Smoke and heat effects on soil seed bank germination for the re‐establishment of a native forest community in New South Wales

The effects of plant‐derived smoke and of heat on the emergence of seedlings from seeds were assessed. Seeds had been stored in forest topsoil used for mine site rehabilitation. The study was carried out in a dry sclerophyll, spotted gum (Corymbia maculata), forest community at the Mount Owen open‐cut coal mine in the Hunter Valley region of New South Wales. Samples of the surface 2.5 cm of topsoil were either exposed to cool smoke from eucalypt foliage for 60 min, heated to 80°C, or left untreated. Seedling emergence from the seed bank in this soil was then monitored in a glasshouse. Within the first month, smoke alone promoted a 4.3‐fold increase in the density of seedlings relative to control. There were 540 emergents per m² in the control and 2309 per m² in the smoke treated topsoil. Many annual and perennial herbs emerged but grasses responded most strongly to smoke. Germination in seven of the 20 grass species was promoted by smoke. Smoke promoted the germination of some introduced species as well as native species, and accelerated the rate at which seedlings emerged, although these differences sometimes declined with time. Heat also stimulated germination but smoke and heat stimuli appeared to be complementary in their promotion of seedling emergence from the topsoil seed bank. Each treatment increased the density of different species, enhanced the species richness of different components of the seed bank, and had different effects on the rate of emergence. The results suggest that increased seed germination in the field immediately following a moderate intensity fire may sometimes be the result of smoke stimulation and sometimes the result of heat stimulation of the soil seed bank. These findings may have important implications for minesite revegetation programs where topsoils are replaced after mining and rapid germination of seeds stored in these soils is required during short periods when conditions are favourable for germination.     

The independent effects of heat,smoke and ash on emergence of seedlings from the soil seed bank of a heathy Eucalyptus woodland in Grampians (Gariwerd) National Park,western Victoria

Abstract The independent effects of smoke, ash, and wet and dry heat treatments on seedling emergence from the soil seed bank were tested for soils from fire-prone heathy woodlands in western Victoria. A total of 763 individuals from 56 species were recorded from the surface soil samples (which covered a total area of 1 m²). Both species richness and density of seedlings was greater for smoke- and heat-treated samples than for controls and ash-treated samples. However, only the density differences were significant. Mean seed bank densities for the smoke and heat treatments ranged from 855 ± 70m^?2 to 1080 ± 58 m^?2 and are similar to estimates obtained elsewhere in Australia for heat-treated soils from dry sclerophyll communities. Of the 56 species recorded, 46 occurred in the smoke and heat treatments but only 33 in the control and ash treatments. The sudden increase in surface soil pH, exchangeable cations and extractable phosphorus which was associated with the ash treatment did not act as a trigger for germination in any of the species recorded here. Chemical constituents from smoke do appear to provide a stimulus separate from the effects of heat, but were not identified with any particular taxa.     

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.     

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.     

Effect of flooding and draw‐down disturbance on germination from a seashore meadow seed bank

Abstract. The objective of this study was to investigate the effects of flooding and draw‐down on the germination from the coastal grassland seed banks and to determine whether the effect of flooding varies between the delta and the seashore. Seed bank samples were collected from three shore transects in SW Finland, two on the shore of the Baltic Sea and one on the delta of River Kokemäenjoki. Samples were germinated in non‐flooded and flooded conditions for over a month, after which both treatments were maintained in non‐flooded conditions. A total of 9267 seedlings of 47 species germinated and mean density of seeds in the soil was ca. 84 000/m². Most of the seedlings were monocots (98%) and perennials (98%). Ca. 30–40% of the species found in the above‐ground vegetation had a seed bank including the majority of the most abundant species. The number of seeds and species richness increased as the organic layer became thicker. The organic layer was thicker in the seashore samples and the seed bank was significantly larger than in the delta. The flooding and draw‐down treatment significantly increased the number of germinating seedlings in the seashore and also increased species richness in two transects, one in the delta and the other in the seashore. Two species, Schoenoplectus tabernaemontani and Typha latifolia, had significantly higher germination in the flooded treatment than in the non‐flooded. Apparently, many species in these coastal grasslands have adapted to flood disturbance and for seeds of some species flooding may work as a positive signal, possibly breaking dormancy.     

Tree seed rain and seed removal,but not the seed bank,impede forest recovery in bracken (Pteridium aquilinum (L.) Kuhn)‐dominated clearings in the African highlands

Considerable areas dominated by bracken Pteridium aquilinum (L.) Kuhn occur worldwide and are associated with arrested forest recovery. How forest recovery is impeded in these areas remains poorly understood, especially in the African highlands. The component processes that can lead to recruitment limitation—including low seed arrival, availability and persistence—are important determinants of plant communities and offer a potential explanation for bracken persistence. We investigated key processes that can contribute to recruitment limitation in bracken‐dominated clearings in the Bwindi Impenetrable National Park, Uganda. We examined if differences in seed rain (dispersal limitation), soil seed bank, or seed removal (seed viability and persistence) can, individually or in combination, explain the differences in tree regeneration found between bracken‐dominated areas and the neighboring forest. These processes were assessed along ten 50‐m transects crossing the forest–bracken boundary. When compared to the neighboring forest, bracken clearings had fewer seedlings (bracken 11,557 ± 5482 vs. forest 34,515 ± 6066 seedlings/ha), lower seed rain (949 ± 582 vs. 1605 ± 335 tree seeds m^?2 year^?1), comparable but sparse soil seed bank (304 ± 236 vs. 264 ± 99 viable tree seeds/m²), higher seed removal (70.1% ± 2.4% vs. 40.6% ± 2.4% over a 3‐day interval), and markedly higher rodent densities (25.7 ± 5.4 vs. 5.0 ± 1.6 rodents per 100 trapping sessions). Camera traps revealed that rodents were the dominant animals visiting the seeds in our seed removal study. Synthesis: Recruitment limitation contributes to both the slow recovery of forest in bracken‐dominated areas, and to the composition of the tree species that occur. Low seed arrival and low persistence of unburied seeds can both explain the reduced density of seedlings found in bracken versus neighboring forest. Seed removal, likely due to rodents, in particular appears sufficient to constrain forest recovery and impacts some species more severely than others.     

Influence of multiple fire‐related germination cues on three Sydney Grevillea (Proteaceae) species

Abstract Many species found in fire‐prone habitats that possess a soil‐stored seedbank only recruit seedlings in large numbers following a fire. Fire‐related germination cues are presumably used by these seeds as signals that a fire has occurred, and would include the heating that occurs in the soil and the combustion products of burning vegetation, smoke and charcoal. Three Sydney species, Grevillea buxifolia (Sm.) R. Br., Grevillea sericea (Sm.) R. Br. and Grevillea speciosa (Knight) McGillivray, were studied for the interactive effects of these cues on their germination. The germination of all species was found to increase with both smoke and heat treatments. While smoke always had a greater influence than heat, the relationship between the two treatments varied with species. The presence of two fire‐related germination cues should allow these species to take better advantage of the recruitment opportunities of the post‐fire environment.     

Comparative effects of different smoke treatments on germination of Australian native plants

Abstract This study consists of a series of investigations designed to determine the potential of concentrated smoke products to stimulate seed germination and assist in the rehabilitation of degraded landscapes. Aerosol smoke and conventional smoke water products are not practical for use in broad‐scale seed bank regeneration programmes due to infrastructure limitations and the high rate of application required for their use. Concentrated smoke products have been found to be effective in stimulating soil seed banks and are suitable for broad‐scale use due to the lower rate of application required to elicit germination within smoke‐responsive species. The use of concentrated smoke products on undisturbed soil seed banks at 50 and 100 mL m^‐2 increased total seedling numbers 10‐fold, while species richness more than doubled over untreated controls. Although aerosol smoke applied directly to soil promoted 10 times more seedlings compared to concentrated smoke products, species richness levels in each treatment were similar. Concentrated smoke products cause leaf scorch when applied directly over plants in active growth, with limited to nil damage when plants are not actively growing. The affected plants generally recovered following application of concentrated smoke products as a foliar spray.     

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.     

Edge Effects and Seedling Bank Depletion: The Role Played by the Early Successional Palm Attalea oleifera (Arecaceae) in the Atlantic Forest

In this study, we examined the impacts of Attalea oleifera on the structure of seedling bank and discuss potential mechanisms of palm influence. Seed rain, seedling bank, and palm leaf fall were assessed beneath the canopy and in the vicinity of 16 adult palms across the edges of a large fragment (3500 ha) of the Atlantic forest. Moreover, we examined A. oleifera impacts on seed germination and seedling mortality by experimentally submitting seeds and seedlings to prolonged palm-leaf covering. As expected, seedling bank beneath the adults exhibited reduced abundance and species richness at local and habitat scale. Small to large seeds (3.1–30 mm) were underrepresented in the seed rain below adults palms, while experimental leaf covering drastically reduced both seed germination and seedling survivorship. A. oleifera leaf fall occurred over the whole year (3.6±2.7 leaves/individual/yr), which resulted in deep leaf litter mounds (10.7±9.2 cm). Finally, adult palm density (21.6±11.9 individuals/ha) correlated negatively with seedling density across Attalea clusters. Our results suggest that A. oleifera exerts negative effects on the seedling bank by reducing seedling abundance and richness as a consequence of two complementary mechanisms: impoverished and size-biased seed rain plus reduced seed germination and increased seedling mortality due to prolonged covering by fallen leaves.     

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.

     

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.

     

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.     

The effect of concentrated smoke products on the restoration of highly disturbed mineral sands in southeast Victoria

Summary Recent studies have recognized the potential of broad‐scale surface application of smoke compounds for enhancing germination from the soil seed‐bank in fire‐prone vegetation communities. Results suggest that smoke technology may play, in the future, a significant role in the restoration and management of areas supporting indigenous vegetation. An important step in the development of smoke‐based restoration tools is the conduct of in situ field trials in a range of geographical locations and environmental conditions. However, most of the published work on the effectiveness of smoke products in promoting seedbank germination has been conducted at sites in southwestern Australia. The present study examines the effect of commercially available smoke‐water products on the regeneration of a highly disturbed former mine‐site at the Royal Botanic Gardens Cranbourne, in southeastern Victoria. Various combinations of concentrated smoke products and topsoil harvested from a nearby heathy woodland community were applied to exposed, uniform mineral sands to test their effect on seedling density and species richness of regrowth. The trials showed that after 12 months a number of common, herbaceous species including Austrodanthonia setacea, Opercularia varia and Platysace heterophylla were recorded in significantly higher numbers in areas treated with a commercial smoke‐water. However, there was no overall improvement in the density of seedlings or the richness of species as a result of the application of the smoke products. Similarly, total seedling density and species richness were not affected by the addition of topsoil, either alone or in combination with smoke products.     

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.     

The soil seed bank and understorey regeneration in Eucalyptus regnans forest,Victoria

Abstract The soil seed bank and its relation to the extant vegetation in a Eucalyptus regnans F. Muell. forest in the Central Highlands of Victoria were examined. The average seed density was 430 germinable seeds m^?2 to a depth of 2 cm. There was a polynomial regression relationship between the density and species richness of seeds in soil and forest age (0. 6–54 years). Species richness was not significantly different among soil depths (0- 2 , 2- 5 , 5–10 and 10–20 cm) in the forest stand of 54 years old. More seeds germinated from the 5–10 cm depth than from the other depths. Forbs accounted for 73% of the total germinable seeds and there was no germination of E. regnans. The number of species, particularly woody plant species, germinating from the soil seed bank were significantly lower than in the extant vegetation. However, almost all species present in the soil seed bank were present in the vegetation. The soil seed bank provides an important source for the rapid regeneration of understorey vegetation following clear-cutting and slash-burning in the E. regnans forest. The rapid understorey establishment may play an important role in protecting soil from erosion, in nutrient conservation, replacement and redistribution. The soil seed bank may also be a necessary source of maintaining genetic diversity in the forest over the long term.     

Depth distribution and composition of seed‐banks in alien‐invaded and uninvaded fynbos vegetation

South African fynbos vegetation is threatened on a large scale by invasive woody plants. A major task facing nature conservation managers is to restore invaded areas. The aim of this study was to determine the restoration potential of fynbos following dense invasion by the Australian tree Acacia saligna. The impacts of dense invasion on seed‐bank composition and depth distribution were investigated to determine which fynbos guilds and species have the most persistent seed‐banks. Soil samples were excavated at three different depths for invaded and uninvaded vegetation at two sand plain and mountain fynbos sites. Seed‐banks were determined using the seedling emergence approach. Invasion caused a significant reduction in seed‐bank density and richness at all sites. There was a significant, but smaller, reduction in seed‐bank density and richness with soil depth at three sites. Seed‐bank composition and guild structure changed following invasion. Low persistence of long‐lived obligate seeders in sand plain fynbos seed‐banks indicates that this vegetation type will be difficult to restore from the seed‐bank alone following alien clearance. The dominance of short‐lived species, especially graminoids, forbs and ephemeral geophytes, suggests that regenerating vegetation will develop into a herbland rather than a shrubland. It is recommended that seed collecting and sowing form part of the restoration plan for densely invaded sand plain sites. As seed density remained higher towards the soil surface following invasion, there is no general advantage in applying a mechanical soil disturbance treatment. However, if the shallow soil seed‐bank becomes depleted, for example following a hot fire through dense alien slash, a soil disturbance treatment should be given to exhume the deeper viable seed‐bank and promote recruitment.     

Influence of fire on critically endangered Swartland Shale Renosterveld in the Cape Floristic Region

Questions

The degree to which renosterveld shrublands are fire‐dependent is currently unclear. To address this issue, the following questions were asked: (1) does smoke stimulate germination of soil‐stored seeds in renosterveld; (2) does recently‐burned renosterveld display changed composition and higher diversity than unburned vegetation; and (3) how do the species compositions of renosterveld soil seed banks and standing vegetation compare?

Location

Swartland, Cape Floristic Region, South Africa.

Methods

Soil seed bank samples from a north‐ and south‐facing slope were smoke‐treated and germinated to test for smoke‐stimulated germination. Burned standing vegetation was surveyed 16 months post‐fire, as was unburned vegetation on the same slopes. Seed bank species richness and density were compared between smoke‐treated and untreated samples within and between slopes. Burned and unburned standing vegetation were compared within and between slopes in terms of species richness, abundance and aerial cover. Compositional similarity of the seed banks and standing vegetation was assessed.

Results

Seed banks were dominated by annuals and graminoids. Smoke treatment had no effect, except for driving significantly higher species richness and seedling density in south‐facing slope perennial shrubs. Species richness and seedling density were significantly higher in seed banks on the south‐facing slope compared to the north‐facing slope. Burned standing vegetation exhibited significantly higher diversity than unburned vegetation. Annuals and graminoids displayed significantly higher species richness and aerial cover in burned renosterveld. The north‐facing slope contained less than half the number of species/m² compared to the south‐facing slope. The seed banks and standing vegetation showed low to intermediate similarity (Sørensen = 31%–53%), but grouped close together on an NMDS plot, suggesting intermediate similarity overall.

Conclusions

Elevated germination of perennial shrubs in smoke‐treated seed bank samples and increased diversity of post‐fire standing vegetation suggest the renosterveld in this study shows elements of a fire‐driven system. Certain species only recruited in burned sites, suggesting fire‐stimulated germination. Aspect had a major influence on plant community composition, with the mesic south‐facing slope being more diverse than the xeric north‐facing slope. The similarity between the seed banks and standing vegetation was higher than previously shown for renosterveld, and appears to be higher than for fynbos.     

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.