Time of germination and establishment success after fire of three obligate seeders in a Mediterranean shrubland of central Spain

Aim Determining how differences in time of germination can affect plant establishment in plant communities that, after a disturbance, must reestablish from seeds under climatic conditions subject to extremes, such as the Mediterranean. Although early germination may be beneficial for survival in summer, when drought is severe, this may expose the seedlings to winter extremes, thus to higher mortality. Understanding how sensitive is the establishment of different species to temporal patterns of germination will help to understand the factors that control species distribution and community stability in disturbance‐prone environments, as well as its sensitivity to changes in weather patterns as climate changes. Methods An experimental fire was made in early fall in an old Cistus–Erica shrubland in Toledo (central Spain). After fire, germination, survival and growth of the three dominant seeder species (Cistus ladanifer, Erica umbellata and Rosmarinus officinalis) were monitored during the first 3 years after fire. Seedlings were tagged to identify their time of emergence, and divided into cohorts according to their month of germination. Differences in survival of the various cohorts were evaluated by means of a Wilcoxon (Gehan) statistic. Height of surviving, tagged plants was compared among cohorts by means of a Kolmogorov–Smirnov test. Results The year following fire was one of the driest on record, while the next one was one of the wettest. Germination was more abundant during the first than during the second year. Establishment was mainly from first‐year germination, as the majority of second‐year germinated seedlings died. Temporal patterns of germination within a year and between years varied between species. Seedling mortality was highest immediately following germination, not in summer. Mortality was related to time of germination: during a given period of time, the mortality of younger seedlings was higher than that of older ones. However, survival was not highest for the first cohorts. In general, the earlier the seedlings germinated the more vigourous they became, more clearly so for Cistus than for Rosmarinus, but differences tended to disappear with time. Overall, time of germination varied between species and affected differently seedling survival and vigour of the various species. Rosmarinus and Cistus had sufficient survivors to reestablish the initial population. Erica, despite abundant germination, suffered a strong population reduction. Main conclusions Mediterranean shrub species differ in their temporal patterns of germination and survival after fire. The effect of time of germination is complex: germinating early is advantageous since old seedlings fared better than younger ones when confronted with the same rigours. However, germinating early might expose the seedlings to greater hazards and the first cohort might not survive best. The temporal window for establishment is narrow and mainly restricted to the first year after fire. Second year seedlings, irrespective of most favourable conditions, survived very little. Missing the window of establishment might virtually lead to a population collapse, despite having very high germination, as found for Erica.     

Early emergence increases survival of tree seedlings in Central European temperate forests despite severe late frost

Global warming is expected to result in earlier emergence of tree seedlings that may experience higher damages and mortality due to late frost in spring. We monitored emergence, characteristics, and survival of seedlings across ten tree species in temperate mixed deciduous forests of Central Europe over one and a half year. We tested whether the timing of emergence represents a trade‐off for seedling survival between minimizing frost risk and maximizing the length of the growing period. Almost two‐thirds of the seedlings died during the first growing period. The timing of emergence was decisive for seedling survival. Although seedlings that emerged early faced a severe late frost event, they benefited from a longer growing period resulting in increased overall survival. Larger seedling height and higher number of leaves positively influenced survival. Seedlings growing on moss had higher survival compared to mineral soil, litter, or herbaceous vegetation. Synthesis. Our findings demonstrate the importance of emergence time for survival of tree seedlings, with early‐emerging seedlings more likely surviving the first growing period.     

亚热带中山常绿阔叶林木本植物幼苗数量动态及其与生境的相关性

以百山祖自然保护区5 hm2永久样地150个幼苗监测站木本植物幼苗为研究对象,2008—2011年定期调查样方中幼苗的种类、数量、萌发、死亡等,探究亚热带中山常绿阔叶林幼苗种类组成、数量动态及其与生境的相关性。结果表明:1)百山祖样地在2008年至2011年出现的幼苗属于26科,40属,共53个物种,不同物种萌发时段有异;2)2009年样地幼苗存活比率为7.7%,2010年为-20.8%,2011年则是-0.3%,幼苗存活比率不高,种类和数量呈减少趋势;3)存活幼苗中有明显的优势物种,分别为光亮山矾(Symplocos lucida)、尖连蕊茶(Camellia cuspidata)、浙闽新木姜子(Neolitsea aurata var.undulatula)、尖叶菝葜(Smilax arisanensis)和短尾柯(Lithocarpus brevicaudatus),5个物种之和占幼苗总数比例50%;4)种子的萌发与生境有极显著的相关性,且与生境因子中水分关联最大;5)存活幼苗数与样站坡位、水分、落叶层厚度呈现显著相关性,水和光照是影响幼苗存活的主要因素。     

Soil seed densities and emergence patterns in pastures in the seasonally dry tropics of northeastern Australia

Germinable seed densities in the surface (0–10 cm) soil of pasture communities growing at Lansdown, near Townsville, were measured during the late dry season before the first germinating rain and again during the following wet season after germination but before the input of new seed. Seedlings emerging in the field were counted at approximately weekly intervals during this period to determine emergence patterns. Twelve communities were sampled in 1980–81 and six were re-sampled in 1981–82. During the late dry season germinable seed densities ranged from 5000 to 40 000 seeds m^-2. Seeds of the introduced legume, Stylosanthes hamata, were present in all pastures. There were many seeds of annual grasses (Digitaria ciliaris and Brachiaria miliiformis) and sedges (Cyperus and Fimbristylis spp.) but only few seeds of perennial grasses (both native and introduced). Soil seed densities were much lower during the wet season than during the preceding dry season, particularly for the grasses. Emergence commenced and approximately 70% of all seedlings emerged on the first major rainfall of the wet season. The subsequent emergence pattern varied between years. In 1980–81 there was a gradual and continuous increase in seedling numbers under the continuously moist conditions which prevailed. In 1981–82 further emergence occurred in discrete events related to rainfall and intervening dry periods. Maximum seedling densities exceeded 34 000 seedlings m^-2 including 29 000 grass seedlings (mainly annual species). The implications of these results for species survival and pasture composition are discussed.     

ECOLOGICAL CONSTRAINTS TO SEEDLING ESTABLISHMENT ON THE PUMICE PLAINS,MOUNT ST. HELENS,WASHINGTON

For both Epilobium angustifolium and Anaphalis margaritacea, the two dominant species in primary succession on the Pumice Plains on Mount St. Helens, density of the seed rain far exceeds the density of colonists. To test the hypothesis that colonization rate is limited by processes occurring during seedling establishment, we investigated the effects of subsurface moisture (using a wet and a dry site), microtopography (fine and coarse pumice particles), and seed size (five size classes), and monitored the fates of seedlings originating from experimentally sown seed for 2 years. Subsurface moisture had the strongest effect on seedling emergence and survival. By the second year there were over 20 times as many seedlings in the wet site as in the dry site; survival in the latter site was nearly zero. Coarse pumice plots had greater establishment than did fine pumice plots. Emergence rate increased significantly with increasing seed size, but this initial difference disappeared by the second year. A. margaritacea established under a wider range of conditions than did E. angustifolium. We conclude that colonization by these species, and hence rate of primary succession, is limited by availability of safe sites for germination and establishment and not by seed dispersal. Safe sites are defined primarily by the level of subsurface moisture and secondarily by microtopography and seed size, and have a greater effect on seedling emergence than on seedling survival.     

Differential effect of shade,water and soil type on emergence and early survival of three dominant species of the Atacama Desert

Understanding the regeneration niche of species may allow us to gain insight into how communities are structured. In deserts, the regeneration niche is usually related to spaces beneath shrubs where shade cast by shrubs creates microenvironments that benefit seedlings and where even small amounts of rain may favour germination and establishment. Shade and water may also interact with different types of soils. However, species may have different requirements for germination and seedling survival. We could expect that shrub species with different drought tolerances exhibit different responses to the combination of these factors. We ask if responses of dominant species of the Atacama Desert to abiotic factors (shade, water and soil type) are related to their drought tolerance, a topic not exhaustively explored in shrubs growing in true deserts. We conducted two factorial experiments. The first one was designed to evaluate how shade (microhabitat) in combination with water may affect germination (emergence) and early survival. In the second experiment, we assessed the influence of shade in relation to soil type. Each species responded distinctively to the three variables under study, but in general, their emergence responses were more influenced by water (more water, greater emergence) than by microhabitat or soil type. Survival was influenced both by microhabitat and by water and was higher under shade and abundant water. Soil type affected only one of our species in terms of emergence. Species responses in general depended on their tolerance to stress. In one species, there was indication of a seed–seedling conflict. Our results show similar species responses to environmental constraints but also more or less unique responses that are related to their tolerance to drought and which may ultimately permit species coexistence. We found that shade may not be important for germination but may be crucial for survival in dry years.     

Episodic recruitment of the seedling banks in balsam fir and white spruce

• Premise of the study: In ecosystems where seed production is low and masting years are sporadic, or with species that have short-lived seeds, regeneration is assured by seedling banks rather than seed banks. Seedling establishment and survival play a critical role in determining the composition of these plant communities by supplying new individuals for their maintenance. Seedling emergence and mortality were investigated to test the hypothesis that recruitment into the seedling bank is periodic. • Materials and methods: Seed production and seedling emergence and survival was monitored during 1994-2007 in balsam fir (Abies balsamea) and white spruce (Picea glauca) in four pristine stands of the boreal forest of Quebec, Canada. Measurements were collected twice per month by sampling one permanent plot of 20 × 20 m per stand. • Key results: Seed-rain abundance reached 9 × 10³ seeds m⁻² year⁻¹, and was characterized by synchronous sequences of low and high seed production. New seedlings appeared only during the year following a seed production of at least 1 × 10³ and 1.5 × 10² seeds m⁻² year⁻¹ for balsam fir and white spruce, respectively. Seedlings emerged in July and survived 34-52 d on average, with balsam fir showing a longer lifespan and lower mortality, although 85–99% of seedlings died before completing one year of life. • Conclusions: The emergence of young seedlings was coupled with massive seed rains, which allowed synchronous replenishment of the seedling banks among stands and species, and generated different cohorts, yielding a discontinuous age structure.     

Facilitation and interference at the intraspecific level: Recruitment of Kageneckia angustifolia D. Don (Rosaceae) in the montane sclerophyllous woodland of central Chile

Most of the studies that have evaluated the interplay between interference and facilitation have been done at the interspecific level, whereas studies at the intraspecific level are scarce. The montane sclerophyllous forests of central Chile are dominated by the tree Kageneckia angustifolia, a semi-deciduous species that lose part of its foliage during summer. It has been reported that during winter snow accumulates in lower amounts beneath the canopy of K. angustifolia favoring the recruitment of new individuals compared to open areas (i.e., facilitation effect). However, it has also been reported that the leaf litter accumulated beneath parental trees contains allelopathic compounds that decrease seed germination, suggesting that recruitment beneath parental plants can be disfavored (i.e., interference effect). Hence, this system seems appropriate to assess the net-outcome between facilitative and negative effects during the emergence and survival of seedlings during the first year. In this study, we asked (i) what is the net-outcome between facilitative and interfering effects for K. angustifolia? (ii) does this net-outcome varies with the distance to parental trees? (iii) are positive and negative effects consistent through the seedling emergence and first year seedling survival phases? (iv) what are the main mechanisms behind the observed net-outcome? and (v) which is the optimal microhabitat for successful recruitment of this species? In an experimental plot of 10,000 m², we selected ten K. angustifolia trees and evaluated the effect of leaf litter on the emergence and survival of seedlings produced by experimentally sown seed seeds in three different microhabitats: beneath adult trees, edge of canopy and in open areas. In addition, we sampled three K. angustifolia stands to evaluate the microhabitat where the natural recruitment of this species actually occurring. Results showed that (1) seedling emergence was greater beneath canopy, intermediate in canopy edge and low in open areas, (2) whilst leaf litter significantly reduced seed germination, the magnitude of this negative effect was lower than the positive effect of beneath canopy microhabitat, (3) seedling survival was affected by microhabitats but not by the presence of leaf litter, (4) that the main mechanisms behind the observed patterns are the lower and delayed emergence of seedlings in open areas due to the longer duration of snow cover, decreasing the time to growth before the onset of summer drought, and (5) the greatest natural recruitment of K. angustifolia seedlings occurs beneath parental plants. Therefore, our findings suggest that the net-outcome between facilitative and interfering effect during the first year is mostly facilitative, indicating that adult trees of K. angustifolia are exerting a conspecific nurse effect on the recruitment of new individuals, a form of parental care in plants.     

Linking seed dispersal,germination and seedling recruitment in the invasive species <Emphasis Type="Italic">Berberis darwinii</Emphasis> (Darwin’s barberry)

Seedling recruitment is a multi-phased process involving seed production, dispersal, germination, seedling establishment and subsequent survival. Understanding the factors that determine success at each stage of this process is of particular interest to scientists and managers seeking to understand how invasive species spread and persist, and identify critical stages for management. To understand the factors and processes influencing recruitment of the invasive species Berberis darwinii Hook. (Darwin’s barberry), temporal and spatial patterns of seed dispersal, germination and seedling establishment were examined. Seed dispersal from a large source population was measured over two fruiting seasons, and subsequent patterns of seedling emergence and survival within each cohort were measured. Seed longevity was tested under both natural and artificial conditions. Seeds were widely dispersed by birds, up to 450 m from the source population. Dispersal was essential to seedling establishment, as few seedlings survived beneath the parent canopy. Seeds were relatively short-lived in the soil under both field and glasshouse conditions, with few surviving for more than 1 year. Patterns of newly emerged seedlings largely reflected patterns of seed rain, but seedling survival was significantly affected by distance from source population, seedling density and light environment. These results suggest that recruitment of B. darwinii is dependent on dispersal of seeds to favourable microsites. Management priorities should include the removal of fruiting plants, and seedling control in highlight areas.     

Mechanisms of structural change derived from patterns of seedling emergence and mortality in a semi-natural meadow

Questions: Is seedling emergence limited by the set of viable seeds, by incompatibility between the phenology of seed shedding and timing of mowing, or by dry weather in germination periods? Does seedling mortality fluctuate with season and weather? Location: Negrentino, southern Alps, Switzerland. Methods: Fecundity estimates of the dominant grass Bromus erectus; highly frequent counts of spontaneous seedlings by species and calculation of a community-level average mortality rate across 5 years; species-level records of seed shedding date and measurements of seed mass; measurement of soil moisture. Results: B. erectus produced 143.9 viable seeds/m²/year while the density of its seedlings was a 55 times smaller fraction. Grasses had fewer seedlings than forbs and their phenology of seed shedding was less compatible with mowing date. Soil moisture was a strong determinant of seedling emergence in spring and less so in autumn. Average seedling mortality declined with age of the populations and reached a maximum in an extremely dry summer. In relatively wet summers establishment success was positively related to seed mass. Conclusion: Community structure is susceptible to drought through mechanisms that selectively reduce recruits of coexisting plant functional groups. We propose that (1) more frequent intense droughts tend to reduce species that depend on frequent recruitment from seed, hence favour long-lived clonally spreading species, (2) drought timing selects between species with different germination phenology and drought resistance, and (3) drought impacts can be mitigated by changing management regimes that affect seed shedding.     

Conditions for seedling establishment and probable function of seed in the clonal sedge Carex rugulosa Kük. in riverside marshes

Carex rugulosa Kük. forms large meadows in moist zones along estuarine riversides. These meadows are usually maintained by rhizomatous ramet production (clonal growth), and the appearance of seedlings is uncommon. We studied the conditions required for seedling establishment. In areas in which clonal ramets were dense (906–1050/m²), and the relative light intensity at ground level was low (0.8–3.8%), seed germination was entirely suppressed. In contrast, many seedlings (288–513/m²) were observed in areas in which clonal ramets had been considerably reduced (13–269/m²) by continuous inundation in the previous year. In these areas, the relative light intensity had increased (20.5–79.3%). It was inferred that seeds resisted the prolonged inundation that killed the ramets, and that germination was induced under these open conditions. These results suggest that the main functions of sexual reproduction in C. rugulosa are recovery after severe damage to vegetation using seeds from the seed bank and the dispersal of seeds to areas without dense vegetation. Most seedlings died when the marshy meadow was temporarily inundated by heavy rain. However, transplanted seedlings survived better at artificially elevated sites that were not submerged. This shows that inundation during the seedling stage impedes seedling survival. Therefore, seeds can contribute to recovery after ramet decline or colonization to open area when (i) the rainfall is not heavy to cause seedling submergence or (ii) seeds are dispersed to higher area which is not inundated.     

毛乌素沙地4种沙生植物种子萌发及出苗对沙埋及单次供水的响应

飞播固沙作为一项快速而有效的现代化治沙措施,已成为毛乌素沙地植被恢复与重建的重要手段。但此项技术仍存在一些问题,如飞播后成苗率较低等。以毛乌素沙地优势植物柠条（Caragana korshinskii）、羊柴（Hedysarum laeve）、油蒿（Artemisia ordosica）与籽蒿（Artemisia sphaerocephala）为研究对象,人工控制7个沙埋深度和9个水分梯度,研究4种植物种子萌发和幼苗出土对沙埋和单次供水的响应,阐明多大的单次降雨才能保证种子萌发与幼苗出土及出土幼苗的维持时间。结果表明,对于柠条和羊柴质量相对较大的种子而言．种子主要在供水量10．20mm,埋藏深度为0．5．2cm的条件下出苗,但在埋藏深度为5cm时这两个物种仍有少量种子能够出苗;而对于油蒿和籽蒿质量相对较小的种子而言,种子主要在供水量10．20mm,埋藏深度为0．5cm时出苗,当埋藏深度为1．5cm和2cm时不能顺利出苗。沙埋深度0．5cm条件下,4个物种具有最高的出苗率和出苗速率。在试验期内（30d）,柠条、羊柴和油蒿出土幼苗分别在单次供水量为5、7．5mm和7．5mm时全部死亡,在单次供水量高于15mm时4个种的出土幼苗死亡率大多低于50％,且随供水量的增加出土幼苗死亡率降低。结合毛乌素沙地降雨特点与本次实验结果,0．5cm的沙埋及7．5mm以上的单次降雨是上述4种植物自然条件下出苗较好的重要条件。     

Acacia karroo invasion of grassland: environmental and biotic effects influencing seedling emergence and establishment

Acacia karroo Hayne is the most important woody invader of grassland in South Africa, and can greatly reduce the productivity of grassland. A field experiment was conducted to test the hypotheses that emergence, growth and the 1st year's survival of Acacia karroo would be enhanced by (1) defoliation of the grass sward, (2) increased irradiance, (3) increased moisture availability and (4) its germination within cattle dung pats. The study was conducted on one site above and one below the natural altitudinal treeline of this species in grassland of the eastern Cape, South Africa. Not one seedling emerged from dung pats. Neither location nor the other treatments affected the density of emerging seedlings, although only 40.4 seedlings m^–2 emerged of the 200 seeds m^–2 planted. Shading dramatically increased the density of surviving seedlings. In the open, only 3 and 1.5 seedlings m^–2 remained respectively at the end of the growing season or the beginning of the next, compared to 23.3 and 19.5 seedlings m^–2 under shading for these respective times. This was attributed to the effect of shade on moisture availability in a season which received only 54% of average rainfall. Seedling survival until the end of the growing season was enhanced (30%) by shade at both sites, but also by supplemental water (24%) and defoliation of the sward (7%) at the site above the treeline. Across sites and treatments, seedling survival was related to moisture availability, with no or poor survival for < 500 mm rainfall, indicating this species can only establish in certain rainfall years. Seedling survival over winter was not influenced by treatment, but was greater for larger seedlings. Treatments affected seedling size, in particular seedlings growing under shade and within a dense grass canopy were etiolated. A. karroo seedlings are capable of establishing and surviving within a dense grass sward for at least a year, tolerant of low irradiance and of interference, which, because most seeds do not persist for much longer than a year, suggests this species forms predominantly a seedling bank. This has implications for the invasion of grassland by woody species.     

Effects of vegetation canopy and climate on seedling establishment in Mediterranean shrubland

Abstract. Question: Does the influence of plant canopy on seedling establishment interact with climate conditions, and particularly, do intensified drought conditions, enhance a positive effect of the vegetation canopy on seedlings in Mediterranean‐type ecosystems. Location: Mediterranean shrubland near Barcelona, Spain at 210 m a.s.l. Methods: Over the course of four years we recorded seedling emergence and survival in open areas and below vegetation under control, drier and warmer experimental climatic conditions. Results: Seedling emergence is more sensitive to climate conditions than later stages of growth. When considering the whole set of species, the total number of established seedlings at the end of the experiment was lower in the drought and warming stands than in control ones, and vegetation canopy increased the number of these seedlings in the drought stands. Drought reduced seedling emergence but not warming, while the interaction between climate treatments and vegetation canopy was not significant. Seedling survival was lower in the warming treatment than in the control. Under drought conditions, vegetation canopy increased seedling emergence of the dominant Globularia alypum. In control stands, vegetation canopy reduced their survival. Vegetation canopy increased the survival of the dominant Erica multiflora in warming stands, and it reduced the survival of G alypum in drought stands. No significant effects of drought and warming were observed in the seed rain of these two species. Conclusions: The balance of the facilitation‐competition interactions between vegetation canopy and seedling establishment in Mediterranean‐type ecosystems determined by water availability, and drought conditions enhance the positive effect of vegetation canopy. This interaction is species‐specific and shows important between‐year variability.     

The influence of climatic variability on local population dynamics of Cercidium microphyllum (foothill paloverde)

This study investigated correlations among climatic variability, population age structure, and seedling survival of a dominant Sonoran Desert tree, Cercidium microphyllum (foothill paloverde), at Tucson, Arizona, USA. A major goal was to determine whether wet years promote seedling establishment and thereby determine population structure. Plant age was estimated from basal circumference for a sample of 980 living and dead trees in twelve 0.5-ha plots. Ages ranged from 1 to 181 years. Age frequency distribution showed that the population is in decline. Most (51.2%) of the 814 living trees were 40-80 years old; only 6.5% were younger than 20 years. The average age of the 166 dead trees was 78 years. Fifty-nine percent of dead trees were aged 60-100 years. Survival of newly emerged seedlings was monitored for 7 years in a 557-m² permanent plot. Mean survival in the 1st year of life was 1.7%. Only 2 of 1,008 seedlings lived longer than 1 year. Length of survival was not correlated with rainfall. Residual regeneration, an index of the difference between predicted and observed cohort size, showed that regeneration was high during the first half of the twentieth century and poor after the mid-1950s. Trends in regeneration did not reflect interannual variation in seasonal temperature or rain before 1950, that is, in the years before urban warming. Taken together, the seedling study and the regeneration analysis suggest that local population dynamics reflect biotic factors to such an extent that population age structure might not always be a reliable clue to past climatic influences.     

Irregular rain cues and the germination and seedling survival of a Panamanian shrub (Hybanthus prunifolius)

Summary Seeds of the Panamanian shrub, Hybanthus prunifolius (Schult.) Schulze (Violaceae) are dispersed at different times in different years ((March to June) and are exposed to the irregular rainfall of the dry season in some years. Fluctuations in soil moisture in the dry season represent suboptimal conditions for germination and seedling survival. There are no mechanisms to prevent germination prior to the arrival of consistent rains in the wet season.Among three natural cohorts of seeds followed in two years, the cohort experiencing the longest time from sowing to consistent rains had the highest germination, but it also had the longest time lag from sowing to, beginning of germination, longest germination period, and lowest survival of seedlings 3 months after sowing.Seeds were also induced experimentally to germinate under 14 different moisture patterns. The patterns encompassed 1) varying lengths of moisture before a dry period, 2) inconsistent moisture, and 3) varying lengths of dryness prior to any moisture.Mortality of seeds by fungal infection occurred if the wet period was delayed. But germination was less affected by fluctuations than was seedling survival. Length of the first wet period and frequency of occurrence of the wet period both affected germination levels. Survival and development of seedlings was influenced by the number of days exposed to dry conditions and by the stage of development at the beginning of the dry period. Young seedlings suffered attrition due to drought stress, and older seedlings died from fungal attack.Results from field and experimental sowing of seeds both indicate that this perennial species has minimal defense against germination when conditions are suboptimal for seedling survival. Undoubtedly there is more recruitment in some years than in others.     

Demography of woody plants in relation to fire: Banksia ericifolia L.f. and Petrophile pulchella (Schrad) R.Br.

Plants of Banksia ericifolia and Petrophile pulchella are sensitive to fire. Changes in population size under different fire regimes were estimated, based on measurements of post-fire seedling emergence, seedling survival, survival and seed production in established plants of differing ages, survival of seeds held in serotinous cones and seed-release in the periods between fire. Seeds were first available at 5 years in P. pulchella and 6 years in B. ericifolia. Exact replacement would be possible when burnt at these ages, if seedling establishment were very high. Low establishment would delay replacement to 13 years of age in both species. Late summer/autumn fires of high intensity favour high establishment. Such fires at 8–10 year intervals would be tolerated without any sustained decline in numbers. Fires at 10–15 year intervals could occur regardless of season or intensity with little risk of a population decline. Large increases in numbers and density would follow fires spaced at 15–30 years. Enough seeds would be available for replacement up to about 50 years in both species. Viable seed-release in unburnt conditions was sufficient to compensate for deaths in stands over 20 years old, even with very low levels of establishment. Two variables accounted for the biggest changes in numbers and density between generations interspersed by fires; namely the age at which a stand is burned and the proportion of seeds which emerge as seedlings. A comparison with other similar species showed similarities in controls on emergence and establishment, lengths of primary juvenile periods and life spans.     

Recruitment in a Mediterranean annual plant community: seed bank, emergence, litter, and intra- and inter-specific interactions

A small‐scale field study was conducted to explain seedling emergence and recruitment of reproductive individuals in a four‐year‐old Mediterranean annual plant community. The analyzed levels were populations, functional types, and total number of individuals in the community. We hypothesized that the number of germinable seeds positively affects the number of emerged seedlings, which positively affects the number of reproductive individuals. We hypothesized as well that litter mass, biomass of established adults, and number of non‐conspecific emerged seedlings negatively affect the numbers of emerged seedlings and reproductive individuals. The results supported our multivariate causal explanation of plant recruitment, although concordances with the initial hypotheses were more frequent at the community level. The effect of the number of germinable seeds on the number of emerged seedlings was found to be robust only at the community level. At the population level, this relationship did not have a consistent tendency and depended on species identity and range of seed density experienced. Litter had negative effects on total number of individuals in the community, and usually non‐significant effects at the population level. Litter effects were found to be inversely related to seed mass, and sharper cotyledons did not improve seedling emergence likely by favoring litter mat penetration. Interactions among plant species appeared not to have any effect on seedling emergence and survival, and had positive effects on certain populations. Biomass of established adult plants exerted no influence on seedling emergence and survival, probably due to their low abundance in this community during the germination period. Results permitted the inference of the main stages in the recruitment process and causal factors. They provided evidence for the greater importance of germination and emergence in comparison with seedling survival to insure reproductive recruitment in this community.     

Facilitation and competition influence succession in a subtropical old field

The effects of neighboring vegetation and soil fertility on the establishment, survival and growth of tree species were studied in a subtropical old-field area in south Brazil. Seed damage, germination and seedling establishment of four tree species plus growth and survival of two transplanted tree species were monitored under factorial combinations of the following treatments: (1) pioneer vegetation (presence and absence); (2) soil fertility (addition of NPK and control). Facilitation was the main process affecting plant performance. The presence of pioneer vegetation significantly improved germination, establishment, growth and survival of most study species. Around 90% of sown seeds were damaged and the removal of pioneer vegetation significantly increased seed damage for all species studied, decreased germination in three out of four species, and decreased establishment in one species. Moreover a significantly higher seedling growth rate of Inga virescens was found after the first year of the experiment in plots where vegetation was present. The presence of vegetation significantly increased seedling survival of I. virescens by protecting seedlings from leaf loss due to winter frosts. Competition was detected by the second year when a higher growth of transplanted seedlings of the species Araucaria angustifolia occurred in plots where vegetation was present and fertilizer were applied. A lower growth rate was detected in plots where vegetation was present but fertilizer was not applied. These results indicate a balance between competition for soil nutrients and protection by neighbor vegetation. Damage of seedlings by leaf cutter ants was an important barrier for plant survival. Damage occurred in 80% of the A. angustifolia seedlings and 58% of these damaged seedlings died. The presence of neighboring vegetation tended to protect seedlings from ant damage. Although competition occurred, facilitation seems to be the main process driving early successional changes in this subtropical old field. This was mainly due to the improvement of local microclimatic conditions and protection against herbivores by neighboring vegetation. Facilitation occurred during establishment and growth phases in a subtropical area that is considered a productive, low stress environment. Our results indicate that facilitation may be more frequent in productive environments than previously thought.     

Seedling emergence patterns and dormancy/germination physiology of <Emphasis Type="Italic">Primula modesta</Emphasis> in a subalpine region

On the basis of the germination/dormancy responses of seeds to temperature and light, and local seed rain, we attempted to interpret the seedling emergence patterns of Primula modesta Bisset et Moore (Primulaceae) in two different types of habitats in a subalpine zone of Mt. Asama: an oligotrophic flat moor and a grassland with relatively dense herbaceous vegetation cover. The seasonal pattern of seedling emergence was well explained by the dormancy/germination physiology revealed in laboratory germination tests. The seeds were demonstrated to have a strict light requirement even after experiencing moist chilling, which might facilitate the incorporation of the seeds into the soil seed bank. Despite sufficient seed production, the seedlings emerging were far less at the grassy site than the moor site, but the number of seedlings was significantly dependent on the seed rain within previous season and on the litter cover of the microsite in both sites. Therefore, the spatiotemporal patterns of seedling emergence in the habitats could be well explained by the spatiotemporal patterns of seed rain and safe-sites for germination.