns imply that the fire-related cues that break seed dormancy generally declined with soil depth. For Acacia, seed dormancy is broken by heat shock, a fire-cue that declines with soil depth. Some 250 species (approx 15% of the fire-prone flora) in the region are thought to have dormancy broken by heat shock. For Grevillea, where seed dormancy is broken by the interaction of smoke and heat shock, at two sites, we suggest three possibilities: (i) the smoke cue declined with soil depth; (ii) both heat and smoke are obligatory for breaking seed dormancy; or (iii) the cues may be independent and additive and below the zone of soil heating, only a proportion of available seeds had dormancy broken by smoke alone. At a third site (no residual seed bank detected) the smoke cue was predicted not to have declined with soil depth. Up to 900 species (just under half the fire-prone flora) in the study region are thought to have seed dormancy broken by the interaction of heat and smoke during the passage of a fire.     

Evidence for physiological seed dormancy cycling in the woody shrub Asterolasia buxifolia and its ecological significance in fire‐prone systems

• Dormancy cycling is a key mechanism that contributes to the maintenance of long‐term persistent soil seed banks, but has not been recorded in long‐lived woody shrub species from fire‐prone environments. Such species rely on seed banks and dormancy break as important processes for post‐fire recruitment and recovery.
• We used germination experiments with smoke treatments on fresh seeds and those buried for 1 year (retrieved in spring) and 1.5 years (retrieved the following late autumn) to investigate whether Asterolasia buxifolia, a shrub from fire‐prone south‐eastern Australia with physiologically dormant seeds, exhibited dormancy cycling.
• All seeds had an obligation for winter seasonal temperatures and smoke to promote germination, even after ageing in the soil. A high proportion of germination was recorded from fresh seeds. but germination after the first retrieval was significantly lower, despite high seed viability. After the second retrieval, germination returned to the initial level. This indicates a pattern of annual dormancy cycling; one of the few observations, to our knowledge, for a perennial species. Additionally, A. buxifolia’s winter temperature and smoke requirements did not change over time, highlighting the potential for seeds to remain conditionally dormant (i.e. restricted to a narrow range of germination conditions) for long periods.
• For physiologically dormant species, such as A. buxifolia, we conclude that dormancy cycling is an important driver of successful regeneration, allowing seed bank persistence, sometimes for decades, during fire‐free periods unsuitable for successful recruitment, while ensuring that a large proportion of seeds are available for recruitment when a fire occurs.