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.     

Post‐burning germination responses of woody invaders in a fire‐prone ecosystem

It has been frequently recognised that there is a positive feedback between plant invasion and fire underlying invasion success in fire‐prone ecosystems. Accordingly, the response of woody alien species germination to fire may have direct implications on their invasiveness in those ecosystems, particularly when fruit ripening occurs in the fire season. Here, we experimentally evaluated the germination response of some of the main woody invaders of the Chaco Serrano dry woodlands (Gleditsia triacanthos, Cotoneaster glaucophyllus, Ligustrum lucidum, Pyracantha angustifolia and Melia azedarach), which fruit in the seasons of highest fire frequency. Seeds were subjected to heat‐shock treatments that simulated a range of heat intensities, and the species were classified according to their germination response as heat sensitive, tolerant or stimulated. Since Gleditsia triacanthos has indehiscent fruits that fall from the plant and might be exposed to flames, its germination response was also assessed of seeds exposed to fruit burning. Germination responses to heat varied among the invasive species. G. triacanthos seeds experienced increased germination under very low and low heat indexes; it was therefore classified as heat stimulated. The other four species showed no change in germination under very low heat indexes and were therefore considered heat tolerant. However, all species were sensitive to high heat as indicated by their significant decline in germination. G. triacanthos would have limited capacity to recruit from seeds following flaming combustion of its fruits. The prevalence of heat‐tolerant rather than heat‐stimulated germination responses suggests that the occurrence of frequent and seasonal fires in this subtropical savanna system might delay rather than boost the expansion of these invasive species in the system. Yet, the presence of heat‐stimulated germination in one of the studied species warns against generalisation, even within the same ecosystem, and further supports the idiosyncratic nature of invasion success. Abstract in Spanish is available with online material.     

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.     

Endozoochory and Fire as Germination Triggers in Neotropical Dry Forests: an Experimental Test

Endozoochory and fire are crucial ecological factors determining germination success and recruitment in many plant species. Fire is a well‐known germination trigger while endozoochory may allow seed dispersal along with an increase in germination. Their interaction has rarely been addressed, however, even though both factors are pervasive in human‐transformed ecosystems like most Neotropical Dry Forests (NDF). For three common Mesoamerican tree species (Acacia pennatula, Enterolobium cyclocarpum, and Guazuma ulmifolia), we used feeding trials to assess the preference of cattle, which are their main seed dispersal agent. We also experimentally tested the interaction between gut passage and fire as triggers of germination. The fruits of the three species were eaten by cattle, but the small seeds of G. ulmifolia were ingested 10‐fold more than those of the other species. While gut passage did not have any effect on germination, heat‐shocks above 90 °C increased the number of germinating seeds by 15 percent. These results suggest that cattle may be a key dispersal vector in NDF, but that fire may be an important germination trigger. Physical dormancy in these species may have been selected for by extinct megaherbivores because it was a key trait ensuring seed survival after gut passage. However, in light of the recent expansion of cattle‐ranching and fire occurrence in NDF, it has become a useful exaptation facilitating the colonization of disturbed areas.     

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.     

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.     

Comparative effects of desiccation,heat shock and high temperatures on seed germination of savanna and forest tree species

Although forest and savanna biomes predominate in tropics regions, the factors that control their distribution remain unclear. South American savannas occur in regions that are considered warm and humid enough to support forests, indicating that agents other than climate determine the occurrence of one or the other physiognomy. Herbivory, fire and water deficit have been considered environmental filters that limit the forest species encroachment in savanna physiognomies, but the effects of these filters on the capability of these species to recruit from seeds remain poorly understood. In this study we investigated how stress factors characteristic of savanna environments, such as soil desiccation, heat shocks and high temperatures affect the survival and germination of seeds from savanna and forest tree species. We found that desiccation (to 5%) reduced the germination percentage of forest seeds, but had no effect on the germination of savanna seeds. Forest seeds were less tolerant to heat shocks of 140°C and 200°C, and showed lower germination percentage at temperatures of 35 and 40°C, when compared with savanna seeds. Savanna seeds presented longer germination times and higher germination variance than forest seeds, indicating a risk‐spreading germination strategy among savanna species. The low tolerance of forest seeds to desiccation, heat shock and high temperatures may explain the low recruitment of forest trees into savanna physiognomies. Climate change models predict lower soil moisture, higher temperatures and higher fires frequency for South America biomes. Our results suggest that savanna species are likely to be more capable of withstanding the effects of these changes than forest 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.     

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.     

Seed germination cues and the importance of the soil seed bank across an environmental gradient in the Serengeti

Soil seed banks are an important source of new individuals for many plant populations and contribute to future genetic variability. In general, the size and persistence of soil seed banks is predicted to be greater where growth occurs in unpredictable pulses, where opportunities for disturbance‐related recruitment are frequent and where the probability of recruitment failure is high. In savanna ecosystems, characterized by disturbance from fire and unpredictable water availability, soil seed banks should be relatively important sources of recruitment. However, the few studies conducted in savannas are inconclusive about the importance of soil seed banks and, more specifically, how seed banks should change across environmental gradients. We determined the number of viable seeds in the soil seed bank across savanna‐grasslands in the Serengeti, an ecosystem characterized by frequent fire and seasonal drought. Soils were exposed to a combination of smoke and heat, cues which may be required to break seed dormancy in such ecosystems. Our a priori expectation was to observe large seed banks in regions characterized by seasonal drought and comparatively smaller seed banks in regions of higher moisture availability and high fire frequencies. In contrast to our hypothesis, seed germination increased strongly with precipitation and fire frequency. In addition, there was a significant interaction effect between fire and rainfall: low rainfall sites with frequent fire had greater seed germination than low rainfall sites with low fire frequency. Moreover, in laboratory experiments, heat had a negative, smoke a positive effect on final seed germination numbers. Together, these findings suggest that fire may be a key factor in driving herbaceous seed bank dynamics in tropical savannas.     

Heat and smoke pre‐treatment of seeds to improve restoration of an endangered Mediterranean climate vegetation type

Invasive alien plants impact ecosystems, which often necessitates their removal. Where indigenous species recovery fails following removal alone, an active intervention involving reintroduction of seed of native species may be needed. This study investigated the potential for a combination of the fire cues of smoke and heat as a pre‐treatment of seeds in breaking dormancy and facilitating increased germination. Species were selected to represent different functional types within Cape Flats Sand Fynbos; a fire‐prone, critically endangered vegetation type in South Africa. Seeds were exposed to either a heat pulse (temperatures between 60 and 300°C for durations of between 30 s and 20 min) or dry after‐ripening (1 or 2 months at milder temperatures of 45°C or less). Thereafter, seeds were soaked in smoke solution for 18 h and subsequently placed on agar at 10/20°C for germination. Most species fell into one of two main groups: Seed germination in the first group was greatest following a lower temperature (60°C) heat pulse, an extended period of mild temperature (20/40°C or 45°C) exposure, or no pre‐treatment with heat. Seed germination in the second group was promoted after brief exposure to higher (100°C) temperatures. No germination occurred in any species following heat treatments of 150°C or higher. Species which responded better to higher temperatures were mainly those possessing physical dormancy, but seed morphology did not correlate with germination success. This study showed that heat stimulation of seeds is more widespread in fynbos plant families than previously known and will enable the development of better seed pre‐treatment protocols before large‐scale sowing as an active restoration treatment after alien plant clearing.     

Influence of heat shock on seed germination of plants from regularly burnt savanna woodlands and grasslands in Ethiopia

The effect of heat shock on the germination of seeds of 21 plant speciesfrom fire-prone wooded savanna ecosystems in western Ethiopia was analysed inorder to examine the possible implications of fire upon plant regenerationafterthis disturbance. Seeds were subjected to 6 different heat intensities (20, 60,90, 120, 150 and 200°C) for 1 or 5 minutes, in ordertosimulate the situation in the upper soil layers or on the soil surface duringfires. Germination tests were carried out in pots in a greenhouse over 20weeks.After 9 weeks no more seedlings emerged. There was wide interspecific variationin the responses of seeds to the different treatments. In all species,germination was significantly affected by the temperature treatment level.Shortexposure of seeds to high temperatures generally stimulated germination whereasprolonged exposure reduced seed germination. However, some species eventolerated 5 min treatment at 200°C. Seedheat resistance was positively correlated with seed length and mass among thespecies. Hence, production of large seeds with protective tissues promotessurvival in fire-prone savanna areas. Also, the seeds of some species showedboth a low and a high temperature optimum which ensures that at least someseedsgerminate in the absence of fire, but also that viable seeds still remain ifsubsequent late fires kill emerging seedlings. Frequent and light burning inwooded savanna grasslands seems to stimulate and enhance germination of most ofthe studied plant species.     

Heat shock effects on seed germination of five Brazilian savanna species

Fire is considered an important factor in influencing the physiognomy, dynamics and composition of Neotropical savannas. Species of diverse physiognomies exhibit different responses to fire, such as population persistence and seed mortality, according to the fire frequency to which they are submitted. The aim of this study is to investigate the effects of heat shocks on seed germination of Anadenanthera macrocarpa (Benth.) Brenan, Dalbergia miscolobium Benth., Aristolochia galeata Mart. & Zucc., Kielmeyera coriacea (Spreng.) Mart. and Guazuma ulmifolia Lam., which are native species of the Brazilian savanna. The temperatures and exposure times to which the seeds were submitted were established according to data obtained in the field during a prescribed fire: 60 °C (10, 20 and 40 min), 80 °C (5, 10 and 20 min) and 100 °C (2, 5 and 10 min). Untreated seeds were used as controls. Seeds of A. galeata and K. coriacea showed high tolerance to most heat treatments, and seeds of A. macrocarpa showed a significant reduction in germination percentage after treatments of 80 °C and 100 °C. Treatments of 100 °C for 10 min reduced germination percentage for all species except G. ulmifolia, which has dormant seeds. For this species, germination was accelerated by heat treatments. The high temperatures applied did not interfere with the time to 50% germination (T₅₀) of the tolerant seeds. Seeds of the savanna species K. coriacea and A. galeata were more tolerant to heat shocks than seeds of the forest species A. macrocarpa. Guazuma ulmifolia, the forest species with seeds that germinate after heat shock, also occurs in savanna physiognomies. Overall, the high temperatures applied did not affect the germination rate of the tolerant seeds.     

Seed dispersal kernel of the largest surviving megaherbivore—the African savanna elephant

Owing to the late Pleistocene extinctions, the megafauna of Europe, Australia and the Americas disappeared, and with them the dispersal service they offered megafaunal fruit. The African savanna elephant, the largest remaining megaherbivore, offers valuable insights into the seed dispersal services provided by extinct megafauna in prehistoric times. Elephant seed dispersal studies have for the most part concentrated on African and Asian forest elephants. African savanna elephants are morphologically distinct from their forest counterparts. Like the forest elephants they consume large quantities of fruit from a large number of tree species. Despite this little is known of the savanna trees that rely on elephants for their dispersal or the spatial scale at which these seeds are dispersed. We combined information from feeding trials conducted on four park elephants with field telemetry data from 38 collared elephants collected over an 8‐year period in APNR/Kruger National Park to assess the seed dispersal service provided by savanna elephants. This study provides the first detailed account of the spatial scale at which African savanna elephants disperse seeds. Our mechanistic model predicts that 50 percent of seeds are carried over 2.5 km, and distances up to 65 km are achievable in maximum gut passage time. These findings suggest the savanna elephant as the longest distance terrestrial vertebrate disperser yet investigated. Maintaining their ecological role as a seed disperser may prove a significant factor in the conservation of large‐fruited tree diversity within the savannas. These results suggest that extinct megafauna offered a functionally unique dispersal service to megafaunal fruit.     

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 production and germination in two rare and three common co‐occurring Acacia species from south‐east Australia

Abstract Seed set, size, viability and germination requirements were investigated for two rare (Acacia ausfeldii and A. willianisonii) and three common (A. pycnantha, A. genistifolia and A.paradoxa) co‐occurring congeners in box‐ironbark eucalypt forests near Bendigo, south‐east Australia to investigate correlates of rarity. Seed size was significantly smaller for the two rare species and germinants were less able to emerge from deeper sowing depths than were the larger seeded common congeners. All species had a strong heat‐stimulated germination response. While the rare A. ausfeldii showed strong germination only at the highest temperature treatment (100°C), the common and widespread A.pycnantha showed strong germination across a broad range of temperatures (60‐100°C), likely to be experienced by soil‐stored seeds during a fire. Seed viability, number of seeds per plant, and number of firm, aborted and eaten seeds per pod varied between species, but the pattern of variation was not related to rarity. Small seed size and a very specific temperature requirement for germination may help to explain rarity in A. ausfeldii, and to a lesser extent in A. willianisonii. Fires are often patchy and heating of the soil is likely to be highly spatially variable, so species with germination responses to a broad range of temperatures have an advantage over those that respond only to a narrow range. A narrower range of soil depths from which seeds can emerge will further reduce the proportion of the seed bank that might recruit following fire. Human impacts on species habitats, such as fragmentation, loss of topsoil through mining, timber harvesting, grazing and urbanization, and consequent reduction in fire intensity, are likely to have further contributed to rarity in these species. The role of pollination and other factors in relation to population size is the subject of further investigation.     

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.

     

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.     

Experimental determination of seed emergence and carry‐over in the soil seed bank of the herbaceous perennial,Trachymene incisa (Apiaceae)

Abstract Many populations of herbaceous perennial plants contain seeds stored in a soil seed bank. The contribution of seeds to population persistence is an important parameter in population models but germination rates of known‐age seeds are difficult to obtain because individual seeds cannot easily be followed. Although Trachymene incisa Rudge plants produce copious seeds that are dispersed into the soil, the existence of a seed bank has not been confirmed. To quantify the potential for a seed bank fresh seeds of T. incisa were sown into experimental seed banks in the eucalypt‐dominated Agnes Banks Woodland in western Sydney, NSW. A recent fire provided the opportunity to compare germination in the burnt and unburnt vegetation. Density of seed sowing and time of maturation/dispersal of seeds were manipulated in 75 seed cages. Emergence of seeds after 5 months was significantly higher for the earliest planting date but after 1 year, germination of seeds planted in the later weeks increased, and the final germination for all weeks was 28%. Density of sowing and the recent fire did not affect emergence. A second experiment planted over a broader time span (9 weeks instead of 3 weeks) confirmed the effect of planting date but also found significant spatial variation on a scale of tens of metres. Laboratory germination rates of over 70% confirmed that the seeds were viable and non‐dormant when sown in the field cages. The carry‐over of non‐germinated seed in the soil seed bank is estimated to be about 70% after 2 years, implying that a cohort of seeds would not be depleted through germination alone for up to 40 years. The potential for a long‐lived seed bank in this species is interesting because the plants are also capable of resprouting from their rootstock after fire, giving them characteristics of both resprouters and seeders.     

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.