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.     

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.

     

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.     

Diversity of germination strategies and seed dormancy in herbaceous species of campo rupestre grasslands

The effects of fire on the vegetation vary across continents. However, in Neotropical fire‐prone grasslands, the relationship between fire and seed germination is still poorly understood, while their regeneration, especially after strong anthropogenic disturbance, is challenging for their conservation. In the present study, we assessed diversity of germination strategies in 15 dominant herbaceous species from Neotropical altitudinal grasslands (locally known as campos rupestres). We exposed seeds to several fire‐related treatments. We also compared germination between regularly and post‐fire fruiting species. Finally, we investigated the diversity of dormancy classes aiming at better understanding the biogeography and phylogeny of seed dormancy. Germination strategies varied among families. Velloziaceae and Xyridaceae produced non‐dormant, fast‐germinating seeds. Cyperaceae and Poaceae showed an extremely low or null germination due to a high proportion of unviable or embryo‐less seeds. The seeds of campo rupestre grasslands are fire resistant, but there is no evidence that fire triggers germination in this fire‐prone ecosystem. Although heat and charred wood did not promote germination, smoke enhanced germination in one grass species and decreased the mean germination time and improved synchrony in Xyridaceae and Velloziaceae. Fire had a positive effect on post‐fire regeneration by stimulating fruit set in some Cyperaceae and Poaceae species. These species produced faster germinating seeds with higher germination percentage and synchrony compared to regularly fruiting Cyperaceae and Poaceae species. This strategy of dispersion and regeneration seems to be an alternative to the production of seeds with germination triggered by fire. Physiological dormancy is reported for the first time in several clades of Neotropical plants. Our data help advance the knowledge on the role of fire in the regeneration of Neotropical grasslands.     

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.     

Combined effects of heat shock, smoke and darkness on germination of Epacris stuartii Stapf., an endangered fire-prone Australian shrub

Epacris stuartii (Epacridaceae) is an endangered heathland shrub in which seedling recruitment occurs almost exclusively after fire. Seed viability and multiple seed dormancy mechanisms were examined to explore why levels of seedling recruitment were low after some fires, despite high rates of pre-fire seed production. The individual and combined effects of heat shock, smoke derivatives and darkness on germination were tested in the laboratory using an orthogonal logit-linear model, a form of analysis ideally suited to hypotheses concerning multiple germination cues. Seed viability (56%) was found not to be limiting. After 89 days of incubation, germination was significantly enhanced by heat shock, smoke treatment and continuous darkness but there were no significant interactions. These effects were equal in magnitude and additive such that maximum germination (42% of viable seeds) was stimulated when all three treatments were combined, significantly less germination occurred in response to any two treatments combined (22–23%), any single treatment stimulated germination at levels (10–11%) significantly less than two treatments and the lowest levels of germination occurred among untreated seeds (4%). Relative to the untreated control, germination was stimulated by smoke derivatives in high concentrations but not in low concentrations. The effect of darkness diminished with time so that after 270 days of incubation darkness had no significant effect on germination, while heat and smoke still enhanced germination significantly, additively and interchangeably. More prolonged germination of seeds exposed to light on the soil surface than buried seed may spread the risk of desiccation of emerging seedlings over multiple rainfall events. The role of heat and smoke derivates suggests that fire management is a crucial tool for influencing seedling recruitment and hence the survival of E. stuartii at its only known location. Received: 3 November 1996 / Accepted: 15 June 1997     

Seed set in Sorghum stipoideum,and not fire,determines the timing of breeding by Gouldian finches (Erythrura gouldiae)

Seeds of native grasses are an important food source for granivorous finches throughout the tropical savannas of northern Australia. The iconic Gouldian finch (Erythrura gouldiae), a threatened species endemic to these savanna grasslands, relies almost exclusively on the grass seeds of annual Sorghum spp. when breeding, and appears to time breeding with the availability of these seeds. Fire is common throughout the savanna grasslands of northern Australia and has the potential to alter Sorghum spp. seed germination and plant reproductive phenology which in turn could alter the Gouldian finch breeding phenology window. This study examines if the temporal relationship between breeding by Gouldian finches in the north‐east Kimberley region of Australia relates to fire and Sorghum stipoideum seed phenology. The availability of S. stipoideum seed was monitored at Gouldian finch breeding sites in association with local fire history and Gouldian finch breeding data. The effect of experimental fire (heat and smoke) on S. stipoideum seed dormancy and germination was also investigated. We found that the first heavy rainfalls preceded germination of S. stipoideum in November/December. Synchronous seed set by S. stipoideum plants the following March/April coincided with the start and duration of the Gouldian finch breeding season. The timing of dry season fires had no relationship with seed dormancy or the phenology of seed germination and seed set. Nor did the effects of smoke or heat affect seed dormancy and germination. These findings support the importance of the seed phenology of annual grass species, such as S. stipoideum, to breeding Gouldian finches, and also suggest that the occurrence of fire at a breeding site in the previous year does not alter the S. stipoideum seed or finch breeding window. The implications of these results for threatened Gouldian finch ecology and management are discussed in relation to previously published fire impacts on S. stipoideum seed nutrition and finch breeding site selection.     

Effects of aqueous smoke solutions and heat on seed germination of herbaceous species of the Sudanian savanna-woodland in Burkina Faso

Smoke generated by burning of plant materials has widely been recognized as a germination cue for some species from both fire prone and fire-free ecosystems. It is an important factor for the understanding of vegetation dynamics and could have potential use for ecological management and rehabilitation of disturbed areas. Individual species, however, seem to have a specific requirement for the type and dosage of smoke treatments. In the present study, six different concentrations of smoke solution were tested on 13 herbaceous species by soaking the seeds for 24 h. The germination of a forb species, Borreria scabra, was significantly stimulated (p<0.05) by the smoke treatment while that of the annual grass species, Euclasta condylotricha, was significantly inhibited (p<0.05) by 100% smoke solution treatment. Contrary to our expectation that another fire-related cue, heat shock, would break the physical dormancy of the species tested, it was not promotive. For non-dormant seeds of B. scabra and Borreria radiata, high temperatures were lethal while low temperature induced physiological dormancy that was overcome in the former species within 30 days of the germination trial. For some species, responses to smoke did not corroborate with the field-observed response to fire, making ecological interpretation challenging. For responsive species, the smoke treatment could be a simple approach for promoting their re-establishment in areas where it is needed. More investigations are needed to assess the spread of response to smoke.     

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.

     

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.     

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.     

Predicting patterns of post-fire germination in 35 eastern Australian Fabaceae

Germination in 35 species from 15 legume genera of southeastern Australia was promoted by a heat treatment which broke the seed coatcaused dormancy. Once the critical temperature was reached, most seeds had their dormancy broken, independent of the duration of heating. Species fell into three classes according to whether their dormancy was broken by a temperature of 40, 60 or 80°C. Highest germination in all species was achieved by heating in the temperature range 80–100°C, although long durations (120 min) at 100°C caused seed death in several species. At 120°C, seeds of most species were killed at all but one minute's duration. A proportion of seeds from 7 species (Acacia myrtifolia, Pultenaea daphnoides, P. incurvata, P. linophylla, P. polifolia, Dillwynia floribunda and Sphaerolobium vimineurn) was not killed at 120°C and had their dormancy broken. This proportion varied markedly and resultant germination levels were significantly less than those at 80 and 100°C, except in S. vimineum. Between-site variations in the 4 species tested (A. myrtifolia, A. suaveolens, A. terminalis and A. ulicifolia) were small. These variations concerned: (i) the minimum temperature required to break seed dormancy in 2 species: 60°C in one population of A. myrtifolia and A. suaveolens, and 80°C in the other; and (ii) the intensity of the germination response. Duration of heating was less important than temperature as a determinant of germination. Ordination techniques revealed that results from one duration across temperatures were comparable with data from multiple durations. This has significant applications in studying rare species, where seed may be in short supply. Predicted germination levels after a moderate intensity fire should far exceed those after a low intensity fire. Little germination was predicted for many species after a low intensity fire and for one species, A. elongata, no germination was predicted. The potential role of indicator species in relation to the maintenance of species in a community is suggested.     

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.     

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.     

The relationship between seed dormancy,seed size and weediness,in Crepis tectorum (Asteraceae)

Summary I examined the germination characteristics of weed and outcrop populations of Crepis tectorum to test the hypothesis that the presumably more ephemeral weed habitat favors the highest levels of seed dormancy. The winter annual habit characterizing most plants of this species was reflected in a rapid germination of seeds sown in late summer. A slightly higher fraction of surface-sown seeds of weed plants delayed germination. Buried seeds of weed plants also survived better than seeds produced by plants in most outcrop populations, supporting the idea that weediness favors seed dormancy and a persistent seed bank. However, the differences in seed dormancy between the two ecotypes were small and not entirely consistent. Furthermore, high levels of seed dormancy were induced during burial in the outcrop group, suggesting that there is a potential for a dormant seed population in this habitat as well. Demographic data from one of the outcrop populations verified the presence of a large between-year seed bank. Possible environmental factors favoring seed dormancy in outcrop populations are discussed. The unusually large seeds of weedy Crepis contrasts with the relatively small difference in seed dormancy between the two ecotypes.     

Heat increases germination of water-permeable seeds of obligate-seeding <Emphasis Type="Italic">Darwinia</Emphasis> species (Myrtaceae)

We examined the response of seeds to heat in four geographically restricted and one widespread species of shrubby Darwinia from the fire-prone region of southeastern Australia. These shrubs are killed by fire and rely on seed germination after a fire to maintain populations. We replicated the germination trials across several sites and several fruiting seasons for most species. Seeds had a high level of viability and were largely dispersed in a dormant state, except in D. glaucophylla, where seed dormancy varied significantly across fruiting seasons. The indehiscent fruit of all species readily imbibes moisture when wet and seeds are not considered to be ‘hard-seeded’. All species had increased seed germination in response to a limited range of heating temperatures (generally 80–100°C). Higher temperatures killed increasing proportions of seeds. This pattern was broadly consistent across species, population and seasons, although the proportion of seeds whose germination was promoted by heat varied from high (D. diminuta, D. fascicularis, D. glaucophylla) to moderate (D. biflora, D. procera). Our work highlights the importance of heat as a mechanism for influencing germination in species that are not hard-seeded. Consequently, soil temperatures during a fire should strongly influence post-fire germination levels in Darwinia. The roles of other cues that promote germination, i.e. smoke, seasonal temperatures and their interactions with heat, remain to be investigated.     

Effects of low temperatures on the loss of innate dormancy and the development of induced dormancy in seeds of Rumex obtusifolius L. and Rumex crispus L.

Abstract It is possible to remove the innate dormancy of seeds of Rumex crispus L and Rumex obtusifolius L. by an initial period of low-temperature stratification, providing the seeds are then transferred to a higher temperature. The lower the initial temperature within the range 1.5°-15°C, the greater the germination; there is no stratification effect at 20°C. Although 10°C and 15°C were shown to be suitable for both stratification and for the process of germination itself, neither temperature results in any germination if given constantly: a change from a lower to a higher temperature is essential. The optimum period for stratification depends on two separate processes which occur during the treatment–a rapid loss of innate or primary dormancy and a slower development of induced or secondary dormancy. Within the range 1.5°-15°C the rate of loss of innate dormancy appears to be independent of light and is probably independent of temperature. In contrast, the rate of induction of secondary dormancy increases with increase in temperature, and is more rapid in the dark than the light. The rate of induction of secondary dormancy during stratification is greater in R. crispus than in R. obtusifolius. As a consequence, maximum germination was obtained in both species after stratification at 1.5°C in the light, the optimum period of treatment being about 4 weeks in R. Obtusifolius and 6 weeks in R. crispus, while the maximum germination obtained and the optimal period of stratification decrease in both species with increase in stratification temperature.     

Seed germination of the carnivorous plant Byblis gigantea (Byblidaceae) is cued by warm stratification and karrikinolide

Byblidaceae is one of the most poorly studied carnivorous plant families, with seed dormancy and germination biology remaining unresolved. This knowledge deficit has significant conservation and management implications, particularly as the most southerly distributed species, the south‐west Western Australian endemic Byblis gigantea, is listed as critically endangered. This study examined the ecophysiology of seed dormancy and germination in B. gigantea in concert with a study of the population dynamics of a single plant community. Mass seedling emergence and plant establishment were observed after a wildfire in 2007 in the natural population, followed by a rapid decline in mature individuals (91%) over a 4‐year monitoring period, with almost no inter‐fire recruitment (1.4% of all emergence) observed. Seeds possessed a fully developed embryo, and the germination characteristics of fresh seeds classified them as showing physiological dormancy. Seed dormancy was partially alleviated by warm stratification (30 °C) for 8 weeks prior to incubation at 15 °C, with c. 40% germination observed. With the additional exposure of seeds to the germination‐active chemical in smoke, karrikinolide, the germination of warm‐stratified seeds increased to 89%. Seeds also displayed orthodox storage behaviour and appeared to be amenable to long‐term seed banking for conservation. These results present the first observation of the stimulation of the germination of a carnivorous plant by a smoke‐derived compound. © 2013 The Linnean Society of London, Botanical Journal of the Linnean Society, 2013, 172 , 143–152.     

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.     

The role of Rhytidoponera metallica (Hymenoptera,Formicidae) in facilitating post‐fire seed germination of three ant‐dispersed legume species

The fire avoidance hypothesis proposes that a benefit of seed dispersal by ants (myrmecochory) is to protect seeds from being killed during fire and to facilitate post‐fire germination of seeds that require heat shock to break their physical dormancy. The aim of this study was to quantify the effect of fire and seed burial by a predominant seed‐dispersing ant, Rhytidoponera metallica (subfamily: Ectatomminae) on germination levels of three ant‐dispersed legume species (Pultenaea daphnoides, Acacia myrtifolia and Acacia pycnantha). Experimental burial of seeds within aluminium cans at a site prior to being burnt and at an adjacent unburnt site showed that fire increased germination levels, particularly for seeds buried at 1‐ and 2‐cm deep and that overall, germination levels differed among the three plant species. To quantify seed burial depths and post‐fire germination levels facilitated by R. metallica ants, seeds were fed to colonies prior to fire at the burnt and unburnt sites. Of the seeds buried within nests that were recovered, between 45% and 75% occurred within the upper 6 cm of the soil profile, although unexpectedly, greater percentages of seeds were recovered from the upper 0–2 cm of nests in the unburnt site compared with nests in the burnt site. Germination levels of buried seeds associated with R. metallica nests ranged from 21.2% to 29.5% in the burnt site compared with 3.1–14.8% in the unburnt site. While increased seed germination levels were associated with R. metallica nests following fire, most seeds were buried at depths below those where optimal temperatures for breaking seed dormancy occurred during the fire. We suggest that R. metallica ants may provide fire avoidance benefits to myrmecochorous seeds by burying them at a range of depths within a potential germination zone defined by intra‐ and inter‐fire variation in levels of soil heating.