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.     

热激对大兴安岭三种松科树种种子萌发的影响

研究了热激对大兴安岭地区兴安落叶松、樟子松、红皮云杉种子萌发的影响,热激温度分别为80℃、100℃、120℃、150℃,每个温度下热激时间分别为1min、3min、5min、10min。研究结果表明:兴安落叶松种子在100℃热激10min,120℃热激5min,10min,150℃热激5min处理下萌发率较对照显著下降(P%0.05),除80℃热激1min、3min、5min、120℃热激1min、150℃热激1min外,其他处理萌发速率指数较对照均显著下降(P%0.05);樟子松种子在高于80℃的所有热激处理萌发率较对照均显著下降(P%0.05),甚至没有萌发,除80℃热激1min外,所有处理下萌发速率指数较对照均显著下降;红皮云杉种子萌发率在80℃热激5min和100℃热激1min处理下较对照显著提高(P%0.05),萌发速率指数在80℃热激1min、3min、5min,100℃热激1min处理下较对照显著提高(P%0.05)。兴安落叶松种子可以在短时间高强度(150℃)热激下维持萌发率,但不能承受长时间热激;樟子松种子对高于80℃的热激反应敏感,热激降低了樟子松种子的萌发能力;热激可以提高红皮云杉种子的萌发能力,短时间(1min、3min、5min)低强度(80℃)热激(包括100℃热激1min)提高红皮云杉种子萌发率和萌发速率指数,红皮云杉种子也可以承受高强度(150℃)瞬时(1min)热激。     

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

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

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.     

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.     

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.     

The effect of heat and smoke on the soil seed banks of heathlands on permanent freshwater swamps

In South Australia, Swamps of the Fleurieu Peninsula are critically endangered due to past vegetation clearance and changes in hydrology, but still contain a high diversity of threatened plant species. This vegetation community provides habitat for 82 threatened ground-stratum plant species, including the nationally critically endangered subshrub Hibbertia tenuis which is endemic to these swamps. With infrequent burning, native ferns and taller shrubs outcompete these species. We conducted glasshouse trials to determine the potential of fire to regenerate threatened and other swamp plant species. Soil samples from eight swamps were used in germination trials with half of each sample treated with heat plus smoke, and half left untreated. Samples were spread onto trays (n = 188), and seedling emergence was recorded for twelve months. Emergence was dominated by native species (2649 seedlings m⁻²) compared to exotics (675 seedlings m⁻²). In total, 48 native and 25 introduced species germinated, with 21% of germinated native species absent from the above-ground vegetation. The dominant native fern Gleichenia microphylla did not emerge from any soil samples, indicating that recovery likely lags behind that of ground-strata species, giving them time to establish and set seed before being outcompeted. Thirty-four rare or threatened species germinated, including five species absent from above-ground vegetation. Of all native species that emerged, only five were confined to heat plus smoke treatments, suggesting most species will regenerate without fire if overstorey competition is reduced. However, seedling emergence of native shrubs/subshrubs more than doubled with the fire treatment. Of particular importance, H. tenuis showed an 18-fold increase in germination when treated with heat plus smoke. This study supports the utility of ecological burns as a management tool to regenerate threatened plant species in long-undisturbed peaty heathlands on permanent swamps.     

植冠与土壤种子库储存种子的萌发特性及策略

为明确黄土丘陵沟壑区植物种子库如何调控种子萌发来提高个体适合度,选择研究区7种具有种子库的主要物种为研究对象,以刚成熟和室内储存种子为对照,比较植冠宿存(5个宿存期)和土壤埋藏(5a埋藏期)对植物种子萌发特性的影响,探讨植冠种子库与土壤种子库储存下的种子萌发策略。结果表明:7种植物种子经过不同种子库储存后萌发特性表现出明显的种间差异,黄刺玫(Rosa xanthina)和水栒子(Cotoneaster multiflorus)种子萌发力表现为植冠宿存不变型、土壤储存增强型,土壤储存明显提高水栒子种子萌发速率;达乌里胡枝子(Lespedeza davurica)和狼牙刺(Sophora davidii)种子萌发力表现为植冠宿存增强型、土壤储存减弱型,种子萌发历时表现为植冠宿存延长型,土壤种子库储存还可加快达乌里胡枝子萌发速率、缩短萌发历时;茭蒿(Artemisia giralaii)和铁杆蒿(Artemisia gmelinii),种子萌发率随植冠宿存时间先升高后降低,随土壤储存时间先降低后升高,土壤储存可推迟其萌发,铁杆蒿种子萌发速率在植冠与土壤储存后均加快;紫丁香(Syringa oblata)种子萌发率随植冠宿存先升高后降低,土壤储存明显加快其种子启动萌发与速率。在黄土丘陵沟壑区,植物种子经过植冠或土壤种子库储存,或增加、加快、提早萌发充分利用有利条件提高占据性,或减少、减缓、推迟萌发分摊不利条件的风险;而且该区植物植冠与土壤储存后种子萌发特性间的关系,体现各自适应环境与应对干扰的分工与协作策略,主要表现为:单一主导型和相辅相成型。     

热激和烟熏对大兴安岭三种灌木种子萌发的影响

研究了大兴安岭地区珍珠梅(Sorbaria sorbifolia)、绣线菊(Spiraea salicifolia)、金露梅(Potentilla fruticosa)种子对热激和烟熏的萌发响应。热激分为干热激和湿热激。干热激处理水平分别为:60℃5 min、60℃15 min、90℃5 min、110℃5 min、150℃5 min;湿热激处理水平分别为:60℃5 min、60℃15 min、95℃5 min;烟熏处理水平分别为:5 min、10 min、20 min。结果表明:(1)在对干热激的响应上,珍珠梅种子的萌发率在60℃干热激处理15 min后显著提高(P<0.05),在110℃、150℃干热激处理5 min后显著下降(P<0.05);绣线菊种子的萌发率在干热激处理后显著提高(P<0.05),平均萌发时间显著缩短(P<0.05);金露梅种子的萌发率在150℃干热激处理5 min后显著下降(P<0.05);(2)在对湿热激的响应上,珍珠梅和绣线菊种子在湿热激处理后停止萌发;金露梅种子在60℃湿热激处理15 min后停止萌发,在95℃湿热激处理5 ...     

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

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

Broad‐scale patterns in smoke‐responsive germination from the south‐eastern Australian flora

Questions

Fire is a crucial component of many ecosystems. Plants whose seeds germinate in response to smoke may benefit from resource availability in the post‐fire environment. Smoke can influence germination timing and success, as well as seedling vigour, resulting in burgeoning research interest in smoke‐responsive germination. Research in this field has largely focused on four key ‘Mediterranean‐type’ fire‐prone ecosystems: the Mediterranean Basin, South African fynbos, Californian chaparral and Western Australia. There are far fewer studies from south‐eastern Australia, a fire‐prone but not “Mediterranean‐type” region. How does smoke‐responsive germination in this region vary according to ecological, phylogenetic, and methodological variables?

Location

South‐eastern Australia.

Methods

We investigated patterns of smoke‐promoted germination in south‐eastern Australian plants across habitat types, growth forms, fire response strategies, phylogeny, taxonomic levels and smoke application methods. We compiled and interrogated data comprising 303 entries on germination responses to smoke in 233 south‐eastern Australian plant species, from 33 different sources.

Results

Smoke‐responsive germination occurs at a lower rate (~41% of tested species) in south‐eastern Australian flora than it does in fynbos and Western Australian floras, and there is clear patterning within these data. Obligate‐seeding species were more likely to respond, Leguminosae and Rubiaceae were less likely to respond (although we question the generality of these results), while Poaceae were more likely to respond to smoke. Finally, studies using aerosol smoke and studies conducted in situ were most likely to find smoke‐promoted germination.

Conclusions

Obligate seeders and Poaceae may be selected for in habitats with higher fire frequencies, consistent with literature suggesting that short inter‐fire intervals favour grasslands over forests. These findings may be particular to south‐eastern Australia, or more widely applicable; more broad‐scale comparative research will reveal the answer. By synthesizing the south‐eastern Australian smoke germination literature we broaden our understanding beyond the better‐studied Mediterranean‐type floras.

     

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.     

Effects of smoke, heat, darkness and cold stratification on seed germination of 40 species in a cool temperate zone in northern Japan

The effects of smoke, heat, darkness and cold stratification on seed germination were examined for 40 species with various life history attributes. These species establish in early successional stages on a volcano and are distributed in cool temperate zones of northern Japan. Smoke decreased seed germination in 11 species and increased it in one species, Leucothoe grayana . Germination of Polygonum longisetum was enhanced by a combination of smoke and cold, and that of Aralia elata by smoke and heat. Heat increased germination for three species and decreased it for one. Cold stratification broke dormancy in seeds of 11 species. Continuous darkness decreased germination of 22 species and did not increase germination for any species, showing that approximately half of the species require light for maximum germination. Although most species are sun plants that establish in early stages of succession and/or in disturbed areas, smoke and heat do not enhance germination of these species after disturbance, even when the disturbance is fire. Germination of slender and/or large seeds tends to be decreased more by smoke, probably because of their larger surface area. Light is more important than smoke and heat for detection of disturbance and for seed germination in this region. However, despite the low fire frequency in the region, germination of a few species was increased by fire-derived stimuli.     

生物土壤结皮对荒漠土壤种子库和种子萌发的影响

研究了腾格里沙漠东南缘在不同自然条件（风、温度、水分）下,人工固沙植被区（24龄、41龄、50龄）和相邻天然植被区的两种生物土壤结皮对荒漠土壤种子库和种子萌发的影响。结果表明,荒漠土壤种子库在苔藓结皮上的储量显著高于藻类结皮。随着生物土壤结皮的发育,种子萌发量在苔藓结皮上增加,在藻类结皮上减少。生物土壤结皮层的含水量对种子萌发有显著的影响（p〈0.05）,植物种子在湿润处理的生物土壤结皮上的萌发量高于干燥处理的生物土壤结皮上的种子萌发量。生物土壤结皮表层温度和亚表层温度对荒漠植物种子萌发无显著影响（p〉0.05）,但总体而言,对于苔藓结皮,植物种子在较高温度下的萌发量略高于在较低温度下的萌发量,而对于藻类结皮,植物种子在较低温度下的萌发量略高。     

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.     

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.     

Comparative seed germination ecology of Austrostipa compressa and Ehrharta calycina (Poaceae) in a Western Australian Banksia woodland

Austrostipa compressa, a native ephemeral of southwest Western Australia was stimulated to germinate under a range of temperatures, in the presence of light, and exposure to smoke-water. This combination of environmental cues results in winter-maximum germination in immediate postfire and disturbed-soil environments of this Mediterranean-type climate. In contrast, Ehrharta calycina, an introduced perennial grass from southern Africa that has invaded Banksia woodlands, germinated under a wide range of temperature and light conditions, but showed no promotive response to smoke-water. Although A. compressa seeds tolerated heat shock better than E. calycina, the self-burial mechanism of A. compressa seeds ensures protection from fire. High-intensity fire could have a greater impact on E. calycina, as the seeds of this species tend to accumulate in the top of the soil profile where they are more susceptible to high temperatures. Although seeds of E. calycina are more susceptible to high temperatures, survival of mature individuals by postfire resprouting ensures continued survival in native woodlands. Estimates of soil seed bank densities showed extreme variability, but some recently burnt areas of the Yule Brook Botany Reserve contained up to 8000 seed m^?2 of A. compressa and nearly 75 000 seeds m^?2 of E. calycina. Viable soil seed bank densities of A. compressa are reduced with time-since-last fire, but areas of greater than 45 years since the last fire, still contained up to 119 seeds m^?2. In both species, only about half their soil seed bank germinates following fire, thus ensuring the potential for later recruitment. Massive soil seed populations of E. calycina in native Banksia woodlands pose a major problem to management of this plant community type.     

土壤种子库的分类系统和种子在土壤中的持久性

对国际上已经发表的10个土壤种子库分类系统的内容进行了总结和阐述,并对土壤种子库分类系统进行了评述,其中Thompson ＆ Grime在1979年提出的把土壤种子库分为短暂土壤种子库（Transient soil seed bank）和持久土壤种子库（Persistent seed bank）的二元分类系统以及Thompson等人提出的把土壤种子库分为（1）短暂土壤种子库,（2）短期持久土壤种子库（Short term persistent seed bank）,（3）长期土壤种子库（Long termp ersistent seed bank）的三元分类系统在生态学文献中已被广泛采用。在此分类的基础上产生了植物种子在土壤中的持久性（Persistence）概念,持久性是指植物的一种特性,是指植物的种子在土壤中能够存活超过1a的特性;植物种子的持久性被认为是一种对环境的进化适应,它可以在多个生长季节萌发从而分担环境震荡的风险,持久土壤种子库不仅在不稳定的环境里占有优势;即使在稳定的环境里,也被认为能够减少种内和种间的竞争;造成持久性的原因可分为环境因子和种子本身的特性比如休眠等两个方面,持久土壤种子库的出现使得土壤种子库的研究与进化生物学结合起来,使得土壤种子库的研究进入一个新的领域,更易激发人们的兴趣。关于种子的大小、形状及持久性的关系问题已经引起了相当的争论,基本上有4种格局：一是种子大小和形状与种子在土壤中的持久性有关,小而圆或扁的种子在土壤易存活持久;二是种子大小与种子在土壤中的持久性有关,小种子在土壤中易存活持久,但种子形状与持久性无关;三是种子大小、形状与种子在土壤中的持久性无关;四为较大的种子在土壤易存活持久,而种子形状与种子在土壤中的持久性无关。影响种子在土壤中的持久性因子比较复杂,总结过去的文献发现主要有以下几个因子：①种子的散布方式,②捕食,③植被的物种组成,④风,⑤土壤基质,⑥火,⑦干扰等。通过比较分析和研究,提出影响种子大小和在土壤中的持久性关系格局的关键因子是气候,特别是生态系统所在地的雨量;湿润气候下容易产生前两种格局,而干旱环境下的生境容易产生后两种格局。