The effects of date of planting on field establishment of serotinous cape Proteaceae

Season of fire have marked effects on the germination and establishment of serotinous shrubs of the family Proteaceae in fynbos vegetation. To investigate reasons for this, we simulated the effects of different fire seasons by planting seeds into cleared fynbos and then followed their progress. Four species of Proteaceae were planted monthly at four sites over two and a half years. Exclosures were used to exclude rodent seed predators. Germination was confined largely to the three winter months (June–Aug.). Seeds planted from January–June had higher germination than those planted in the second half of the year. Higher levels of regeneration noted after fires in the first half of the year, were previously hypothesised to be results of predation. However, we obtained similar results despite the exclusion of seed predators. Monthly minimum temperature was strongly correlated with germination percentage but monthly rainfall was not. Loss of seed viability may be important, in determining post-fire seedling densities. Differential seedling mortality of earlier and late germinants appears to be unimportant in determining establishment levels. Our results nevertheless support the current practice of restricting management fires in fynbos to the summer-autumn period.     

The promotive effect of smoke derived from burnt native vegetation on seed germination of Western Australian plants

Exposure of dormant seed to cold smoke derived from burnt native vegetation had a positive influence on germination in one or more seed provenances in 45 out of 94 species of native Western Australian plants that are normally hard to germinate. When tested under controlled conditions some species showed earlier germination in smoke treatments than controls; in others smoke-treated seeds continued to germinate for several weeks after controls had achieved full germination. In the remainder, treated and control seeds germinated to similar time schedules. A group of 23 species which responded positively had previously been recorded as extremely difficult or impossible to germinate using conventional techniques. These included members of the genera Geleznowia (Rutaceae), Hibbertia (Dilleniaceae), Stirlingia (Proteaceae), Verticordia (Myrtaceae), Actinostrobus (Cupressaceae) and Pimelea (Thymelaeaceae). Both large- and small-seeded species were encountered amongst the positively responding taxa, which encompassed representatives of 15 families and 26 genera of dicotyledons, 5 families and 8 genera of monocotyledons and the gymnosperm Actinostrobus acuminatus. Sowing seeds on smoke-fumigated filter papers or watering with aqueous eluates of smoke elicited similar degrees of stimulation of germination, as did exposure to gaseous smoke in a readily germinating species Anigozanthos manglesii (Haemodoraceae) and the normally intractable species Lysinema ciliatum (Epacridaceae). Exposing recently burnt and unburnt natural bushland sites to smoke, smoked water or smoked dry sand elicited a significant germination response in 15 species. Over one third of the species sampled in the burnt site exhibited germination additional to that caused by the fire. Data are discussed in relation to previous germination studies on Australian and other taxa.     

Patterns of seed persistence in South African fynbos

In fire-prone communities such as fynbos, many species rely on regeneration from seed banks in the soil. Persistent seed banks are particularly important for species with life spans shorter than the average fire cycle, in order to counter local extinction. Persistent seed banks also give potential for restoring ecosystems following disturbances such as alien plant invasion. This study investigated the seed persistence patterns of 25 perennial species, representing several growth forms and life histories, during a three-year burial. Long-term persistence (i.e., seed bank half-life exceeding two years) was found in the hard-seeded Fabaceae and Pelargonium, and the nut-fruited Proteaceae. In this group, germinability was low and dormancy increased further following burial, resulting in a highly viable, dormant seed bank after three-year's burial. A second group with potentially long-term persistent seeds includes four taxa (Pseudopentameris, Passerina, Elegia and Restio) that either have low germinability or develop secondary dormancy following burial. Dormancy in the latter group was partially countered by exposure to smoke-seed primer. Of the small-seeded species, only two Erica species with high initial dormancy had long-term persistent seed banks. The other species mostly displayed high initial germinability and short-term persistent seed banks (i.e., seed bank half-life less than two years). This group included taxa with short to medium life-spans (Syncarpha, Roella) that were expected to have long-term persistent seeds in order to buffer against local extinction following average to long fire-return intervals. We hypothesize that light may play a role in overcoming secondary dormancy in those species, and could have resulted in an underestimate for seed persistence in this study. Alternatively, those short to medium life-span species persist via inter-fire recruitment in gaps or long-distance dispersal (of the smallest seed). No correlations were found between seed persistence and seed mass or variance in seed dimensions. Nor was a correlation found between seed persistence and phenol concentration. In fynbos, seed burial of larger seeds by ants and rodents are major processes that operate in conjunction with passive burial of small seeds. Selection for persistence can be expected to operate across all seed sizes and shapes in fire-prone communities.     

Population structure and reproductive status of two Banksia shrubs at various times after fire

Populations of the obligate-seeder, Banksia ericifolia, were even-aged. Seedling recruitment occurred only after fire. Mean genet size (height + canopy diameter; H+D) increased progressively with elapsed time since fire in stands last burnt 2–23 years before 1981. Populations of a co-occurring resprouter, B. oblongifolia, were mixed-aged. Genet size varied significantly between stands, but this variation was not explained by regressions of H+D on years since fire. In addition B. oblongifolia seedlings were recruited both after fire and in patches of heath unburnt for 16 years.Most flower and seed production in B. oblongifolia occurred in the stands last burnt less than 10 years previously. More than 30% of genets had not produced cones since the last fire, irrespective of how many years had elapsed. In contrast, few B. ericifolia genets had produced cones five years after fire, but by 16 years after fire nearly 100% had. Overall, about 51% of B. ericifolia inflorescences and about 28% of B. oblongifolia inflorescences set seed. The number of seeds in seed-bearing cones was not significantly different between species.Resprouting B. oblongifolia genets began flowering sooner after fire, but B. ericifolia subsequently overtook them in accumulating a bank of serotinous seeds. In the stand unburnt for 23 years the largest B. ericifolia genets had more than twice as many cones as the largest co-occurring B. oblongifolia. However, when accumulated cone production was compared for genets of equal H+D over all stands, there was no difference between species.We thank the New South Wales National Parks & Wildlife Service for permission to do this work in Ku-ring-gai Chase National Park. We are grateful to Don Adamson, Lynn Day, David Haig and James Sim for constructive comments on earlier drafts.     

Establishment of microscale vegetation pattern in maritime chaparral after fire

We quantified microscale pattern in vegetation and seed assemblages along a 24 m transect before and for two years following a controlled burn in chamise (Adenostoma fasciculatum) chaparral in central coastal California. Our objective was to document scale-dependent correlation between pre-burn seed assemblages, pre-burn canopy cover, microtopography, soil temperatures during burning, post-burn seed assemblages and post-burn vegetation. Scale-dependent correlations among pre- and post-burn seed densities, maximum soil temperatures during burning, microtopography and post-burn vegetation were measured based on two-term local covariance analysis.Seed distribution varied among species prior to fire, with seeds of some annual species concentrated in gap areas. Maximum soil temperatures during burning ranged from less than 50 °C to 225 °C, and were generally lowest in gaps in the pre-burn canopy. These gaps were associated with local topographic depressions. After burning, readily germinable seeds were concentrated in or near gaps in the pre-burn canopy. Germination of different species was variously enhanced, diminished or unchanged by the passage of fire. Post-burn vegetation was very patchy, with some areas nearly devoid of seedlings and other areas, especially pre-burn canopy gaps, supporting numerous seedlings. Seedling recruitment patterns in the second year were generally highly correlated with patterns in pre-burn seed banks and first year vegetation. Although many species exhibited similar recruitment patterns, several different mechanisms may have been responsible for the origin of those patterns.     

Impact of fire on Picea mariana and Pinus banksiana seedlings in subarctic lichen woodlands

Abstract. The demography of Picea mariana (black spruce) and Pinus banksiana (jack pine) seedlings was monitored through five censuses over 13 months in four different seed bed types after fire of four severity levels in lichen woodland. Most seeds germinated just before early frost in late summer 1990 or immediately after snow thaw in early spring 1991; the germination rate subsequently decreased. For both species, germination rate decreased along a gradient of fire severity. The proportion of Pinus seeds that produced a seedling surviving 13 months after sowing was 4.3 % and 0.4 % respectively in the intact lichen mat and in the mineral soil seed bed type. For Picea these values are 3.2 % and 0.2 % respectively. The low germination rate in a severely burned seed bed type appeared to be associated with the formation of a water-repellent crust at the soil surface following the fire. Seedlings were contagiously distributed and were more frequent in flat and hollow microsites, where there is probably more water available than on bumps or among pebbles. Properties of experimentally burned seed bed types may differ from those under natural fires where regeneration by seed generally occurs following dispersal. However, the high germination rate observed in the intact lichen mat suggests that scattered lichen woodland patches may respond to increased seed input by a higher frequency of seedling establishment.     

The invasion of Pteridium aquilinum and the impoverishment of the seed bank in fire prone areas of Brazilian Atlantic Forest

Several studies have documented that fires are widespread in the tropics. Because fire in the Atlantic Rain Forest is rare, fire events and their consequences at the community level have not been evaluated. This study describes the composition of different seed banks in areas of Atlantic Forest. The study was carried out in the National Park of Tijuca in the state of Rio de Janeiro. Four different areas were chosen considering their differences on fire history. Fifteen soil surface samples (23 cm (centimeter) diameter ×7 cm height at 5 cm deep) were obtained from random locations within each site. The dominant family at all study sites except in the most preserved one were Dennstaetiaceae, followed by Melastomataceae and Poaceae. The family Dennstaetiaceae was exclusively represented by Pteridium aquilinum (Klf.) Herter. There were no significant differences among areas when comparing the number of seedlings excluding P. aquilinum. However, the number of P. aquilinum was higher in the most disturbed area, while Melastomataceae and Rubiaceae were more frequent in a less impacted area. Our results suggest that the seed bank in disturbed areas of the Atlantic Forest possibly will not contribute for forest restoration after disturbance.     

Fire increases aboveground biomass, seed production and recruitment success of Molinia caerulea in dry heathland

During the last decades, the perennial tussock grass Molinia caerulea has shown an increased abundance in European heathlands, most likely as a result of increased nitrogen deposition and altered management schemes. Because of its deciduous nature, Molinia produces large amounts of litter each year, which may affect the intensity and frequency of accidental fires in heathlands. These fires may influence plant population dynamics and heathland community organization through their effects on plant vital attributes and competitive interactions. In this study, fire-induced changes in competitive ability and invasiveness of Molinia through changes in biomass production, seed set and seed germination under both natural and laboratory conditions were investigated. We found that fire significantly increased aboveground biomass, seed set and germination of Molinia. Seed set was twice as high in burned compared to unburned heathland. Two years after fire, seedling densities in natural conditions were on average six times higher in burned than in unburned heathland, which resulted in increased abundance of Molinia after burning. The seed germination experiment indicated that seeds harvested from plants in burned heathland showed higher germination rates than those from unburned heathland. Hence, our results clearly demonstrate increased invasive spread of Molinia after large and intense fires. Active management guidelines are required to prevent further encroachment of Molinia and to lower the probability of large fires altering the heathland community in the future.     

Ecological correlations between the persistence of the soil seed bank and several plant traits,including seed dormancy

It was empirically showed that seed size and life history correlate with the formation of a soil seed bank. Although no empirical data are available that indicate a close relationship between seed dormancy and the soil seed bank, dormancy has been considered essential to the formation of a soil seed bank. I have considered the formation of the soil seed bank and survival of seeds for more than a year in the soil, and the persistence and survival of the seed bank for more than 5 years. These periods were derived from the definition of a persistent seed bank and the criterion for seed banks of long-term persistence. Plant traits that are closely related to the formation or persistence of a seed bank and their relationships to dormancy were analysed using two pre-existing databases of seed longevity in soil and comparative ecology. The integrated database comprised 18 plant traits and seed bank formation or persistence data. This approach was used to identify more reliable general empirical rules. The results of a regression tree analysis and common statistical tests of plant traits indicated that only life history and seed size were closely related to seed bank formation, and dormancy was not essential for the formation and persistence of a seed bank. However, the contribution of dormancy differed slightly between dormancy types. Scarification or dry storage requirements to break dormancy slightly enhanced the formation and persistence of a seed bank, whereas a chilling requirement decreased the formation and persistence of a seed bank. In contrast, fluctuating temperature requirements clearly contributed to the formation and persistence of a seed bank.     

A comparative analysis of seed and cone characteristics and seed-dispersal strategies of three pines in the subsection Sabinianae

The seed-dispersal systems of Coulter pine (Pinus coulteri), gray pine (P. sabiniana), and Torrey pine (P. torreyana), all of the subsection Sabinianae, are not well understood. These pines occur in arid and semi-arid foothills and mountains of California that are subjected to frequent fires. Cone and seed traits of these three California pines are compared to those of four species of pines (sugar pine, P. lambertiana; Jeffrey pine, P. jeffreyi; ponderosa pine, P. ponderosa; and lodgepole pine, P. contorta) that occur in more mesic environments in the nearby Sierra Nevada mountains. The cones of the Sabinianae pines are large with thick, dense scales, and the scales of gray and Coulter pines are armed with sharp, recurved spines. The seeds of all three species are large, and those of gray and Torrey pines are nearly wingless. In contrast, the Sierra Nevada pines have small to medium-sized seeds with large wings that are initially dispersed by the wind. Heavy wing loading of the Sabinianae pine seeds causes them to fall rapidly, and they are not dispersed far by wind. However, animals remove the fallen seeds rapidly, and rodents and jays scatter hoarded many seeds in the soil. This caching activity results in seedling establishment. The unusual morphology of the cones and seeds of the Sabinianae pines is interpreted as a combination of traits that attract animal dispersers, thwart the foraging activities of seed predators, and promote the survival of seeds in an environment subject to frequent fires.     

Effects of altitude and seed size on germination and seedling survival of heathland plants in north Spain

The effects of altitude and seed size on germination and seedling survival were studied in Calluna vulgaris, Erica cinerea and Erica vagans. Experiments were carried out in the laboratory over a one year period. Seeds collected from heathlands of different altitudes were divided in two size classes. They were sowed on moist filter paper inside Petri dishes which were placed in chambers at 20 °C and a photoperiod of 12 h light/12 h darkeness. The seeds of Calluna vulgaris were the first to begin germination and had the highest rate and percentage of germination. The germination of Erica vagans was moderate, while seeds of Erica cinerea germinated later and the germination was very low. Seeds of Calluna vulgaris and E. cinerea collected at the highest altitudes had the highest germination percentages. Seed size in Calluna vulgaris and E. cinerea did not affect germination. However, large seeds of Erica vagans had higher germination rates and percentages than small seeds. The large seed size of Calluna vulgaris contributes to a better survival and growth of its seedlings.     

Effects of snow accumulation on survival of beech (Fagus crenata) seed

The escape of beech seeds from seed predators and winter desiccation due to snow accumulation was studied by comparing two sites in Japan: one site that experiences much snow and another site that experiences less snow cover. At the site with greater snow cover, about 70% of the beech seeds escaped seed predation by rodents during winter and about 70% of surviving seeds germinated successfully in spring. At the site with less snow cover, however, all of the beech seeds were eaten by rodents, and all seeds that were protected from feeding were killed by winter desiccation. We confirmed that snow prevents beech seeds from predation by rodents because it conceals their sight and scent. These effects are one of the main reasons why beeches in snowy areas regenerate constantly and those in less snowy areas do not.     

Smoke-induced germination of succulents (Mesembryanthemaceae) from fire-prone and fire-free habitats in South Africa

Previous studies have shown germination to be stimulated by fire-related cues such as heat and charate extract and, more recently, plant-derived smoke extract. However, smoke extract has not been tested on non-fire-prone species. We hypothesized that smoke-induced germination would be restricted to fire-prone species and investigated responses of members of the family Mesembryanthemaceae which has genera confined to fire-prone and to non-fire-prone habitats as well as genera with species across both habitats. Results of germination trials of smoke effects did not support the hypothesis. Plant-derived smoke extract stimulated germination in both fire-prone and non-fire-prone species. These results cast some doubt on the ecological significance of smoke as a fire-related cue and we suggest that investigation of effects of fire-related cues on non-fire prone species may throw light on general mechanisms of germination.     

Convergent seed germination in South African fynbos and Californian chaparral

California chaparral and South African fynbos are fire-prone communities dominated by species exhibiting remarkable similarities in germination response. In both regions there are a substantial number of species with germination stimulated chemically by charred wood and smoke. This type of germination behaviour has arisen independently in distantly related families and is interpreted as convergent evolution. Heat-shock is also an important germination trigger that is widespread, although in both regions it is most common in the same families. Phylogeney may play an important role in the presence of this postfire germination cue in both regions, but a much more rigorous analysis is required to show that this trait represents a single unique event in each lineage. In both regions, germination response is not randomly distributed across growth forms and there are marked regional similiarities in the type of germination behaviour associated with certain growth forms. Geophytes largely lack refractory seeds, which require fire-type cues for germination, but the presence of fire-stimulated flowering of bulbs and corms may time recruitment to subsequent postfire years. Annuals that cue germination to postfire conditions are predominantly triggered by chemicals from smoke and/or charred wood.     

Effects of fire on forest dynamics in southern Switzerland

Abstract. We analysed the influence of fire on species composition, floristic diversity and indicator values of the vegetation in forests of the Canton Ticino (South Switzerland). Both frequency of fire and time since last fire were taken into account using variance analysis. Results show a strong interaction of these two parameters, i.e. the reaction of the vegetation after a fire depends on the frequency of past fires. A rapid recovery is generally observed after fire, involving both herbaceous and woody species. However, a long-term tendency towards floristic and edaphic impoverishment is observed with increasing frequency of fires. This phenomenon appears to be linked to the progressive dominance of a few fire-tolerant species.     

Biodiversity and resilience of arthropod communities after fire disturbance in temperate forests

Changes in ecosystem functions following disturbances are of central concern in ecology and a challenge for ecologists is to understand the factors that affect the resilience of community structures and ecosystem functions. In many forest ecosystems, one such important natural disturbance is fire. The aim of this study was to understand the variation of resilience in six functional groups of invertebrates in response to different fire frequencies in southern Switzerland. We measured resilience by analysing arthropod species composition, abundance and diversity in plots where the elapsed time after single or repeated fires, as determined by dendrochronology, varied. We compared data from these plots with data from plots that had not burned recently and defined high resilience as the rapid recovery of the species composition to that prior to fire. Pooling all functional groups showed that they were more resilient to single fires than to repeated events, recovering 6–14 years after a single fire, but only 17–24 years after the last of several fires. Flying zoophagous and phytophagous arthropods were the most resilient groups. Pollinophagous and epigaeic zoophagous species showed intermediate resilience, while ground-litter saprophagous and saproxylophagous arthropods clearly displayed the lowest resilience to fire. Their species composition 17–24 years post-burn still differed markedly from that of the unburned control plots. Depending on the fire history of a forest plot, we found significant differences in the dominance hierarchy among invertebrate species. Any attempt to imitate natural disturbances, such as fire, through forest management must take into account the recovery times of biodiversity, including functional group composition, to ensure the conservation of multiple taxa and ecosystem functions in a sustainable manner.Electronic Supplementary Material Supplementary material is available to authorised users in the online version of this article at .     

Effects of rabbit gut passage on seed retrieval and germination of three shrub species

It has been known for a few decades that European rabbits consume seeds and fleshy fruits of native woody plants, but relevant factors in the endozoochory processes such as seed predation (chewing and digestion), sexual differentiation, or the temporal pattern of seed recovery have been little evaluated until now. In this study, we examined seed dispersal of three Mediterranean shrub species by wild rabbits through monitoring of seed retrieval and germination after gut passage. Twelve adult wild rabbits (Oryctolagus cuniculus; six males and six females) of similar size and age were fed seeds of three shrub species with fleshy fruits (Crataegus monogyna, Myrtus communis and Pistacia lentiscus). After ingestion of fruits, seeds were retrieved from dung every 12 h for a day and a half. The viability and germination of retrieved seeds were tested along with that of uneaten seeds. Between 5% and 76% of ingested seeds were retrieved from dung, with significant differences between species and sex. Most M. communis seeds were retrieved with 12–24 h after ingestion; almost all C. monogyna seeds were recovered with 0–12 h after ingestion; no seeds of P. lentiscus were recovered. Only in the case of M. communis seed was the recovery rate greater in female than in male. Passage through the rabbit gut significantly increased seed germination in M. communis, and decreased it in C. monogyna. In conclusion, wild rabbits acted in this study as predators of C. monogyna and P. lentiscus seeds and potential dispersers of M. communis seeds.     

Seed yield,seed size and germination behaviour in the annualPogogyne abramsii

Summary Models of the evolution of seed dormancy reveal that dormancy is favoured either when opportunities for establishment vary over time and when there is wide variation in the probability of success, or when the probability of success is limited by frequency dependence. Empirical evidence supporting the temporal heterogeneity hypothesis exists, but there is scant evidence for dormancy being favoured by frequency dependent competition among seedlings. We test the hypothesis that the intensity of between-sib competition should favour a positive relationship between maternal fecundity and seed dormancy. This hypothesis is supported for the rare, vernal pool annual,Pogogyne abramsii: the proportion of dormant offspring was significantly higher among high fecundity mothers than among other mothers. Dormancy inP. abramsii is controlled by the seed coat, a maternal tissue, so delaying germination favours the inclusive fitness of mothers by reducing the potential for competition among siblings. Seed weight and time to first germination varied significantly amongP. abramsii plants and mean seed weight increased linearly with plant biomass. Seed weight and seed number are independently regulated by plant size. Overall, seed weight varied 10-fold and variability in seed weight within mothers was not explained by plant biomass, seed yield or mean seed weight. GerminableP. abramsii seeds were significantly heavier than dormant seeds, and germinable seeds heavier than 0.20 mg germinated more rapidly than those smaller than 0.20 mg.     

Germination response to fire-related factors of seeds from non-serotinous and serotinous cones

Pinus halepensis, a Mediterranean pine tree, is a partially serotinous species: individual trees of this species carry both non-serotinous and serotinous cones. Serotinous cones open mainly after fire, whereas non-serotinous cones open in absence of fire. In this study we addressed the question, whether or not this cone response is linked with the germination response of seeds to fires. Two main factors associated with fire are heating of seeds and soil pH. A combination of high heat and high pH simulates a scenario with fire, whereas low heat and low pH simulates a scenario without fire. We assessed the separate and combined effects of heat and pH on the germination rate and the percentage of germination of seeds from non-serotinous cones and two age classes of serotinous cones of P. halepensis. Heat had no effect on the percentage of germination of seeds from any of the cone types, but did positively affect the germination rates of seeds from both age-classes of serotinous cones. High pH negatively affected the germination rate of seeds from all cone types as well as the percentage of germination of seeds from non-serotinous cones. The combinations of heat and pH had different effects on the three cone types: percentage of germination and rate of germination of seeds from non-serotinous cones was higher in the combination high heat-high pH than in the combination low heat-low pH. In the combination high heat-high pH, seeds from serotinous cones germinated better than seeds from non-serotinous cones. The different germination responses of seeds from non-serotinous and serotinous cones could not be attributed to differences in cone age. Our results indicate that the cone response is linked to the germination response of the seeds in P. halepensis, with seeds from serotinous cones being more tolerant to fire related factors. This revised version was published online in June 2006 with corrections to the Cover Date.     

Seed size and seed germination in the Mediterranean fire-prone shrub <Emphasis Type="Italic">Cistus </Emphasis><Emphasis Type="Italic">ladanifer</Emphasis>

The germination response of different sized seeds from individuals of a Mediterranean fire-prone shrub (Cistus ladanifer) was investigated in relation to pre-germination heating. A control (no heating), a low temperature during a short exposure time (50°C during 5 min), a high temperature during a short exposure time (100°C during 5 min) and a high temperature during a long exposure time (100°C during 15 min) were applied to seeds from different individual plants with different mean seed weight. These pre-germination treatments resemble natural germination scenarios for the studied species, absence of fire, low intensity pasture fire, typical Mediterranean shrub fire, and severe fire with high fuel load. Mean seed weight only showed a marginally significant positive correlation with the proportion of germinated seeds whatever the pre-germination treatment. These results suggest that seed dormancy is unrelated to seed size and that under the experimental conditions used in this study, the effect of seed size on seed germination is low. Nevertheless, larger seeds could be favoured in natural conditions, especially under the high competition scenario which arise after wildfires. Control seeds showed a negative correlation between seed size and germination velocity suggesting that lighter seeds could take advantage from early germination in recruitment events in the absence of wildfires. Nevertheless, even the lower pre-germination heating treatment turns this correlation in not significant, suggesting a strong selection pressure (unrelated to seed size) for early germination after fire events. In our study, different sized seeds of C. ladanifer seem to perform better under different germination scenarios suggesting that seed size variation could be maintained by the alternation of recruitments without wildfires and recruitments after wildfire events.