Litter effects on seedling establishment interact with seed position and earthworm activity

Seedling establishment is influenced by litter cover and by seed predators, but little is known about interactions between these two factors. We tested their effects on emergence of five typical grassland species in a microcosm experiment. We manipulated the amounts of grass litter, seed sowing position and earthworm activity to determine whether: (i) the protective effect of litter against seed predation depends on cover amount and seed sowing position, i.e., on top or beneath litter; (ii) seed transport by earthworms changes the effect of seed sowing position on seedling emergence; and (iii) seeds transported into deeper soil layers by earthworms are still germinable. Litter cover and presence of earthworms lowered seedling emergence. The impact of seed position increased with seed size. Emergence of large-seeded species was reduced when sown on the surface. Additionally, we found an important seed position × earthworm interaction related to seed size. Emergence of large-seeded species sown on top of the litter was up to three times higher when earthworms were present than without earthworms. Earthworms also significantly altered the depth distribution of seeds in the soil and across treatments: on average 6% of seeds germinated after burial. In contrast to the seed position effect, we found no size effect on mobility and germinability of seeds after burial in the soil. Nevertheless, the fate of different-sized seeds may differ. While burial will remove large seeds from the regeneration pool, it may enhance seed bank build up in small-seeded species. Consequently, changes in the amount of litter cover and the invertebrate community play a significant role in plant community composition.     

Effects of experimental and real land use on seedling recruitment of six fen species

Seedling emergence and early establishment of six fen species differing in seed mass and growth form were investigated under experimental land use with changed vegetation structure and under real land use in a calcareous fen. Seeds of all six species were sown in plots with different experimental land-use treatments: summer and autumn mowing with or without litter removal, trampling and abandonment. Additionally, emergence and survival of experimentally sown seeds was investigated under real land use on adjacent sites managed by mowing, grazing, intense trampling or abandonment.On abandoned plots and on plots without litter removal of the land-use experiment, emergence rates of all species were negatively affected either by high litter and moss cover or by tall canopy. No differences were found between autumn and summer mowing. Gap creation by experimental trampling did not increase germination rate. Under real land use, establishment of seedlings of most species was positively affected by litter cover and tall canopy. Trampling, in contrast, had a severe negative effect on seedling survival.The investigated species differed in their germination ability which was tested in the germination chamber and in their response to land use. Succisa pratensis with the highest seed mass germinated well in the chamber and in the field more or less regardless of land use. The low germination rate of Parnassia palustris in the germination chamber indicated a limitation of viable seeds. In the field, however, seedling emergence was additionally limited by microsite availability. Seeds of Serratula tinctoria and Primula farinosa germinated well in the germination chamber, but seedling recruitment in the field was hampered in the presence of a high litter or moss cover. Seeds of Tofieldia calyculata and Pinguicula vulgaris were strongly dependent on the availability of suitable microsites in the field. They hardly germinated under natural conditions, in spite of a high number of germinable seeds in the germination chamber.     

Seedling recruitment of Succisella inflexa in fen meadows: Importance of seed and microsite availability

Abstract. The effects of different forms of land use on germination and establishment of the rare fen species Succisella inflexa were investigated in seed introduction experiments in a mown and an abandoned fen meadow in SE Germany. Treatments included abandonment, mowing in fall and mowing with creation of gaps in the moss and litter layer. Floating capacity of seeds was tested in order to estimate potential dispersal by water. On the mown meadow, gaps had a slightly positive effect on germination rates, while greatly increasing seedling survival until the next spring. At the abandoned site, litter inhibited germination, whereas mosses had a negative effect on germination and a positive effect on survival rates during the first year after germination. Both germination and seedling establishment were negatively affected by the presence of slug herbivores. On the abandoned site, no seedlings at all survived until the next spring. Even though seeds of Succisella inflexa were capable to float for several weeks and to germinate thereafter, the situation at the field sites indicates that longdistance dispersal is highly unlikely. Our results showed that not only direct effects of abandonment, such as accumulation of litter, may have led to poor germination and poor seedling establishment of the species. Additionally, indirect consequences of changes in land use, such as higher seedling herbivory by slugs and successional vegetation changes due to abandonment, were important in determining habitat quality and availability of microsites for seedling recruitment. Furthermore, early mowing imposed seed limitation on plant populations.     

Establishment and reproduction of Aeschynomene virginica (L.) (Fabaceae) a rare,annual, wetland species in relation to vegetation removal and water level

Aeschynomene virginica is a rare annual plant found in freshwater tidal wetlands of the eastern United States. We hypothesized that standing vegetation and water inundation were two important environmental factors in its population dynamics. To test these hypotheses, we sowed seeds into plots with undisturbed vegetation or plots with all aboveground vegetation removed in 1998 and 1999. Presence/absence of seedlings was noted and seedling survival to reproduction, final size, and seed set were measured throughout both growing seasons. Seedling establishment from germination to the first true leaf stage increased with decreasing water depth. Vegetation removal plots had greater seedling establishment, higher seedling survival, and higher seed set per plant than non-removal plots. In a greenhouse study designed to test the effects of water level on seed germination and seedling establishment, no seedlings established in submerged soils, and seed germination and seedling establishment were lower in waterlogged soil than in wet soil. Physical stress associated with deeper water likely limits the distribution of A. virginica to higher elevations, where seeds that colonize patches with low vegetative cover are more likely to produce reproductive adults that produce more seeds relative to patches with established vegetation. A. virginica appears to be a fugitive species specializing on open habitat patches in tidal wetlands. This species may be dependent on disturbances for population establishment and maintenance.     

Seed size and seedling emergence in 16 temperate forest herbs and one dwarf-shrub

The impact of seed size (seed mass) on seedling emergence beneath a leaf litter layer and post-dispersal seed predation was investigated in two field experiments including 16 forest herbs and one dwarf-shrub in southeastern Sweden. In the first experiment, I studied the relationship between seed mass of eight forest herbs (0.3–16.7 mg) and seedling emergence after removal of litter and reduction of seed predation (rodents and insects). Removal of litter and reduction of seed predation did not affect seedling emergence. However, regardless of treatment, species with large seeds (> 3 mg) had a higher seedling emergence than those with small seeds (< 1 mg). In the second experiment, I investigated the relationship between seed mass of 12 species (0.007–18.4 mg) and seedling emergence after removal of litter, reduction of seed predation (insects) and seedling herbivory (molluscs). Total emergence over three years was significantly higher in species with large seeds (> 3 mg) than in those with small seeds (< 2 mg). Removal of litter increased total seedling emergence, while application of insecticide and molluscicide had no effect. Similar results were obtained from both a deciduous and a mixed coniferous forest, but seedling emergence was in general higher in the mixed coniferous forest. Seedling emergence in temperate forest herbs and dwarf-shrubs seems to be higher in species with large than in those with small seeds, and it is often enhanced by disturbance.     

Experimental demography of the rare Gentianella germanica: seed bank formation and microsite effects on seedling establishment

The persistence of populations of short-lived species requires regular reproduction and seedling establishment. A persistent seed bank can buffer populations against extinction in unfavourable years. We experimentally investigated seed fate in Gentianella germanica, an endangered biennial species characteristic for species-rich nutrient-poor calcareous grasslands in central Europe. We studied the effect of experimental gaps on seedling establishment from sown seeds and the fate of seeds buried in bags over two years. In December 1993 experiments were established at seven calcareous grassland sites in the Swiss Jura mountains. In spring 1994 seedlings emerged in all plots where seeds had been sown, including previously unoccupied patches. This suggests that limited dispersal within sites contributes to small population sizes. Significantly more seedlings emerged at sites with current populations of G. germanica than at unoccupied sites (5.95% vs 3.40%). Because this difference was not explained by germinations from the natural seed bank it indicates differences in habitat quality. Clipping of the vegetation and disturbance of the soil reduced vegetation cover in the following spring and enhanced seedling emergence. In undisturbed plots 4.5% of seeds sown produced a seedling in spring 1994, whereas in plots with clipped vegetation 9.9% and in disturbed plots 12.7% produced seedlings (p>0.01). This suggests that management measures which create gaps in the vegetation (e.g. grazing) could positively influence population size and persistence of G. geymanica. On average, we recovered 7.55% viable seeds after one year of burial in bags, and 4.05% after two years, indicating that G. geymanica has a persistent seed bank. The demographic data indicate that the number of viable seeds in the seed bank exceeds the number of established plants in a population at least by a factor of 20. Restoration of extinct populations of the species from the seed bank may thus be possible if appropriate management measures are taken within a few years.     

Seed deposition in drift lines: Opportunity or hazard for species establishment?

Arrival at suitable sites for germination and establishment is crucial for hydrochorous plants. Seed deposition in riparian landscapes occurs mainly within litter in drift lines. We investigated whether conditions within drift lines are suitable for seed survival and germination and whether drift lines act as seed sources or sinks for species diversity in riparian fen grasslands.We analysed drift litter in a small river valley in Northern Germany (Upper Eider valley) after typical late winter floods in 2000, 2002 and 2004. The content of viable seeds in drift lines was studied via a germination trial with drift litter samples taken from five fen grassland sites representative of the vegetation of floodplain. Further, seedling recruitment of both abandoned and moderately grazed fen grasslands affected by drift line deposition was compared with equivalent sites without drift lines. Additionally, the decline of viable seeds after the deposition of drift litter was studied. Finally, the relative importance of seed-containing drift litter and microsite availability for seedling recruitment was investigated in a field experiment with the factors disturbance (mowing, large gaps, small gaps, undisturbed control) and addition of (seed-containing) drift litter (yes, no) in an abandoned riparian fen.Seedlings which recruited from drift litter collected at different sites differed in species composition. A nearly 50% reduction of the number of viable seeds in drift litter was observed already during 2 weeks after deposition. Naturally deposited drift lines had a positive effect on the number of seedlings, although an increase of seedling recruitment in areas with drift litter was only observed at the grazed site. Adding drift litter reduced the number of seedlings in the field experiment, while creating gaps enhanced seedling recruitment.These results indicate that in riparian fen grasslands, drift lines can affect species diversity either negatively by hampering germination and/or establishment or positively by increasing seed availability. As the studied river system is comparatively species poor and flooding extension is limited, the role of drift lines as seed source might be of minor importance in the study area.     

Are the best dispersers the best colonizers? Seed mass,dispersal and establishment in <Emphasis Type="Italic">Carduus</Emphasis> thistles

Negative correlations between dispersal and establishment are often reported in the plant literature; smaller seeds tend to disperse better but germinate less well, and produce smaller seedlings. However, because dispersal capacity is often quantified using proxies, such as the settling velocity of wind-dispersed seeds, little is known about the exact shape of this negative relationship, and how it is modified by other plant traits and environmental conditions. We studied the dispersal-establishment relationship in two wind-dispersed thistles (Carduus nutans and Carduus acanthoides). We applied a mechanistic wind dispersal model (WALD) to seeds released under a range of environmental conditions, and tested germination and seedling growth under standardized conditions in a greenhouse. Dispersal distance and establishment (germination and seedling growth) were not significantly correlated, although in both species smaller seeds dispersed farther, and showed lower germination and lower seedling growth rates. This apparent paradox can partly be explained by the significant influence of other factors such as release height and environment (wind and vegetation), which explained more variation in dispersal than did terminal velocity. Another potential explanation is the variation in seed traits: germination is strongly positively related to seed mass, weakly positively related to plume loading, but not significantly related to terminal velocity. This weakening of the correlation with germination is due to additional layers of trait (co)variability: for instance, seed mass and pappus size are positively correlated, and thus big seeds partially compensate for the negative effect of seed mass with larger pappi. Our mechanistic approach can thus lead to a better understanding of both potentially opposing selection pressures on traits like seed mass, and diluting effects of other seed, plant and environmental factors.     

森林凋落物对林地微生境的影响

评述了凋落物改变林地微生境的效应．凋落物减弱了林地地表及表层土壤(2mm厚)中的光照,改变了光谱特性,降低了PPFD和Pfr．缩小了日温差,提高了土壤湿度和肥力,增加了毒素和土壤动物的活动等,但凋落物改变生境的效应强度与凋落物组成、厚度、分解状态和周围生境(如植被、气候、地形和地貌等特性)等有关,在自然森林生态系统中,凋落物对林地微生境的效应因凋落物、植被、气候、地形和地貌的多样化变得更为复杂,而凋落物对植被天然更新的影响也因其对生境改变效应的复杂性而具有多样性和综合性,最后,对凋落物改变生境效应的复杂性和有关研究进行了展望。     

Seed characteristics and soil surface patch type interact to affect germination of semi-arid woodland species

Biological soil crusts are common in many arid and semi-arid regions and they can alter microenvironments which are likely to directly and indirectly influence vascular plant establishment. The effect of biological soil crusts on germination is also influenced by the biological characteristics of the seeds themselves but rarely have the effects of both crust type and seed morphology on germination been examined in the same study. In this study, seed of five semi-arid woodland species with contrasting seed morphology were sown on top of patch types that commonly occur in natural woodlands (foliose lichen, short-turf moss, tree leaf litter, disturbed crust) and their emergence was followed. Percent germination varied between patch types and, for the largest-seeded species (Maireana excavata), final germination was significantly lower on the biological soil crust and litter patch types because they strongly acted as a physical barrier to seed penetration into the soil. The remaining four species showed no significant differences in final percent germination with patch type because most seeds either completely or partially penetrated the surface layer. Germination time courses, however, showed that biological soil crusts delayed the timing of germination of these species. Hence, soil crusts might differentially affect the spatial patterning of species in semi-arid woodlands by their subtle influence on seedling emergence that is determined by differences in seed morphology and subsequent positioning within crusts.     

Conspecific versus heterospecific litter effects on seedling establishment

Plant litter is an important determinant of seed germination and seedling establishment. Positive effects of litter have received considerable attention, but few studies have explicitly tested whether seedlings are more facilitated by conspecific litter compared to heterospecific litter. In order to contrast conspecific and heterospecific facilitative effects on seedling establishment, we used Anthriscus sylvestris, Angelica sylvestris, Pimpinella saxifraga and different combinations of their seeds and litter seedbeds as a model system. Although litter had a significant species-specific effect on seedling emergence, we found no evidence of strictly conspecific facilitation. Anthriscus sylvestris displayed a positive response to all types of litter. In contrast, there was a clear negative effect of conspecific litter in Pimpinella saxifraga. Activated carbon did not modify the negative effect, indicating that chemical compounds were not the cause. Our study suggests a high level of idiosyncrasy in response to litter at the species level.     

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.     

Effects of bryophytes and grass litter on seedling emergence vary by vertical seed position and seed size

Establishment of plants through seeds is often constrained by the quality of microsites, which is in part controlled by the nature and amount of ground cover. The latter consists of living shoots of vascular plants or bryophytes and/or the dead remains of the dominant species. In the present article, we report the results of a controlled pot experiment with five species characteristic of floodplain grasslands. We manipulated the amounts of grass litter and/or mosses to study (1) differences between ground cover types with respect to their effects on microenvironment and seedling emergence and (2) how these effects interact with seed size and seed sowing position. Increasing amounts of both cover types led to increasing soil humidity, whereas temperature amplitude and illumination were decreased. However, since grass litter decomposed much faster than bryophytes, light conditions for germination under grass litter improved considerably with time. Although seedling emergence varied significantly between species, ground cover types and cover amounts, seed position alone explained about 50% of the variation in the data set. Additionally, we found an important interaction between seed size, seed position and cover type: large-seeded species showed a fitness advantage when seeds were situated beneath a cover, irrespective of cover type, which disappeared when seeds were shed on top of a cover layer. We suggest that this interaction may be ecologically and evolutionarily relevant because it may lead to changes in species composition and diversity of plant communities as a consequence of changes in the amount and type of ground cover.     

Effects of the litter layer of Pteridium aquilinum on seed banks under experimental restoration

Questions: Does the litter layer of Pteridium aquilinum (bracken) act as a barrier to certain species in the seed bank? Does bracken control/restoration treatment affect seed transfer through the litter layer? Location: Five experiments at three sites across the UK covering two major vegetation types; acid‐grassland and heath‐land. Methods: At each experiment a range of bracken control and vegetation restoration treatments were applied for about ten years. The seed bank was sampled in both the bracken litter and the soil. The cover (%) of each species in the vegetation and the bracken litter abundance (cover and depth) was also estimated. Results: The bracken litter layer acts as an inert barrier as it contained a large proportion of seeds available in the litter‐soil profile (38%– 67% of the total). Bracken litter depth and cover also influenced significantly the seed bank composition in both the bracken litter and the soil. These effects were site‐specific, and species‐specific. The application of treatments changed significantly the balance between seed inputs and outputs in the bracken litter layer for some species. This was either a positive or negative response relative to the untreated control plots. Conclusion: For heathland and acid‐grassland restoration, the bracken litter layer may be an important seed source, but it must be disturbed particularly before seed addition.     

Seed predation limits post-fire recruitment in the waratah (<Emphasis Type="Italic">Telopea speciosissima</Emphasis>)

Seed predation may reduce recruitment in populations that are limited by the availability of seeds rather than microsites. Fires increase the availability of both seeds and microsites, but in plants that lack a soil- or canopy-stored seed bank, post-fire recruitment is often delayed compared to the majority of species. Pyrogenic flowering species, such as Telopea speciosissima, release their non-dormant seeds more than 1 year after fire, by which time seed predation and the availability of microsites may differ from that experienced by plants recruiting soon after fire. I assessed the role of post-dispersal seed predation in limiting seedling establishment after fire in T. speciosissima, in southeastern Australia. Using a seed-planting experiment, I manipulated vertebrate access to seeds and the combined cover of litter and vegetation within experimental microsites in the 2 years of natural seed fall after a fire. Losses to vertebrate and invertebrate seed predators were rapid and substantial, with 50% of seeds consumed after 2 months in exposed locations and after 5 months when vertebrates were excluded. After 7 months, only 6% of seeds or seedlings survived, even where vertebrates were excluded. Removing litter and vegetation increased the likelihood of seed predation by vertebrates, but had little influence on losses due to invertebrates. Microsites with high-density vegetation and litter cover were more likely to have seed survival or germination than microsites with low-density cover. Recruitment in pyrogenic flowering species may depend upon the release of seeds into locations where dense cover may allow them to escape from vertebrate predators. Even here, conditions suitable for germination must occur soon after seed release for seeds to escape from invertebrate predators. Seed production will also affect recruitment after any one fire, while the ability of some juvenile and most adult plants to resprout after fire buffers populations against rapid declines when there is little successful recruitment.     

种子重量的生态学研究进展

作为植物生活史中的一个关键性特征,种子重量与其它许多植物性状和生态因子有关,种子重量的分异与其它一些植物性状及环境的变化关系在进化生物学上已经成为一个非常有意义的研究内容,且具有一定的实践意义。种子重量被发现与下列的一些植物学和群落学性状有关:植物的生活型、种子的散布能力、种子的散布方式、植物的高度、植物的冠幅、植物的比叶面积、植物的寿命、动物的捕食、植被中植物的数量或多度、土壤中种子的数量或多度、种子的休眠、种子在土壤中的持久性和植物的净初级生产力等,另外生态因子如降雨、温度、坡向、海拔、经度、纬度、光强和干扰等都影响种子的重量。种子的重量被认为是在大量小种子和少量大种子之间的进化折衷,在一定的能量限度内,较大重量的种子一般具有较少的数量,而较小重量的种子一般数量较多,这是种子重量和数量方面具有的一种反向关系。与其它性状相比,很多研究都表明种子重量和植物的生活型的关系密切。没有散布结构或风散布的种子比以动物和水作为散布媒介的种子重量要小。种子重量与捕食的关系现发现有3种格局。种子重量和形状与种子在土壤中的持久性的关系有4种格局。在干旱和阴暗的环境条件下,种子有变大的趋势。大重量种子比小种子赋予幼苗较优势的竞争地位,其原理尚有争论,尚不清楚是否是幼苗阶段的竞争决定了世界上大部分植被类型的物种组成。未来的研究方向主要有以下几个方面:1) 种子重量与植物系统学相结合,探索种子重量的变化规律;2)调查群落三向(纬度、经度和海拔)性的种子重量谱变化规律;3) 群落演替与群落种子重量谱的变化;4) 种子重量与群落中植物个体和种子的数量的关系及机理研究;5) 微生境、微地形如坡向、坡位和林间隙等对种子重量的影响;6) 全球气候变化和种子重量变化的关系。     

Hydrocarbon Tolerance Correlates with Seed Mass and Relative Growth Rate

The abundance and vegetation cover of plant species with different seed masses growing on crude oil contaminated and uncontaminated field plots were examined. In addition, seedling mass and relative growth rate of eight plants in four seed mass classes were compared in contaminated and uncontaminated soils. Species producing seed in the second largest seed class (1.0 to 9.9 mg) were more common on contaminated than uncontaminated field plots. Species with seed in the smallest seed class (< 0.1 mg) were less common on contaminated than uncontaminated plots. Species with seed masses between 1.0 and 9.9 mg formed 15% more vegetation cover, and those with masses between 0.1 and 0.9 mg formed nearly 15% less vegetation cover on contaminated compared to uncontaminated field plots. In the growth chamber, species with larger seeds produced approximately 25% less seedling mass and exhibited a 32% reduction in relative growth rate, when grown in contaminated compared to uncontaminated soil. Small-seeded species had reductions of 95% in seedling mass and 72% in relative growth rate. Species with the highest relative growth rates in uncontaminated soil had the lowest seedling mass in contaminated soil. Seed mass and relative growth rate were negatively correlated (r = 0.866).     

Dung beetle activity affects rain forest seed bank dynamics and seedling establishment

Dung beetles relocate vertebrate feces under the soil surface, and this behavior has many ecological consequences. In tropical forests, for example, seeds defecated by mammals that are subsequently buried by dung beetles are less likely to suffer predation. While the effects of dung beetles on the fate of defecated seeds have been relatively well studied, their effect on seeds already buried in the soil has not. To contribute to fill this gap, we designed a study with three objectives: (a) Describe the vertical re‐distribution of soil seeds that occurs due to dung beetle activity; (b) Determine if beetle activity favors establishment of seedlings from the soil seed bank; and (c) Determine if the effect of dung beetles is stronger in sites of recurrent mammal defecation. We carried out three complementary field experiments, one with artificial seeds (plastic beads) of three sizes buried at known depths, one with two species of seeds buried at those same depths, and one with the natural soil seed bank in sites of single vs. recurrent defecation. Buried beads were moved by dung beetles along the vertical axis, both upwards (9.5%) and downwards (11.5%); smaller beads were more frequently moved downwards while the contrary occurred for larger beads. Dung beetle activity caused an increase in seedling establishment, both from experimentally buried seeds and from the natural seed bank. Defecation recurrence had no effect on seedling establishment. We conclude that dung beetle activity affects seed bank dynamics with important consequences for seedling establishment in tropical forests.Abstract in Spanish is available with online material.     

Fire,canopy cover and seedling dynamics in Mediterranean shrubland of northeastern Spain

Abstract. Seedling establishment and survival of the main woody species of early-successional shrubland in northeastern Spain were studied from 1992 to 1995 with emphasis on the importance of vegetation cover (existence of open areas and the situation beneath the plant canopy) and microhabitat (occurrence of stones, litter and bare soil). In the absence of fire, vegetation cover (measured at a scale of 30cm × 30 cm) was not correlated with seedling emergence of most species, nor with growth and survival of seedlings up to one year old. Seedlings older than one year showed a similar pattern: their density was not significantly different in both kinds of habitats. The emergence of seedlings was mainly associated with the presence of mature plants – which can provide seeds – and with the absence of unsuitable microhabitats (large stones, deep litter and bare soil). In April 1994 a wildfire burned the study area. This enabled a study of the pattern of post-fire establishment. After the wildfire, seedling emergence of several species increased and most species produced seedlings with higher survival and growth rates than in the period before the fire. Pre-fire cover, however, did not show significant effects on post-fire seedling dynamics in most cases.     

Allometric relationships between seed mass and seedling characteristics reveal trade-offs for neotropical gap-dependent species

A seed size–seed number trade-off exists because smaller seeds are produced in greater number but have a lower probability of establishment. This reduced establishment success of smaller-seeded species may be determined by biophysical constraints imposed by scaling rules. Root and shoot diameter, root growth extension rate (R _GER) and shoot length at death for dark-grown seedlings are predicted to scale with the cube root of seed embryo and endosperm mass (m). We confirmed this expectation for ten neotropical gap-dependent tree species with an embryo and endosperm dry mass >1 mg. However, for nine smaller seeded species (m < 1 mg) with photoblastic germination, root and shoot diameters were larger than expected, and consequently, R _GER was slower than expected. The maximum shoot thrust of seedlings from seeds with masses ≥1 mg was comparable to the estimated force required to displace overlying litter, supporting the hypothesis that photoblastic behaviour only occurs in seeds with insufficient shoot thrust to displace overlying leaves. Using the model soil water, energy and transpiration to predict soil drying in small and large gaps, we showed that: (1) gaps that receive a significant amount of direct sunlight will dry more quickly than small gaps that do not, (2) compared to the wet-season, soil that is already dry at depth (i.e. the dry-season) will dry faster after rainfall (this drying would most likely kill seedlings from small seeds) and (3) even during the wet-season, dry periods of a few days in large gaps can kill shallow-rooted seedlings. We conclude that the smaller the seed, the more vulnerable its seedling would be to both covering by litter and soil drying because it can only emerge from shallow depths and has a slow R _GER. Consequently, we suggest that these allometrically related factors contribute to the reduced establishment success of smaller-seeded species that underpins the seed size–seed number trade-off.