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 litter on seedling establishment: an indoor experiment with short-lived Brassicaceae species

Recruitment by seeds is essential both in vegetation dynamics and in supporting biodiversity in grasslands. The recruitment by seeds is feasible in suitable vegetation gaps from the seed rain and/or by establishment from persistent soil seed banks. Cessation of grassland management results in litter accumulation, which leads to the decline of species diversity by the decreased availability of open patches. Low amounts of litter is often beneficial, while high amounts of litter is detrimental for seed germination and seedling establishment of short-lived species. In a designed indoor experiment, we explored the effect of litter on seedling establishment by germinating six short-lived Brassicaceae species with both increasing seed mass and litter cover. We found that both seed mass and litter had significant effect on germination and establishment of the sown species. Small-seeded species were significantly negatively affected by the 300 and/or 600 g/m² litter layers. No negative litter effect was detected for species with high seed masses (Lepidium spp.). No overall significant positive litter effect was found, although for most of the species cumulative seedling numbers were not the highest at the bare soil pots. Our results suggest that the negative effects of litter are less feasible on the large-seeded short-lived species than on that of small-seeded ones.     

Experiments on the mechanism of tree and shrub establishment in temperate grassy woodlands: Seedling emergence

Field experiments were designed to examine tree and shrub seedling emergence in temperate grassy woodlands on the New England Tablelands. The effects of study sites, intensity of previous grazing, removal of ground cover by fire or clearing, burial of seeds and ant seed theft on seedling emergence were tested in two field experiments. Six tree and seven shrub species were used in the experiments and their cumulative emergence was compared with laboratory germination studies. All species used in field experiments had lower cumulative emergence than those in laboratory germination studies despite prolonged periods of above average rainfall before and after seeds were sown. Eucalypt species emerged faster in the field than the shrub species and generally attained higher cumulative emergence than the shrubs. Spatial effects of sites and patches within sites, and of previous grazing history did not strongly influence patterns of seedling emergence in most species. Ground and litter cover generally did not enhance or suppress the emergence of seedlings, although the removal of cover in recently grazed areas enhanced the emergence of some species. Burning enhanced the emergence of some tree and shrub species where plots had more fuel and intense fires, but this effect was not strong. Compared with other treatments, seedbed manipulations produced the strongest effects. In the absence of both invertebrate and vertebrate predators, seedling emergence was lower for surface‐sown seed, compared with seed sown on scarified soil surfaces. Higher seedling emergence of buried seeds in the presence of invertebrate predators probably resulted from the combined effects of predator escape and enhanced moisture status of the germination environment. Some promotion of emergence was achieved for all species in most sown treatments probably as a result of a prolonged above average rainfall. In contrast, the natural recruitment of trees and shrubs was negligible in experimental plots, highlighting the importance of seed supply and dispersal as ultimate determinants of recruitment.     

Germination and emergence of annual species and burial depth: Implications for restoration ecology

Due to the high content of viable seeds, topsoil is usually spread on ground left bare during railway and motorway construction to facilitate the regeneration of vegetation cover. However, during handling of the topsoil, seeds are often buried deeply and they cannot germinate or the seedlings cannot emerge from depth. This study experimentally explores the predictive value of seed mass for seed germination, mortality and seedling emergence at different burial depths for 13 common annual species in semiarid Mediterranean environments. We separate the effect of burial depth on germination and emergence by means of two experiments. In the germination experiment, five replicates of 20 seeds for each species were buried at depths ranging from 0 to 4 cm under greenhouse conditions. Germinated and empty or rotten seeds were counted after 8 weeks. In the emergence experiment, five replicates of four newly-germinated seeds per species were buried at the same depths under controlled conditions and emergence was recorded after 3 weeks. The effect of burial depth on percentage of germination and seedling emergence was dependent on seed size. Although all species showed a decrease in germination with burial depth, this decrease was greater for small-than large-seeded species. Percentage of emergence was positively related to seed mass but negatively related to burial depth. Seed mortality was higher for small-than large-seeded species, but there was no general effect of burial depth on this variable. Thus, the current practice of spreading 30 cm deep layers of topsoil in post-construction restoration projects is unadvisable. In this restoration scenario, thinner layers of topsoil should be used to achieve the maximum potential of the topsoil for germination and seedling establishment.     

Exotic Ecosystem Engineers Change the Emergence of Plants from the Seed Bank of a Deciduous Forest

The anthropogenic spread of exotic ecosystem engineers profoundly impacts native ecosystems. Exotic earthworms were shown to alter plant community composition of the understory of deciduous forests previously devoid of earthworms. We investigated the effect of two exotic earthworm species (Lumbricus terrestris L. and Octolasion tyrtaeum Savigny) belonging to different ecological groups (anecic and endogeic) on the emergence of plants from the seed bank of a northern North American deciduous forest using the seedling emergence method. We hypothesized that (1) exotic earthworms change the seedling emergence from the plant seed bank, (2) L. terrestris increases the emergence of plant seedlings of the deeper soil layer but decreases that of the upper soil layer due to plant seed burial, and (3) O. tyrtaeum decreases plant seedling emergence due the damage of plant seeds. Indeed, exotic earthworms altered the emergence of plant seedlings from the seed bank and the functional composition of the established plant seedlings. Surprisingly, although L. terrestris only marginally affected seedling emergence, O. tyrtaeum changed the emergence of native plant species from the seed bank considerably. In particular, the number of emerging grass and herb seedlings were increased in the presence of O. tyrtaeum in both soil layers. Moreover, the impacts of earthworms depended on the identity of plant functional groups; herb species benefited, whereas legumes suffered from the presence of exotic earthworms. The results highlight the strong effect of invasive belowground ecosystem engineers on aboveground ecosystem characteristics and suggest fundamental changes of ecosystems by human-spread earthworm species.     

Bamboo as bioresource in Ethiopia: management strategy to improve seedling performance (Oxytenanthera abyssinica)

Seedling emergence and subsequent survival and growth are vital for natural forest restoration or plantation establishment by means of seeds. Such information is lacking for the African bamboo species. Two experiments were carried out in a greenhouse to evaluate the influence of seed orientation and sowing depth of the lowland bamboo Oxytenanthera abyssinica on seedling emergence, survival and growth. A randomised complete block design was used. Seedling emergence in the seed orientation experiment followed the order embryo-end-up>lay-flat>embryo-end-down. Survival rate after 62 days decreased in the order lay-flat>embryo-end-down>embryo-end-up. Mean seedling height and number of leaves per seedling followed a similar pattern. Seeds sown on top of the soil surface and at 2.5 mm depth achieved faster and higher seedling emergence than those sown at 5 and 10 mm depths. However, mean seedling height and number of leaves per seedling were higher in 5 and 2.5 mm depths than surface and 10 mm depths. There were significant quadratic relationships between sowing depth and seedling height (p=0.034) as well as number of leaves per seedling (p=0.032), both peaking around 5 mm soil depth. Lay-flat orientation, which is the most frequent position in broadcast sowing, is recommended at 5 mm sowing depth for the lowland bamboo based on overall performance in seedling emergence, survival and growth.     

Seed germination in Chrysophyllum sp. nov., a large-seeded rainforest species in north Queensland: effects of seed size,litter depth and seed position

Leaf litter affects seed germination in many ways and past studies have shown greater impacts on relatively small seeds, both within and among species. In this shade-house experiment I examined the impact of forest litter on seed germination in Chrysophyllum sp. nov. (Sapotaceae), a large-seeded (2.4 g) rainforest tree from north Queensland. Seed mass varies more than 30-fold in this species, making it useful for studying the role of litter as a possible selective pressure in the evolution of seed size in large-seeded species. Seeds of varying size (small, medium, large) were sown in planting boxes containing one of three litter levels (low, medium, high) and placed either below or on top of the litter. Seed size and litter biomass had no significant impact on the number of germinating seeds or the time to germination but seeds placed below the litter germinated around twice as frequently, and 20% sooner, than seeds placed on top of the litter. There were no significant interactions between any of the three factors. This shade-house experiment suggests that leaf litter is not an important selective pressure in the evolution of seed size in this species. However, if litter disturbance under field conditions differentially affects the probability of seed germination in relation to seed size and/or litter biomass, then litter could still act as a selective pressure in the evolution of seed size in Chrysophyllum and other large-seeded species.     

The combined effects of a newly designed dibber drill, irrigation and seed covering treatments on lettuce and calabrese establishment

A comparison was made between seedling establishment following sowing with the Mk2 dibber drill and a conventional coulter drill at three sowing occasions with both lettuce and calabrese. Different seed covering materials used to cover dibber-drilled seed were also compared. With lettuce, the rate and uniformity of seedling emergence and the numbers of seedlings emerging were greater following sowing with the dibber drill than with the coulter drill. Seedling establishment was further improved with irrigation especially when seeds were sown with a small quantity of the hydrophillic polymer, Alcosorb, which avoided associated problems of increased soil impedance. Alcosorb was a more effective seed cover than a peat and vermiculite mix and better suited for mechanical metering. Results indicate that the dibber drill can be considered as a realistic and cheaper alternative to transplanting for establishing later-season lettuce crops. With calabrese, however, percentage seedling emergence on all dibber-drilled plots was lower than that on plots sown with the coulter drill. This result may have been due to seed-coat damage caused as seeds are plunged into the soil during sowing with the dibber drill.     

Seed–litter–position drives seedling establishment in grassland species under recurrent drought

Changes in land use and climate interfere with grassland ecosystem processes. Here I experimentally investigated the combined effects of land‐use change related litter cover and contrasting water supply on seedling emergence. In this context, the role of the initial relative position of seeds, i.e. seeds on top of the litter versus seeds beneath the litter in interaction with water supply has not been investigated so far. I hypothesised that facilitative effects of litter on seedling emergence occur when seeds are covered by litter and deteriorate when litter covers the ground and seeds fall on it (seeds on top of the litter). Further, I hypothesised that the importance of seed position for seedling emergence will increase under conditions of recurrent drought. I performed a controlled pot experiment on seedling emergence of three common European grassland species (Pimpinella saxifraga, Leontodon autumnalis, Sanguisorba officinalis) by experimental manipulations of litter and water availability. Seedling emergence under moist conditions showed no significant differences between each litter position compared to the control across species. In contrast, under recurrent drought, seedling emergence was significantly higher below the litter compared to seeds on top of the litter and the control (i.e. no litter). In abandoned land, seedling emergence may be limited when seeds fall on ground‐covering litter. In contrast, in grasslands with regular low‐intensity land use, seedling emergence may be enhanced when a moderate level of litter covers seeds at the end of the growing season. Protective mechanisms that occur with seeds positioned beneath litter are particularly important under recurrent drought.     

Seed positioning in extruded pellets: does it matter?

Extruded pellets containing activated carbon (AC) can be used to sow native seeds while simultaneously applying herbicide to control invasive species. Incorporating AC in pellets has been demonstrated to protect native seeds; however, there may be unintended detrimental impacts to seedling emergence. We aimed to optimize seed position within pellets to maximize emergence and survival of the perennial shrub Jacksonia furcellata. Seeds were positioned at 2 mm (top), 6 mm (middle), and 12 mm (bottom) within pellets (with or without AC), sown on or below the soil surface, and compared to non-pelleted seeds sown under the soil surface in the equivalent positions (2, 6, and 12 mm depth). Trays were treated with a pre-emergent herbicide (Simazine) or left unsprayed. Emergence (without herbicide) was significantly higher from seeds positioned at the bottom of pellets without AC sown on the soil surface (70%), compared to non-pelleted seeds sown at the bottom (12 mm below the soil surface; 57%). However, emergence was inhibited when seeds were positioned in the middle (6 mm) of pellets with AC (32%). When treated with Simazine, survival was highest from seeds positioned at the bottom of pellets with AC (60%), compared to pellets without AC (15%) and non-pelleted seeds sown at the bottom (12 mm below the soil surface; 15%). Jacksonia furcellata seeds positioned at the bottom of pellets, sown on the soil surface, shows promise to minimize negative impacts to emergence, and maximize herbicide protection. Further testing with additional species is required to refine pellet production (e.g. recipe, extrusion, and shape) for optimal emergence.     

Non-native earthworms alter the assembly of a meadow plant community

Non-native earthworms can alter ecosystems by modifying soil structure, depredating seeds and seedlings, and consuming soil organic matter, yet the initial responses of plant communities to earthworm invasions remain poorly understood. We assessed the effect of non-native earthworms on seedling survival during germination and after establishment using six native and six non-native plant species grown from seed in single- and multi-species experimental mesocosms. We examined the extent to which earthworms (1) influenced seedling survival, (2) selectively depredated native versus non-native plants, (3) impacted establishment based on seed size and/or root morphology, and (4) shaped community assembly. The effect of earthworms on seedling survival varied temporally and among species but inconsistently with respect to species origin. Differences in seed/seedling survival translated to changes in community assembly. Earthworms tended to reduce species abundance, richness, evenness, and diversity in multi-species mesocosms and led to the divergence of communities by treatment. In general, species with large seeds and fibrous roots dominated communities with earthworms present, whereas species with small seeds and taproots only persisted in multi-species mesocosms without earthworms. Our findings suggest that earthworms act as ecological filters in the early stages of invasion to shape community composition based on plant morphological traits.

     

Recruitment filtering by a moss layer disadvantages large-seeded grassland species

Calcareous grasslands are rich in biodiversity and thus receive much attention in nature conservation. In such grasslands, the formation of moss layers is perceived as a management problem. However, its impacts on the community level are complex, as not only inhibition but also facilitation of vascular plant recruitment occur. Possible filters of recruitment are shading by mosses, isolation from soil resources and the resulting desiccation. To understand how seed size and shape moderate moss effects, a combined glasshouse and field experiment was conducted in southern Germany. Seeds of 14 species from calcareous grasslands were sown either on top or underneath a moss layer, or on bare soil without moss. We determined the total number of emerged and established seedlings, i.e. the ones that survived until the end of the experiment. Both measures were reduced for most species sown on moss, while mortality was slightly enhanced by moss presence. Seed size explained a significant proportion of the moss effects on plant recruitment. Inhibitive effects on recruitment increased with seed size when seeds were placed on top of moss. When germinating underneath moss, the effect on emergence changed from negative in small-seeded plant species to positive in large-seeded species, but this effect was insignificant for establishment. The positive response in large seeds was probably due to their higher moisture requirements for imbibition, and lower dependency on light for germination. However, moss-seed-interactions were not affected by seed shape. Seedling establishment of plants with large seeds strongly depended on their initial position in moss, while small-seeded species showed no effect. Hence, moss layers constitute a selective recruitment filter by modifying seed penetration and hence germination conditions. This effect could increase beta diversity of vascular plant species in calcareous grasslands, and thus conservation should aim at intermediate and patchy moss abundance.     

Seed and microsite limitation in Rheum nobile,a rare and endemic plant from the subnival zone of Sino-Himalaya

Background: Species persistence, particularly in monocarpic species, depends on the successful recruitment of individuals. An understanding of the factors that limit the recruitment of rare monocarpic plant species is therefore vital for their conservation.

Aims: To identify the factors limiting the recruitment of Rheum nobile, a rare and highly specialised monocarpic giant herb endemic to the high eastern Himalayas.

Methods: Seed sowing (seeds added or not added) and seedling transplanting experiments were conducted in disturbed (vegetation removed) and undisturbed plots in the vicinity of established populations of R. nobile to explore the mechanisms of recruitment limitation. Four levels of photosynthetically active radiation (0, 15, 30 and 50 μmol m^?2 s^?1) and two sowing positions (beneath and above grass litter or moss layer) were manipulated in the laboratory to determine how ground cover limited seedling emergence.

Results: Seed addition increased seedling recruitment. Disturbance significantly increased seedling emergence and establishment. Seed germination significantly decreased with the reduction of light availability, but 31.7% of all seeds germinated in complete darkness. Seedling emergence was close to zero when seeds were sown on top of a layer of grass litter or moss, but rose to 34.5% when the seeds were sown beneath such layers.

Conclusions: Our results indicate that the recruitment of R. nobile is limited by a combination of seed and microsite availability. Therefore, in order to conserve this species, we suggest adding seeds to suitable sites and implementing soil disturbances in existing populations to create suitable microsites.     

Vertical seed distribution in the soil constrains regeneration of Bromus pictus in a Patagonian steppe

Question: What is the effect of vertical seed distribution (i.e. seeds on soil surface or buried) and the relative importance of vertical and horizontal seed distribution (i.e. seeds near established vegetation or near bare soil) for seedling emergence? Location: Patagonian arid steppe. Methods: To evaluate the effect of vertical seed distribution on seedling emergence, four sowing treatments were used in field and greenhouse experiments. Vertical vs. horizontal seed distribution and the occurrence of seedlings were analysed in the field. Effects of trampling and mechanical drilling on seedling emergence were investigated experimentally in the field. Results: In field and glasshouse experiments buried seeds showed a higher emergence and larger seedling size than seeds lying on the soil surface. The observational field study indicated that natural emergence was not associated with particular microsites or plant cover (horizontal distribution), but that most seedlings emerged from buried positions (vertical distribution). Buried seeds represented less than 10% of the sampled seed cohort but account for almost 60% of the seedlings. The trampling/drilling experiment showed that sheep trampling and mechanical drilling increased seedling emergence compared to non‐treated controls. Conclusions: The population dynamics of Bromus pictus is strongly constrained by the vertical position of seeds, which largely affects emergence, and seed horizontal distribution, which largely affects seedling survival. Our results showed that sheep trampling after seed dispersal may be a low‐input technique for increasing grass recruitment.     

死、活地被物对红松伴生树种天然更新影响的实验研究

1963年在小兴安岭丰林自然保护区红松林及有关的皆伐迹地上设置了61组,325个实验样方,定期观察到1965年,以研究死、活地被物对人工播种的红松林伴生树种种子发芽和幼苗初期存活、生长的影响。实验表明死、活地被物,特别是死地被物的干湿变化剧烈,影响幼苗的成活,甚至使种子或幼根无法接触土壤,因而种粒越小,或轻而带翅,受影响越大。此外本文还指出了桦树种子的强迫休眠和延迟发芽现象。讨论了实验结果在红松林演替和人工促进天然更新中的意义。     

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.     

The chemical nature of P accumulation in agricultural soils��implications for fertiliser management and design: an Australian perspective

Sand burial, persistent seed bank and soil water content (SWC) are three factors that potentially can affect regeneration in sand dune plant populations. To evaluate the effects of these three factors on population regeneration of Eremosparton songoricum, a rare and endangered legume, we investigated seed germination, seedling emergence and seedling survival in greenhouse and controlled field experiments in different sand dunes microsites. Freshly matured seeds are physically dormant, and the highest germination was only 9.3?±?5.8% at 25/10°C. Seed germination occurred at burial depths from 0 to 10 cm, but the maximum depth from which seedlings emerged was 6 cm; from 1 to 6 cm, the deeper the burial, the lower the percentage of seedling emergence. Only 36.2% of the total soil seed banks occurred at depths of 0?C6 cm. For artificially sown seeds at different dune microsites, mean seedling emergence percentage was 6.8%. Of 150 seedlings that emerged in the field investigation at the study site, only those germinating in flat sandy areas survived, and mean survival percentage was only 2.0%. Thus, the proportion of non-dormant seeds in soil seed banks that developed into seedlings and survived to the end of the growing season was only 0.2%. Minimum SWC for seed germination, seedling emergence and seedling survival was 2.0%. During monitoring of emergent seedlings in the field, low seedling recruitment was at least partly due to the rate of root extension (1.6?±?0.3 cm day^?1) into the sandy soil, which was slower than that of the downward movement of plant-available moisture (2.8?±?0.6 cm day^?1). Thus, population regeneration under natural conditions rarely occurred via sexual reproduction, owing to the limited water resources available for seedling establishment. Rational field seeding practices, including manually scarified or dry stored seeds before sowing, sowing the seeds at right time and suitable place, are suggested for ecological restoration of endangered E. songoricum populations.     

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.     

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 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.