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.     

Establishment and growth of seedlings of Eucalyptus obliqua: Interactive effects of litter,water, and pathogens

We studied how leaf litter, water and fungal pathogens affect the establishment, survivorship, and growth of Eucalyptus obliqua using glasshouse and field experiments. In a glasshouse experiment, the presence of leaf litter of E. obliqua increased the establishment of seedlings when the seeds were placed beneath the litter, but dense litter layers reduced seedling emergence when seeds were placed on top. Results from a complementary experiment suggest that the effect of litter may be mediated by the high humidity microenvironment it creates, which may enhance imbibition of unburied seeds. In another glasshouse experiment, litter in combination with excessive water applications reduced seedling emergence and increased seedling mortality, but the addition of fungicide reverted these effects. Litter with lower water application or fungicide added increased seedling emergence. In all cases higher biomass in seedlings growing in litter treatments was associated with earlier emergence. In two recently burnt sites, the density of seedlings of E. obliqua was positively correlated to the presence of litter. Experimental addition of litter in the field reduced soil temperature during the summer, and together with watering, increased soil water content, and the water potential of established seedlings. However, neither treatment had any positive effect on their growth or survivorship. On the contrary, litter addition reduced the height and number of leaves, probably as a result of increased herbivory. We conclude that in Mediterranean-type systems litter may produce various effects on seedling establishment depending on their developmental stage, and the season.     

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.     

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.     

Negative and positive interactions among plants: effects of competitors and litter on seedling emergence and growth of forest and grassland species

Living plant neighbours, but also their dead aboveground remains (i.e. litter), may individually exert negative or positive effects on plant recruitment. Although living plants and litter co‐occur in most ecosystems, few studies have addressed their combined effects, and conclusions are ambivalent. Therefore, we examined the response in terms of seedling emergence and growth of herbaceous grassland and forest species to different litter types and amounts and the presence of competitors. We conducted a pot experiment testing the effects of litter type (grass, oak), litter amount (low, medium, high) and interspecific competition (presence or absence of four Festuca arundinacea individuals) on seedling emergence and biomass of four congeneric pairs of hemicryptophytes from two habitat types (woodland, grassland). Interactions between litter and competition were weak. Litter presence increased competitor biomass. It also had positive effects on seedling emergence at low litter amounts and negative effects at high litter amounts, while competition had no effect on seedling emergence. Seedling biomass was negatively affected by the presence of competitors, and this effect was stronger in combination with high amounts of litter. Litter affected seedling emergence while competition determined the biomass of the emerged individuals, both affecting early stages of seedling recruitment. High litter accumulation also reduced seedling biomass, but this effect seemed to be additive to competitor effects. This suggests that live and dead plant mass can affect species recruitment in natural systems, but the mechanisms by which they operate and their timing differ.     

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.     

格氏栲幼苗叶绿素荧光和非结构性碳水化合物积累对种子散布位置的响应

植物种子从母树掉落形成土壤种子库时,凋落物或土壤是其最初接触的物理环境,种子所处位置(种子在凋落物上层、土壤表层或凋落物下层)影响了幼苗天然更新进程。模拟格氏栲种子在凋落物上层(种子下层铺垫2和4 cm凋落物)、土壤表层(无凋落物)及凋落物下层(种子上层覆盖2、4、6和8 cm凋落物)等3种不同散布位置,探讨种子散布位置对幼苗叶绿素荧光特性、非结构性碳水化合物、比叶面积、叶干物质含量和养分含量的影响。结果表明:不同散布位置的幼苗单位面积的叶氮含量与可溶性糖、非结构性碳水化合物含量呈显著正相关,与比叶面积呈显著负相关。适宜凋落物覆盖(2和4 cm)的幼苗通过提高叶绿素相对含量、可溶性糖含量、非结构性碳水化合物含量、叶干物质含量和单位面积的叶氮含量和叶磷含量,降低比叶面积等的资源获取策略来实现自身快速生长需求。无凋落物和深层凋落物覆盖(6和8 cm)的幼苗采取高单位重量的叶氮含量和比叶面积,低叶干物质含量和非结构性碳水化合物含量的资源保守型策略以截获更多有效光资源,进而弥补深层凋落物带来的郁闭环境,降低幼苗因“碳饥饿”而死亡的几率。下层铺垫凋落物的幼苗通过在叶片储藏淀粉,降低叶片光合组织消耗能量(低PSⅡ最大光化学效率)等维持幼苗生长。熵值法综合分析表明,浅层凋落物覆盖(2 cm)对格氏栲幼苗生长的促进作用最为显著,未来可通过调节天然林凋落物层厚度以促进格氏栲幼苗生长与更新。     

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.     

种子散布位置对格氏栲幼苗根系形态及生长特征的影响

种子从母株掉落于地面萌发后,其根系在不同散布位置(凋落物上层、土壤表层和凋落物下层)的生长形态影响幼苗定居及建成,而目前对其根系形态及生长特征的了解并不充分,限制了对幼苗根系在不同散布下适应策略的理解。为此,以格氏栲(Castanopsis kawakamii)为研究对象,通过模拟种子在凋落物中位置,设置凋落物上层(种子下层铺垫2 cm和4 cm凋落物,U2和U4处理)、土壤表层与凋落物下层(种子上层覆盖0、2、4、6 cm和8 cm凋落物,CK、D2、D4、D6和D8处理)等3种散布,探讨不同散布位置对格氏栲幼苗根系9个生长指标的影响。结果表明：(1)种子散布位置对幼苗根干物质质量具有显著影响,D2处理达最大值。(2)D2处理的幼苗根长、根表面积、根尖数、分枝数和比根长高于其它处理;根系平均直径在D6处理达最大值。(3)相关分析表明根长、根表面积、根尖数、分枝数和比根长与根系平均直径呈显著负相关关系。(4)对根系9个生长指标提取主成分后聚类为4个类群,D2与D4处理各划分一类;U2与U4处理划分一类,其余三个处理划分一类。综上所述,凋落物浅层覆盖(D2处理)适宜格氏栲根系生长;凋落物...     

Effect of grass litter on seedling recruitment of the critically endangered Cistus heterophyllus in Spain

Cistus heterophyllus Desf. is an Iberian-north African endemic plant species with only one wild population left in Spain. A sowing experiment under controlled conditions was designed to assess both the seedling emergence rate and the growth fitness of C. heterophyllus on a litter layer of Brachypodium retusum (Pers.) Beau., which forms dense carpets around the remnant population of C. heterophyllus. In order to test the physical effects of B. retusum litter on Cistus seedling recruitment, six sowing treatments in pots were compared that combined three doses of litter (0, 0.6 and 1.1 kg m⁻²) and different physical locations of the seeds (above, below the litter layer or within a litter–peat mixture). B. retusum litter interfered with the seedling recruitment of C. heterophyllus, negatively affecting both seedling emergence and their early growth. Such an effect was more severe when the seeds were located above a 0.6 kg m⁻² litter layer, although seedlings from seeds sown under the litter showed a growth reduction ranging from 58% to 100% compared to the no-litter treatment. A significant negative effect of the litter–peat mixture on the number of leaves, leaf size and number of branches of seedlings was also inferred. These data provide experimental evidence on the serious threat that B. retusum litter represents for the conservation and regeneration of the last population of C. heterophyllus in Spain.     

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.     

Leaf litter of <Emphasis Type="Italic">Kageneckia angustifolia</Emphasis> D. Don (Rosaceae) inhibits seed germination in sclerophyllous montane woodlands of central Chile

Leaf litter accumulation can have either positive, negative or neutral effects on seed germination and seedling recruitment. In montane woodlands of the Mediterranean zone of central Chile, large amounts of leaf litter accumulate beneath the crowns of the summer semi-deciduous tree Kageneckia angustifolia and no regeneration of this or other plant species has been observed beneath this tree throughout the year. In a sample plot of 5000 m² we selected ten K. angustifolia trees and measured (1) leaf litter accumulation beneath and outside canopy; (2) the effects of time elapsed since burial on viability of K. angustifolia seeds with and without a leaf litter cover; (3) field seed germination with presence or absence of leaf litter and (4) the possible chemical effects of K. angustifolia leaf litter leachates on seed germination of its own seeds and of other two co-occurring native shrubs species (Guindilia trinervis and Solanum ligustrinum). Our results show that a considerable accumulation of leaf litter occurred beneath K. angustifolia, and litter negatively affected seed viability and germination of this species in the field. Under laboratory conditions, K. angustifolia leaf litter leachates inhibited seed germination of its own seeds and of the two native shrub species. Chemical effects are likely involved in the negative effects of leaf litter on the recruitment of K. angustifolia in the montane sclerophyllous woodland of central Chile.     

Effects of leaf litter on woody seedlings in xeric successional communities

We investigated the effect of leaf litter on the establishment of Eucalyptus incrassata, a mallee eucalypt. It has been suggested that litter accumulation may hinder seedling establishment, and that the removal of litter may be one of the mechanisms through which fire enhances recruitment. We conducted factorial experiments testing the effects of three kinds of leaf litter on E. incrassata seeds and seedlings at three contiguous sites with different land use histories. One site was an uncleared E. incrassata open mallee woodland (Mallee site), one a cleared area that had been ungrazed for about five years (Pasture site) and the third an area of mallee rolled some 40 years ago and permitted to regenerate (Regrowth site). Litter had no effect on emergence of planted E. incrassata seeds, but emergence differed between sites. Overall, the percentage of seeds that germinated and emerged was substantial (mean 35.2% ± 25.9%). Seedling shoot biomass did not differ between sites or litter treatments. Although seedlings grown in Pasture litter suffered higher mortality rates, overall mortality rates were low (mean 13.2% ± 15.5%), suggesting that leaf litter has little effect on recruitment rates during winter and spring. We conclude that leaf litter does not affect emergence or growth in young E. incrassata seedlings during winter and spring, when most establishment occurs. Our results emphasize the difficulty in predicting litter effects on recruitment.     

Effects of dung beetles on seedling emergence from herbaceous seeds in the dung of sika deer (<Emphasis Type="Italic">Cervus nippon</Emphasis>) in a temperate Japanese grassland ecosystem

The effects of dung form and condition and of dung beetles on the emergence of seedlings from herbaceous seeds in sika deer dung were examined in a temperate grassland ecosystem dominated by Zoysia japonica and Hydrocotyle maritima. I conducted field experiments to compare seedling emergence between dung exposed to dung beetles and intact dung using both dung pellets and pats during a typical rainy month (June) and the hottest, drier month (August), when large numbers of seeds of the dominant species were present in the dung. The exposed dung was immediately attacked and broken up by dung beetles, whereas dung protected from the beetles remained intact. In June, at least 12 herbaceous species, including Z. japonica, H. maritima, Mazus pumilus, and Plantago asiatica, emerged from the dung, versus at least six species in August. Decomposition rates of the pellets in June and decomposition scores of the pats in June and August were positively correlated with the number of emerging seedlings, suggesting that the acceleration of decomposition by dung beetles can positively affect seed germination. In this system of interactions among sika deer, herbaceous plants, and dung beetles, sika deer dung prevented seeds from germinating, and beetles had an indirect positive effect on seedling emergence by accelerating decomposition of the dung, although the extent of the effect may depend on the dung type, plant species, and environmental factors.     

Rodent seed predation and seedling recruitment in mesic grassland

Seedling recruitment of two grasses (Arrhenatherum elatius and Festuca rubra) and two herbs (Centaurea nigra and Rumex acetosa) was measured in areas with and without rodents to which seeds of each species were sown at three seed densities (1000, 10,000 and 50,000 seeds m⁻²) in two seasons (spring and autumn 1995). Seed removal was measured for 10-day periods and the fate of seedlings was followed for 15 months after sowing. The proportion of seed removed ranged from 6 to 85% and increased with increasing seed density for each species. Rodents had no effect on seedling emergence or survival in the spring sowing. In the autumn sowing, rodents reduced seedling emergence of all four species sown at 1000 and 10,000 seeds m⁻² but had no impact at 50,000 seeds m⁻², presumably because of microsite limitation. We suggest the difference between spring and autumn arose because emergence was seed limited in autumn but microsite limited in spring; microsite availability was higher in autumn because a summer drought killed plants, reduced plant biomass and opened up the sward. Fifteen months after the autumn sowing, fewer A. elatius and C. nigra seedlings survived on plots exposed to rodents. This result reflected not only the reduced seedling emergence but also increased seedling mortality (seedling herbivory) in sites exposed to rodents. In contrast, F. rubra and R.acteosa showed density-dependent seedling survival which compensated for initial differences in seedling emergence, so that no effect of rodents remained after 15 months. The results suggest that rodent seed predation and seedling herbivory exert strong effects on seedling recruitment of A.elatius and C. nigra when recruitment conditions are favourable (conditions that lead to high microsite availability) and may contribute to both species being maintained at low densities in the grassland. The results also demonstrate that highly significant impacts of rodent seed predation at the seedling emergence stage can disappear by the time of plant maturation. Received: 2 March 1998 / Accepted: 28 September 1998     

Lack of recruitment in Lavandula stoechas subsp. pedunculata: a case of safe-site limitation

Lavandula stoechas subsp. pedunculata regeneration depends exclusively on the establishment of new individuals. Seed availability and seedling emergence and survival are therefore critical life stages and processes for species regeneration. In this study, seedling emergence and survival was monitored for two years in the scrub, both in clearings and adjacent to adult plants, and the surrounding perennial grassland, at 1, 3 and 5 m from the scrub. Soil seed bank spatial distribution was also studied for one year in the same two habitats, using the same sampling design. Soil seed availability in the scrub is high regardless of the distance from the adult individuals. On the contrary, the adjacent grassland shows a drastic fall in seed density, and almost no seedlings were observed there. In the scrub, seedling density was negatively related to distance from the three nearest adult plants in the clearings, and positively related to adult plant size beneath the adult Lavandula plants. There was also a negative relationship between seedling density and the percentage of bare soil. Only one seedling survived the first drought period, with no detection of effects of either position with respect to adult individuals or seedling density. We hypothesized that the study populations suffer a lack of appropriate safe sites within the scrubland while in the adjacent perennial grassland, observed low seed availability was added to safe-site limitation. That results in a lack of successful seedling establishments and a poor expansion potential of Lavandula scrublands, whose edges remain static in the short and medium term. As found in other Mediterranean scrubland, recruitment may only occur in years with particularly favourable weather, under disturbance regimes that increase seedling survival probability or when external dispersal agents increased seed availability in adequate places for Lavandula establishment.     

Amplified drought induced by climate change reduces seedling emergence and increases seedling mortality for two Mediterranean perennial herbs

Seedling recruitment is a bottleneck for population dynamics and range shift. The vital rates linked to recruitment by seed are impacted by amplified drought induced by climate change. In the Mediterranean region, autumn and winter seedling emergence and mortality may have strong impact on the overall seedling recruitment. However, studies focusing on the temporal dynamic of recruitment during these seasons are rare. This study was performed in a deciduous Mediterranean oak forest located in southern France and quantifies the impact of amplified drought conditions on autumn and winter seedling emergence and seedling mortality rates of two herbaceous plant species with meso‐Mediterranean and supra‐Mediterranean distribution (respectively, Silene italica and Silene nutans). Seedlings were followed from October 2019 to May 2020 in both undisturbed and disturbed plots where the litter and the aboveground biomass have been removed to create open microsites. Amplified drought conditions reduced seedling emergence and increased seedling mortality for both Silene species but these negative effects were dependent on soil disturbance conditions. Emergence of S. italica decreased only in undisturbed plots (−7%) whereas emergence of S. nutans decreased only in disturbed plots (−10%) under amplified drought conditions. The seedling mortality rate of S. italica was 51% higher under amplified drought conditions in undisturbed plots while that of S. nutans was 38% higher in disturbed plots. Aridification due to lower precipitation in the Mediterranean region will negatively impact the seedling recruitment of these two Silene species. Climate change effects on early vital rates may likely have major negative impacts on the overall population dynamic.     

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.     

Contrasting litter effects on old field tree germination and emergence

Because the fate of seeds is critical to understanding the invasion of old fields by trees, and plant litter is an important component defining the old field microsite of dispersed seeds, I investigated the effects of litter type (Solidago spp./goldenrod,Quercus spp./oak, mixed) and litter amount (100–800 g/m²) on tree seed germination and seedling emergence. I found that at all densities bothSolidago andQuercus litter greatly reduced emergence of the small-seeded, bird-dispersed species,Juniperus virginiana andCornus florida. For one of the large-seeded, mammal-dispersed species,Carya tomentosa, high densitySolidago litter and high density mixed litter treatments reduced emergence. For the other large-seeded species,Quercus rubra, the high density mixed litter treatment and all levels ofSolidago litter reduced emergence.Quercus seedlings emerged twice as often as the other three species in control pots without litter.Carya emerged before the other species but the high density oak treatment delayed the expansion of its cotyledons. My results suggest that litter may contribute to the slow rate of tree invasion and the low probability of tree establishment in old fields. However, old field litter studies taken together point to the difficulty in drawing general conclusions about any net effect of litter on old field tree establishment.     

Alleviation of Summer Drought Boosts Establishment Success of <Emphasis Type="Italic">Pinus sylvestris</Emphasis> in a Mediterranean Mountain: An Experimental Approach

We performed an irrigation experiment to study the impact of summer drought on Pinus sylvestris establishment at its southernmost distribution limit. Watering was done during the first growing season simulating mesic summer conditions, and we monitored the consequences for survival and growth during the first growing season and the delayed consequences on the second growing season. In addition, we considered the heterogeneity created by the microhabitats, where seeds are found after dispersal (bare soil, under shrubs, and under adult pines). Summer drought was the main mortality factor in all the microhabitats. Watering increased emergence and doubled seedling survival compared to non-watered control sites. Differences were even higher when the cumulative effect on emergence and survival was considered, with an overall recruitment of 22.4% in watered plots vs. 7.9% in control. Irrigation increased growth in bare soil and under shrubs, but had scant effect on growth under pines, suggesting that radiation was the limiting factor in this microhabitat. The positive effect of irrigation on growth parameters persisted during the second growing season despite water was not added the second year, showing delayed consequences of drought on seedling performance. Summer drought thus limits Pinus sylvestris establishment in these southernmost forests by reducing both recruitment and growth. This might lead to the development of a remnant dynamic in these relict populations under the current regional increase in dryness and rainfall variability associated with global warming.