Bet‐hedging and germination in the Australian arid zone shrub Acacia ligulata

The diaspore of the Australian arid zone shrub Acacia ligulata is dispersed by birds and ants. To investigate the benefits of providing a dispersal structure attractive to both groups, we compared the germination response and viability of seeds eaten by birds, handled by ants or collected from trees to simulated precursors of germination: scarification, fire and rainfall were simulated. Seed germination and viability were related to the degree of preheating disturbance to the seed coat. Heating increased the germinability of seeds not scarified or eaten by birds. In the absence of heating, ingestion by birds increased germinability. Heating increased the mortality of seeds. Our results suggest that ingestion of seeds by birds may break seed dormancy and hence enable some seeds to germinate soon after dispersal. Alternatively, seeds not eaten by birds are likely to remain dormant until sufficiently scarified by soil or stimulated by fire. Consequently, in areas such as the Simpson Desert, A. ligulata may be able to use a range of seedling establishment ‘windows’ provided by monsoon rains, post‐fire environments and unseasonal winter rains, and also spread the risk of unsuccessful seedling establishment by retaining dormant seeds in the seedbank.     

Post-fire regeneration strategies of Californian coastal sage shrubs

Summary Regeneration methods for coastal sage srub vegetation after fire were studied in the coastal Santa Monica Mountains of southern California. Six sites were sampled two years after a large fire of fall, 1978. The intensity of fire varied. Foliar cover and flowering incidence were recorded for individuals regenerating by resprouting or from seed. Resprouting plants contributed most to post-fire recovery, comprising 95% of the relative foliar shrub cover; 84% of resprout and 47% of seedling cover had flowered. An ANOVA of reproductive mode and fire intensity indicates that resprout total cover and individual size are significantly greater than those of seedlings, regardless of fire intensity. Among sites the average foliar cover of resprouts exceeded that of seedlings by factors ranging from 9 to 63. All coastal sage species examined resprout, although the potential vigor of resprouting appears to vary widely within genera (e.g. Encelia, Eriogonum, and Salvia) and even within species. In the second growing season following fire seedling density increased due to seeds shed by resprouted shrubs. Most of the cover on these stands of coastal sage scrub is destined to be either crown-sprouted individuals or their progeny.     

Fire, soil fertility and delayed seed release: a community analysis of the degree of serotiny

Delayed seed release (serotiny) is a convergent plant trait in fire-prone regions of the world but explaining the degree of serotiny has remained elusive because of the paucity of community data. Selective forces involving seed predators, fire and soil nutrients have been suggested as factors influencing serotiny. We tested whether protection of seeds and/or synchronized dispersal were associated with different levels of serotiny and if resprouting ability influences selection for strong serotiny. We compared the numbers and abundance of 146 woody species with delayed dispersal among five community types varying in combinations of fire severity, fire frequency, soil fertility and seed predators. The strength of the relationship between levels of serotiny and environmental factors was tested among community types ranging from rainforests to heathlands. Highest levels of serotiny were recorded in low nutrient shrublands with intermediate fire return intervals that burn at high severity, while the lowest were recorded in high nutrient, low flammability forests. Both protection of seeds and synchronized seed release were related to fire effects in nutrient-limited environments. Strong serotiny is prominent in species killed by fire whereas weak serotiny is more common in resprouting species. Recruitment failure in the inter-fire interval appears to drive selection for strong maternal care of seeds and synchronized seed dispersal in fire-prone environments. Weak serotiny is proposed as a bet-hedging strategy that relies on resprouting after fire for population persistence and higher probability of inter-fire recruitment. The spectrum of serotiny (weak to strong) in these communities is proposed to be driven by the interactive effect of both fire and soil nutrients on the selection for delayed seed dispersal.     

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.

     

Patterns of reproductive effort with time since last fire in Florida scrub plants

Abstract. In periodically burned ecosystems, fire frequency may be an important selective pressure for the evolution of plant reproductive allocation patterns. We evaluated this hypothesis for Florida (USA) scrub plants by developing three models of reproductive effort with time since last fire given assumptions concerning seed dormancy and seedling establishment. We then examined reproductive effort of five woody, resprouting shrubs at sites representing nine times since last fire (ranging from 0–64 yr). All species showed significant patterns with time since fire in percentage of stems reproductive and fruit production. Stems of all species needed to attain a minimum size before flowering. Four species had the greatest level of reproductive effort (fruit biomass/above-ground biomass) within 5 yr post-fire and best fit the Early Peak Model of reproductive effort (i.e. between-fire seedling recruitment or seed dormancy). A fifth species best fit the Broad Peak Model (i.e. immediate post-fire seedling establishment), peaking in reproductive effort at 7 yr post-fire. Both of these models are based on somewhat variable fire-return intervals, suggesting that the frequency of scrub fires may have been too unpredictable to select for reproductive allocation patterns precisely reflecting particular fire-return intervals. Early peaks in post-fire reproductive effort may be a bet-hedging strategy to allow for greater chances of seedling establishment and survival.     

R.M. Cowling,D.M. Richardson & S.M. Pierce (eds.) 1977. Vegetation of Southern Africa.

We determined the role of bird dispersal in seed and seedling dynamics of the tree Kalopanax pictus from 7 years of observing seed rain and seedling emergence in a broad-leaved deciduous forest in central Japan. We also performed an experiment on the influence of seed pulp on germination of seeds of K. pictus. Seeds of this species can lie dormant for several years, and this causes rather constant yearly seedling emergence in spite of irregular seed production. The spatial distribution of the seedlings that emerged each year (maximum distance from nearest conspecific seed-bearing tree of 90 m) was wider than that of gravity-dispersed seeds (max. distance of 37 m), suggesting seed dispersal by birds in winter. Emerged seedling densities at sites over 20 m from the nearest conspecific seed-bearing tree were highest in the spring of 1991, about half a year after the largest seed fall of the observation period. However, emerged seedling densities within 20 m from seed-bearing trees were highest in 1992, 1.5 years after the largest seed fall. These field observations may be explained by the experimental results on the effects of seed pulp on germination. Intact seeds germinate slowly at low germination rates, while seeds without seed pulp germinate quickly at high germination rates. Fallen seeds with seed pulp thus appear to form a seed bank near seed sources (temporal dispersal), while seeds scattered by birds appear to increase the possibility of reaching the present safe sites in distant areas with quick germination (spatial dispersal).     

Seed and seedling ecology of piñon and juniper species in the pygmy woodlands of western North America

Knowledge of the seed and seedling ecology of the piñon and juniper woodlands of western North America is essential for understanding both the northward migration and expansion of the woodlands during the Holocene (< 11,500 B.P.), and the accelerated expansion of the woodlands since settlement of the West by Anglo-Americans around 200 years ago. We follow the fates of seeds and seedlings of the different piñon and juniper species within the woodlands from seed development to seedling establishment, and discuss the implications of this information for the past and present expansion of the woodlands. While seed development requires about two and one-half years in pinons, it is species-dependent in junipers and can take one, two, or even three years. Substantial seed losses can occur during seed development due to developmental constraints, and before or after seed maturation as a result of insects, pathogens, or predatory animals. In piñon pines, the primary seed dispersers are scatterhoarding birds (corvids) and rodents that harvest seeds from the trees or after seed fall and cache them in the soil. In contrast, most junipers appear to be dispersed primarily by frugivorous birds and mammals that ingest the seeds and defecate them onto the soil surface. We have recently documented that scatter-hoarding rodents also disperse juniper seeds. Disperser effectiveness, or the contribution a disperser makes to the future reproduction of a plant population, may vary among species of piñons and especially junipers. Piñon seeds are short-lived and exhibit little dormancy, and they probably only germinate the spring following dispersal. Juniper seeds are long-lived and seed dispersal can occur over one or more years. Seed germination can be delayed for several years due to impermeable seed coats, embryo dormancy, or the presence of inhibitors. Seedling establishment of piñon pines is facilitated by nurse plants but, while junipers often establish beneath nurse plants, they are capable of establishing in open environments. In the southwestern United States, higher establishment of juniper occurs in open environments due to more favorable precipitation, and competition may be more important than facilitation in determining establishment. When considering the mechanisms involved in the past and present expansion of the woodlands, short-distance dispersal, local population growth, and long-distance dispersal are all important. Different classes of dispersers, some of which appear to have coevolved with the tree species, appear to be responsible for local (short-distance) vs. long-distance dispersal in pinons and junipers. Because ecotones form the interface between the woodlands and adjacent communities, they can provide valuable information on both the seed dispersal and seedling establishment processes responsible for tree expansion. Disturbance regimes and, recently, the effects of humans on those regimes have major effects on the expansion and contraction of the woodlands. Before Anglo-American settlement, fires occurred as frequently as every 50–100 years throughout much of the woodlands. During this century, fire frequencies have been reduced due to the indirect effects of livestock grazing and the direct effects of removing Native Americans from the ecosystem and implementing active fire-prevention programs. The result has been an increase in tree-dominated successional stages at the expense of grass-dominated stages. Various management techniques, including controlled burning and chaining, have been implemented to reduce tree dominance, but their effects depend largely on the life histories of the tree species and the disturbance characteristics. Several areas relating to the seed and seedling ecology of the piñon and juniper require additional research if we are to truly understand the dynamics of the woodlands.     

Seed germination ecology in southwestern Western Australia

Germination responses of species from the native plant communities of southwestern Western Australia can be related to syndromes of life history, fire response, and seed storage, and also to factors related to environmental stress. The Mediterranean-type climate of the region with periodic drought and recurrent fires affects the production of viable seeds in plants of limited stature and rooting depth. Fire response ephemerals and species cued to flower by fire tend to produce viable, readily germinable seeds, but there are instances where seed production is aborted in these predominantly herbaceous life forms. Clonal, rhizomatous species often produce mainly inviable seeds. Production of viable seeds in woody species of these highly diverse communities may also be restricted by limitations to cross pollination. Obligate post-fire seeding species tend to produce a greater proportion of viable seeds than species which are capable of resprouting following fire. Serotinous species, whether post-fire re-seeders or post-fire resprouting species, produce mainly viable seeds, which germinate readily once freed from protective fruits. Species of the legume families and a few others of the soil seed bank produce innately dormant seeds which can be germinated following heat shock treatments which simulate the effects of fire. Heat shock in these species appears mainly as a mechanism to crack the hard seed coats, but the effect of heat to denature seed coat inhibitors has not been eliminated. Western Australian species do not seem to break dormancy when exposed to leachates from burned wood as has been observed in comparable habitats in California and South Africa, but further research is advised. Germination in many native southwestern Australian species is cued by temperatures that correspond to the winter rainfall period. There are also indications that an after-ripening period of warm, dry storage increases percentage of germinable seeds. Stimulation of germination by hormones is almost unresearched in Western Australia, but germination percentages have been increased in a small number of species of horticultural potential. Stimulation of germination by soil nutrient concentrations is almost unresearched in Western Australia, except for the inhibitory effect of excess sodium chloride levels inEucalyptus andMelaleuca. These species only germinate when osmotic effects are reduced to lower levels as would occur when winter rains dilute soil salts. Application of research on seed germination has already enhanced the establishment of seedlings in the restoration of mine sites and is becoming important in aspects of the breeding and selection of native plants for the cut flower, bedding plant and essential oil industries.     

Stem demography and post fire recruitment of Podocarpus drouynianus: a resprouting non-serotinous conifer

Podocarpus drouynianus is unusual in the Podocarpaceae and conifers in general in being a strongly resprouting species which thrives in a fire-prone environment. The species is a dioecious, multi-stemmed shrub endemic to the eucalypt forests of the south-west of Western Australia. Stems are killed by fire but it is able to regenerate its foliage from a lignotuber. The total bud bank seems to be released by burning and death of the shoots, while pruning without fire releases only a proportion of available buds. Growth rate of resprouts is initially rapid, with stems reaching 25% of their mature length in the first year after fire. However, growth of juvenile plants is very slow. Fire promotes cone production on the new stems about one year after fire. The large seed is recalcitrant but timing of germination coincides with the wettest time of the year in the area where the species grows. Pyrogenic coning avoids loss of seed due to severe fires and is a form of mast reproduction that benefits this species by increasing pollen transfer and producing a mass seed crop to attract the primary vertebrate disperser – the emu. The strong resprouting ability ensures the species is resilient to frequent disturbance and to severe predation of the seed crop. Vertebrate dispersal tends to concentrate seed lots, making them vulnerable to predation if they are deposited in open areas. However, being dispersed by a large vertebrate will ensure long-distance dispersal, often to areas beyond that which was subject to fire. These would include vegetated areas, where the large seeds could lodge in safe sites and produce seedlings that are able to survive under the shade of established plants. © 2005 The Linnean Society of London, Botanical Journal of the Linnean Society , 2005, 149 , 433–449.     

Fire season and intensity affect shrub recruitment in temperate sclerophyllous woodlands

The season in which a fire occurs may regulate plant seedling recruitment because of: (1) the interaction of season and intensity of fire and the temperature requirements for seed release, germination and growth; (2) post-fire rainfall and temperature patterns affecting germination; (3) the interaction of post-fire germination conditions and competition from surrounding vegetation; and (4) the interaction of post-fire germination conditions and seed predators and/or seedling herbivores. This study examined the effects of different fire intensities and fire seasons on the emergence and survival of shrubs representing a range of fire response syndromes from a summer rainfall cool climate region. Replicated experimental burns were conducted in two seasons (spring and autumn) in 2 consecutive years and fuel loads were increased to examine the effects of fire intensity (low intensity and moderate intensity). Post-fire watering treatments partitioned the effects of seasonal temperature from soil moisture. Higher intensity fires resulted in enhanced seedling emergence for hard-seeded species but rarely influenced survival. Spring fires enhanced seedling emergence across all functional groups. Reduced autumn recruitment was related to seasonal temperature inhibiting germination rather than a lack of soil moisture or competition. In Mediterranean-type climate regions, seedling emergence has been related to post-fire rainfall and exposure of seeds to seed predators. We think a similar model may operate in temperate summer rainfall regions where cold-induced dormancy over winter exposes seeds to predators for a longer time and subsequently results in recruitment failure. Our results support the theory that the effect of fire season is more predictable where there are strong seasonal patterns in climate. In this study seasonal temperature rather than rainfall appears to be more influential.     

Demography of a non-sprouting and resprouting Hakea species (Proteaceae) in fire-prone Eucalyptus woodlands of southeastern Australia in relation to stand age,drought and disease

Population size-structures, seed production, canopy seed storage (serotiny), and recruitment were investigated in relation to fire, drought and disease for a pair of co-occurring resprouting and non-sprouting shrub species from the genus Hakea (Proteaceae) in fire-prone Eucalyptus woodlands in western Victoria, Australia. The non-sprouter species, Hakea decurrens, showed faster height growth, higher seed production and higher seed viability than the resprouter, Hakea rostrata. Population size structures in stands up to 24 years since last fire showed no evidence of inter-fire recruitment for either species. Following a fire in 1990 in a mixed species stand 15–20 years old, the estimated number of viable seeds released from canopy-stored seed banks was approximately equal for both species. However, the rate of seedling establishment in the first year was about 10 times higher, and seedling suvivorship over the first 5 years was seven times higher, for the non-sprouter. Seedlings of Hakea decurrens and resprouts of Hakea rostrata began to produce seeds within three years of the last fire, while the few surviving seedlings of Hakea rostrata showed no evidence of reproductive maturity after six years. Inter-fire recruitment was recorded for the non-sprouter, Hakea decurrens, in the oldest stand (burned in 1967) between 24 and 28 years since last fire. This was associated with an increased rate of seed release from serotinous fruits due to the onset of high rates of adult plant mortality. High adult mortality and increased seed release correlated with increasing stand age, the occurrence of severe drought, and the likely presence of Phytophthora cinnamomi, a fungal pathogen which damages the root system, reducing water and nutrient uptake. There were no new recruits for the resprouter Hakea rostrata in this stand, but old plants continued to resprout from basal lignotubers and no mortality was observed. While recruitment of strongly serotinous shrub species is commonly described as being restricted to the immediate post-fire period, the present study illustrates that other events (e.g., senescence, drought, disease) can lead to recruitment of serotinous non-sprouters and may be important in the maintenance of populations during unusually long periods without fire.     

Soil temperatures after the passage of a fire: Do they influence the germination of buried seeds?

Abstract Soil temperatures down to a depth of 5 cm were measured in the days following one fire in summer, one fire in winter and in unburnt vegetation during summer. Soil temperatures did not rise above 40°C after the winter fire or in unburnt vegetation during summer. Consequently, no impact on seed dormancy in the soil seedbank was expected. After a summer fire, soil temperatures above 40°C were found up to 4.5 cm in depth, while temperatures above 60°C were found only in the top 0.5 cm of soil. These temperatures are sufficient to break seed dormancy in some legume species in the seedbank. Hence, the season of burn may influence the number of seeds in the soil that have their dormancy broken and subsequent germination levels.     

Ant-mediated expansion of an obligate seeder species during the first years after fire

Most obligate seeder species build up a soil seed bank that is associated with massive seed germination in the year immediately after a fire. These species are also shade‐intolerant and disappear when vegetation cover closes, creating unsuitable conditions for seedling recruitment. The only way for these plants to expand their populations is when habitats suitable for seedling recruitment arise (i.e. in years immediately after a fire). However, short primary seed dispersal of obligate seeders does not allow these plants to colonise the suitable habitats, and these habitats can only be colonised by secondary seed dispersion. We hypothesised that Fumana ericoides, an obligate‐seeding small shrub, not only establishes abundantly in the first year after fire, but also expands its local range in the following years due to secondary dispersal by ants while suitable habitats are still available. We tested this hypothesis using experimental studies and a simulation model of potential population expansion in a recently burned area. Results showed that F. ericoides not only established prolifically in the year immediately after fire, but was also able to recruit new individuals and expand its population in the years following the fire, despite a low germination rate and short primary seed dispersal. Ant‐mediated seed dispersal and availability of suitable habitats were key factors in this phenomenon: ants redistributed seeds in suitable habitats while they were available, which accelerated the expansion of F. ericoides because new plants established far away from the core population.     

How much seed remains in the soil after a fire?

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

Ecological benefits of myrmecochory for the endangered chaparral shrub Fremontodendron decumbens (Sterculiaceae)

Fremontodendron decumbens grows in a single county in central California, USA. Prior research showed that its elaiosome-bearing seeds are dispersed by the harvester ant Messor andrei. I tested several hypotheses regarding the positive role of ant-mediated dispersal to F. decumbens: (1) Does ant-mediated seed dispersal facilitate seed escape from rodent predation?; (2) Does ant processing of seeds stimulate germination?; (3) Are ant middens more suitable microsites for seed or seedling survival in unburned chaparral areas?; and (4) Do survival benefits of dispersal occur post-fire in the form of differences in seedling survival probabilities and, if so, why? Results of tests of each hypothesis were: (1) similar percentages of seeds placed on ant middens and under F. decumbens shrub canopies were destroyed by rodents, but seeds from which elaiosomes had been removed were more likely to escape rodent predation; (2) seeds processed by ants did not germinate more readily than seeds removed directly from shrub branches; (3) seedling predation was a major cause of mortality in unburned chaparral on both ant middens and under shrubs, and overall seedling survival did not differ between the two microsites; (4) post-burn seedling survival was significantly greater for seedlings dispersed away from F. decumbens shrub canopies, because dispersed seedlings were both less likely to be killed by predators and more likely to be growing in a gap created by the fire-caused death of an established shrub. I concluded that the major ecological benefit to F. decumbens of ant-mediated seed dispersal was elevated post-fire seedling survival resulting from enhanced escape by dispersed seedlings from both predation and competition.     

Variation in dormancy and germination in three co-occurring perennial forest herbs

Mesic deciduous forest herbs often disperse seed with morphophysiological dormancy (MPD) that prevents germination during unfavorable periods for seedling survival. However, for seeds of some species with MPD, seasonal separation of root and shoot emergence and variation in dormancy levels can complicate interpretation of seedling emergence timing in the field. We tested whether dormancy-break and germination requirements differed among co-occurring perennial forest herbs, Actaea racemosa, Hydrastis canadensis, and Sanguinaria canadensis, which are wild-harvested for their medicinal properties and known to have MPD. Seeds of all species exhibited a summer → autumn → winter requirement for seedling emergence in spring. However, species differed in seed-bank persistence due to variation in primary dormancy levels and stratification requirement of seeds. A. racemosa and H. canadensis can form short-term persistent seed bank, whereas S. canadensis can form a long-term persistent seed-bank, regardless of whether elaiosomes were removed from seeds prior to burial. A. racemosa seeds are dispersed in autumn with weak physiological dormancy, as seeds germinated to high rates at 15/6°C after 8 weeks. In contrast, most seeds of the summer dispersed species, H. canadensis and S. canadensis, require summer temperatures to overcome physiological dormancy. Consequently, seedling emergence is reduced and delayed by 1 year if seeds are not sown immediately following the period of natural dispersal. Seedling emergence was much lower in the field than in controlled conditions for all species, especially in the small-seeded A. racemosa. Interspecific variation in dormancy levels and germination traits must be considered when establishing populations for conservation purposes and in understanding recruitment limitation in perennial forest herbs.     

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.     

Evolution of recruitment features in plants: a comparative study of species in the Rosaceae

Theory predicts that life history traits that reduce the impact of environmental variation show patterns of negative covariation (trade‐offs). In plants, seed size, seed dormancy and dispersal (in space) interact to reduce risk in a temporally and spatially variable environment. Dispersal in time and space permits escape from unfavourable conditions, whereas a large seed size may improve establishment under unfavourable conditions. However, large seeds may suffer a high rate of loss due to seed predators. The objective of this study was to examine relationships between seed size, seedling establishment and seed dormancy by combining data from field experiments with information of phylogenetic relationships among 11 species in the subfamily Rosoideae (Rosaceae). The predicted relationship between seed size and seed predation was also examined. Emergence from seed, survival and recruitment of seedlings, effects of seed predators (post‐dispersal predation), and seed dormancy were studied experimentally in the field. All species generated seedlings in the experimental plots. Overall, the emergence of seedlings was promoted by disturbance. Several species possessed seeds that were able to germinate after being buried in the soil column for three years. Despite a small data set (ten phylogenetically independent contrasts), the comparative analysis confirmed an expected positive effect of seed size on establishment ability. The emergence of seedlings increased with seed size. The results did not demonstrate the hypothesised positive association between seed size and predation risk, or negative association between seed size and seed dormancy. However, the contrast analysis showed a significant negative relationship between seed dormancy and survivorship of seedlings, i.e. between quantitative data of dormancy and establishment capacity of the species obtained under natural conditions in the field. When not controlling for phylogeny (without contrasts), the only significant effect was a negative association between seed size and seed dormancy.     

植物天然更新过程中种子和幼苗死亡的影响因素

植物天然更新包括有种子搬运、种子库动态、种子萌发和幼苗定居等过程。从种子生产到幼苗定居的更新是植物生活史中最为敏感的阶段之一 ,多种因素的影响种子和幼苗的命运。其中包括 :( 1 )动物取食或病原体侵袭。种子在扩散和搬运过程中 ,易被小哺乳动物或无脊椎动物取食。蛀虫也可以使种子失去萌芽能力。病原体感染种子和幼苗 ,容易引起种子和幼苗的死亡。 ( 2 )异质生境的影响。在不同生境中 ,光照条件、土壤水分和化学成分等因子的组合严重影响种子和幼苗的命运。 ( 3 )干扰的影响。小尺度和大尺度的干扰都可以影响到植物更新时种子和幼苗的命运。林窗作为特殊的干扰体系 ,为不同种类植物提供了更新的机会。 ( 4 )繁殖体特征。种子大小、质量和保护色等特征影响种子和幼苗在更新过程中的生存。种子休眠期间 ,由于生理衰老和腐烂的原因使种子失去活力而不能萌发。 ( 5 )密度和距离制约。母株附近由于密度竞争的影响 ,种子和幼苗死亡率都较高。     

The potential for mycobiont sharing between shrubs and seedlings to facilitate tree establishment after wildfire at Alaska arctic treeline

Root‐associated fungi, particularly ectomycorrhizal fungi (EMF), are critical symbionts of all boreal tree species. Although climatically driven increases in wildfire frequency and extent have been hypothesized to increase vegetation transitions from tundra to boreal forest, fire reduces mycorrhizal inoculum. Therefore, changes in mycobiont inoculum may potentially limit tree‐seedling establishment beyond current treeline. We investigated whether ectomycorrhizal shrubs that resprout after fire support similar fungal taxa to those that associate with tree seedlings that establish naturally after fire. We then assessed whether mycobiont identity correlates with the biomass or nutrient status of these tree seedlings. The majority of fungal taxa observed on shrub and seedling root systems were EMF, with some dark septate endophytes and ericoid mycorrhizal taxa. Seedlings and adjacent shrubs associated with similar arrays of fungal taxa, and there were strong correlations between the structure of seedling and shrub fungal communities. These results show that resprouting postfire shrubs support fungal taxa compatible with tree seedlings that establish after wildfire. Shrub taxon, distance to the nearest shrub and fire severity influenced the similarity between seedling and shrub fungal communities. Fungal composition was correlated with both foliar C:N ratio and seedling biomass and was one of the strongest explanatory variables predicting seedling biomass. While correlative, these results suggest that mycobionts are important to nutrient acquisition and biomass accrual of naturally establishing tree seedlings at treeline and that mycobiont taxa shared by resprouting postfire vegetation may be a significant source of inoculum for tree‐seedling establishment beyond current treeline.