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.     

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.     

浙江天童常绿阔叶林地被层的研究

 对天童常绿阔叶林地被层植物种类组成及群落学作用进行了初步研究。研究表明：组成地被层计有维管植物44科78属114种。蕨类植物优势度最大,里白占绝对优势。39种草木植物,以世界分布科和地面芽植物为主,常绿种类占84.6％,未见亚洲和北美洲间断分布的地理成分。乔灌木幼苗51种,80%的苗量集中于4～6个种中,优势种突出。藤本植物24种,草质藤本占25％,落叶种类42％。地面苔藓植物(22种)与枯枝落叶构成镶嵌分布,发育较成熟的群落中苔藓种类多,盖度大。里白及其枯死物形成层层交错重叠的地被层,阻碍凋落物进入地表,减弱光照,使近地表光照长期低于幼苗光补偿点,抑制了乔灌木优势种的更新。凋落物阻碍种子下移至半分解层,栲树种子下移率为17％至22.3％,冬季种子的动物损耗率平均为98.2％,凋落物中种子的平均腐烂率为28.2％,表层保存下来的栲树种子因缺水萌发速度慢且成活率低,下层萌发速度虽快,但腐烂率高,栲树天然更新幼苗主要来源于草窗。     

The role of seed depth,litter, and fire in the seedling establishment of amphicarpic peanutgrass (Amphicarpum purshii)

Summary Amphicarpum purshii is an annual grass which mostly grows in disturbed areas of the New Jersey Pine Barrens, USA. It is amphicarpic, producing spikelets (and seeds) both above and below the soil surface. Previous research has shown that subterranean seed production ensures reproduction in the event of a major disturbance such as fire and results in rapid post-burn colonization of these sandy habitats. The effects of fire, litter, and seed depth were further examined by planting subterranean seeds at four depths in 16 litter-covered flats buried at ground level and comparing plants arising from burned flats with those in undisturbed litter-covered flats. At 0 and 1 cm depth, rates of seedling emergence were lowest in burned flats. Surface-sown seeds produced seedlings more likely to desiccate. Sowing depth had a greater influence on most measured characters than burning treatments. The mean depth of subterranean seed placement by Amphicarpum is 3.5 cm and this coincides with the seed depth from which plants showed the greatest height growth, shoot biomass, and reproductive output. In a second experiment, subterranean seeds on the bare soil surface in clay pots were more likely to lose viability and less likely to germinate than seeds protected by litter or burial in soil. In addition to providing protection from fire, placement of seeds below ground in the sandy habitat of peanutgrass provides conditions more suitable for seed survival and subsequent seedling establishment.     

动物与植物种子更新的关系Ⅱ．动物对种子的捕食、扩散、贮藏及与幼苗建成的关系

植物的繁殖体总是面临来自各类生物（如昆虫、脊椎动物、真菌）的捕食风险。因动物捕食引起的种子死亡率影响植物的适合度、种群动态、群落结构和物种多样性的保持。种子被捕食的时间和强度成为植物生活史中发芽速度、地下种子库等特征的主要选择压力,而种子大小、生境类型等因素也影响动物对植物种子的捕食。捕食者饱和现象被认为是植物和种子捕食者之间的高度协同进化作用的结果,是限制动物破坏种子、提高被扩散种子存活率的一种选择压力。大部分群落中的大多数植物种子被动物扩散。种子扩散影响种子密度、种子被捕食率、病原体攻击率、种子与母树的距离、种子到达的生境类型以及建成的植株将与何种植物竞争,从而影响种子和幼苗的存活,最终影响母树及后代植物的适合度。种子被动物扩散后的分布一般遵循负指数分布曲线,大多数种子并没有扩散到离母树很远的地方。捕食风险、生境类型、植被盖度均影响动物对种子的扩散。植物结实的季节和果实损耗的过程也体现了其对扩散机会的适应。许多动物有贮藏植物种子的行为。动物贮藏植物繁殖体的行为,一方面调节食物的时空分布,提高了贮食动物在食物缺乏期的生存概率;另一方面也为种子萌发提供了适宜条件,促进了植物的扩散。于是,植物与贮食动物形成了一种协同进化关系,这种关系可能是自然界互惠关系（mutualism)的一种。影响幼苗存活和建成的因子包括种子贮蒇点的微生境、湿度、坡向、坡度、林冠盖度等。许多果食性动物吃掉果肉后,再将完好的种子反刍或排泄出来。种子经动物消化道处理后,发芽率常有所提高。     

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.     

A fungal endosymbiont affects host plant recruitment through seed- and litter-mediated mechanisms

1.  Many grass species are associated with maternally transmitted fungal endophytes. Increasing evidence shows that endophytes enhance host plant success under varied conditions, yet studies have rarely considered alternative mechanisms whereby these mutualistic symbionts may affect regeneration from seed.
2.  We performed a microcosm experiment to evaluate whether infection with Neotyphodium occultans affects recruitment in the annual grass Lolium multiflorum either directly, by infecting the seeds, or indirectly, by altering the suitability of recruitment microsites through the litter shed by host plants. Endophyte effects on establishment were tested for different litter depths and watering regimes under natural herbivory by leaf-cutting ants.
3.  Seed infection increased seedling emergence through the litter as well as final recruitment, irrespective of microsite conditions. However, litter produced by infected plants delayed emergence and decreased density of both infected and non-infected grass populations.
4.  Individual plant biomass did not change with seed infection but was increased under deep litter from endophyte-infected plants. Although seed infection did not protect establishing plants from leaf-cutting ants, herbivory was reduced in the presence of deep litter shed by infected plants.
5.  We conclude that fungal endophytes may affect host plant recruitment across subsequent generations not only by infecting the seeds but also through the host's dead remains. While the former effect entailed an advantage to infected plants, litter-mediated effects did not discriminate by infection status, and generally promoted the establishment of fewer and larger plants. Thus hidden foliar symbionts may play an underappreciated role in maintaining host species dominance through the litter produced by prior patch occupants.     

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.     

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.     

Response of ant communities and ant–seed interactions to bush regeneration

Summary Bush regeneration programmes aim to restore native plant communities by interventions designed to assist natural recovery, supplemented by planting or direct seeding where necessary. The present study compared ground-dwelling ant communities and rates of seed removal by ants in weed-infested, regenerating and undamaged urban bushland in northern Sydney. Three replicate dry sclerophyll sites on sandstone were studied, each containing zones of weed-infested, regenerating and undamaged bushland. Canopy cover, midstorey cover, ground cover and litter depth were measured at each site. Each zone type showed distinctive vegetation and cover characteristics, particularly at the ground level. Ground foraging ants were sampled with pitfall traps, sorted to morphospecies and placed in functional groups. There were no significant differences in ant abundance, number of genera or number of morphospecies between site types. Ant diversity and equitability was highest in the undamaged zones, followed by regenerating zones, then weedy zones. There was a significant difference between zone types in the proportion of ant species in different functional groups. Weedy zones were dominated by Opportunist species whereas regenerating zones had a high proportion of Dominant Dolichoderinae. At undamaged zones, the distribution of ants across functional groups was more even. Differences in the ant communities between the three zone types were reflected by differences in interactions between ants and seeds. Seed removal rates of both native and exotic seeds were significantly lower at weedy zones than at undamaged zones, with the regenerating zones having intermediate rates. Sites subject to regeneration 4–10 years previously were intermediate between undamaged and weedy zones for ground cover, litter depth, ant species diversity and equitability, proportion of Opportunist ant species, and seed removal rates. Thus, the regeneration techniques used have been at least partially successful in restoring some aspects of the invertebrate fauna and ecological interactions typical of undamaged dry sclerophyll bushland.     

The influence of spatial heterogeneity on regeneration by seed in a limestone grassland

Abstract. The spatial distribution of seedlings in fertilized and non-fertilized grassland patches (1 m x 2 m) was examined in alvar vegetation on the island of Öland, Sweden. We established whether variation in seedling distribution in grassland patches reflects the distribution of conspecific adults. We also established the significance of microsite turf attributes for prediction of seedling distribution at a small scale (10 cm x 10 cm), and the significance of conspecific adults abundance, assemblage of adult species and cover of lichens, mosses, litter and bare ground. Our results indicate that fertilization reduces the importance of regeneration by seed of perennial species and enhances seedling recruitment of winter annuals and biennials. At the patch scale, enrichment does not affect species richness. Community patchiness contributes to the maintenance of diversity by providing a variety of environments with differential regeneration of the component species. Cover of lichens, mosses, litter and bare ground, accounts, in part, for the spatial variation in seedling numbers of certain species. The strong association of seedlings of some species with conspecific adults indicates that limitations in dispersal and/or the occurrence of a more suitable environment for emergence and/or establishment close to conspecific adult plants are likely to constrain the spatial distribution of new individuals in many cases. For other species, the lack of association of seedling densities with the cover of their conspecific adults, suggests that long-distance dispersal or a long-lived seed bank can play a role in counteracting the effect of short-distance dispersal.     

花楸树种子散布、萌发与种群天然更新的关系

花楸树是我国东北林区重要的非木质资源树种,其实生天然更新不良.本文通过研究花楸树种子散布、土壤种子库及种子萌发出土过程,分析花楸树实生天然更新的影响因素.结果表明：自然散落的花楸树果实96.1%分布于母株2 m范围内,凋落物层和土壤表层（0～2 cm）的种子数占土壤种子库总数的97.0%;不同季节花楸树土壤种子库种子数量差别很大,当年11月上旬种子数量最多,达(257.7±69.2)粒·m^-2;翌年7月下旬种子数量最少,仅为(2.9±2.9)粒·m^-2;温度不是花楸树种子萌发出土过程的限制因子,0 ℃～5 ℃时幼苗出苗率达(67.5±6.6)%,但对其出苗速率影响显著.土壤含水量为50%时,花楸树出苗率最高,达(74.7±4.2)%;含水量为60%时,幼苗死亡率最低,为(32.6±0.6)%.花楸树种子的散布格局和土壤种子库的时空分布格局影响种子的萌发出土过程,进而影响其种群的天然更新.     

Efficiency of endozoochorous seed dispersal in six dry-fruited species (Cistaceae): from seed ingestion to early seedling establishment

We combined laboratory and nursery experiments to analyse the effectiveness of sheep as endozoochorous seed dispersers of six native shrubby Cistaceae species collected in SE Spain (Helianthemum apenninum (L.) Mill., H. violaceum (Cav.) Pers., Fumana ericoides (Cav.) Grand., F. thymifolia (L.) Spach, Cistus monspeliensis L. and C. laurifolius L.), considering the main stages after seed ingestion, i.e. seed recovery, seed germination, seedling emergence and early seedling establishment. Seed recovery after gut passage was high (around 40%) for all the species, except F. thymifolia (12%). Most seeds (ca. 90%) were recovered within 48 h after ingestion for all the species, although seeds were still recovered up to 96 h after ingestion. Gut passage increased germination up to seven-fold compared to non-ingested seeds. Furthermore, seedling emergence from seeds contained in pellets was overall similar (intact pellets) to or higher (crumbled pellets) than emergence from seeds without dung. Survival of emerged seedlings and mass of seedlings after 20 days were not reduced by dung. Sheep act therefore as effective dispersers of these Cistaceae species by scattering seeds and promoting germination, while faeces do not hamper seedling establishment. We conclude that the interaction between herbivorous ungulates and these dry-fruited species may be considered a mutualism qualitatively similar to the mutualism between frugivorous vertebrates and fleshy-fruited plants.     

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

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

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.     

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.     

Rodent-favored cache sites do not favor seedling establishment of shade-intolerant wild apricot (Prunus armeniaca Linn.) in northern China

Seed dispersal by scatter hoarders is widely assumed to involve directed dispersal, in which microhabitats selected for caching also benefit seedling establishment and growth. However, in many systems, this may not be true if animal-favored cache sites do not match the safe sites for plants, or if cache sites benefit the plant in one life stage and not another. Here, we investigated whether cache sites selected by rodents are favorable for seedling establishment and growth of shade-intolerant wild apricot (Prunus armeniaca Linn.) in northern China. We tracked tagged seeds and compared the germination and growth of seedlings from rodent-cached seeds with that of naturally established seedlings in a secondary forest and shrubland stand. Rodents preferred to cache seeds under shrubs with medium canopy cover (31–60 %) in litter substrate in the secondary forest, and under shrubs with high canopy cover (>60 %) in soil or litter substrate in the shrubland stand, neither of which conveyed an advantage for seedling establishment. Although fewer caches were made along shrub edges, or under low canopy cover (≤30 %) in the secondary forest, or along shrub edges, open areas of grass, or under low canopy cover in the shrubland stand, these cache sites consistently contributed to higher survival rates. The microhabitats of grass, soil, or low canopy cover significantly promoted the emergence, survival, and growth of naturally established seedlings. Our results are best explained by the conflicting demands of rodents for caching seeds in more secure sites and P. armeniaca’s high-light requirements for seedling recruitment. We argue that the relationship between favored cache sites for seed hoarders and safe sites for plants will often not match but may still allow a reasonable rate of establishment and regeneration.     

凋落物和土壤覆盖对动物取食和搬运辽东栎种子的影响

在六盘山区的华北落叶松人工林,研究了清除凋落物、凋落物覆盖和土壤覆盖(以不清除凋落物直接将种子投放于森林地表为对照)等处理对动物取食和搬运辽东栎种子的影响.结果表明:种子释放3d后,凋落物和土壤覆盖处理种子均具有较高的留存率(分别为10.7％和7.0％);释放14 d后,土壤覆盖处理种子的留存率仍最高(0.7％),但凋落物覆盖处理种子的留存率为0.凋落物和土壤覆盖处理种子的就地取食率很高(分别为45.9％和41.5％);清除凋落物处理种子的就地取食率最低(27.0％),但其被搬运后的取食率最高(49.8％);凋落物覆盖、清除凋落物和土壤覆盖处理种子被动物搬运后的埋藏率均显著高于对照(P＜0.01).种子被动物搬运后集中分布于5 m以内,尤其在＜1 m和l～2m两个距离组的分布频率更高;种子被搬运后取食的平均距离大干埋藏的平均距离,以土壤覆盖和凋落物覆盖处理最大,分别为2.38 m±0.55 m和1.44m±0.26 m.     

杉木凋落物覆盖对自身幼苗出土和早期生长的影响

凋落物可通过物理和化学作用显著影响幼苗出土和早期生长,进而影响天然更新.杉木是中国南方重要的造林树种,但存在着天然更新障碍,其原因可能是林下较厚的凋落物层阻碍了杉木幼苗出土和早期生长.本试验利用覆盖自然和塑料凋落物来研究凋落物对杉木幼苗出土和早期生长的影响,并检验其影响是物理作用还是化学作用.本试验设置2种凋落物类型(自然和塑料凋落物)和4个覆盖厚度(对照,0 g·m^-2;浅层,200 g·m^-2;中层,400 g·m^-2;深层,800 g·m^-2).结果表明: 与对照(0 g·m^-2)相比,浅层(200 g·m^-2)凋落物覆盖对出苗率有促进作用但不显著,深层(800 g·m^-2)凋落物覆盖对出苗率和存活率有显著抑制作用.随着凋落物覆盖厚度的增加,幼苗根长不断减小,而茎长逐渐增加.凋落物浅层覆盖下杉木幼苗的根生物量、叶生物量和总生物量均最大,深层覆盖下最小.幼苗的根冠比随着凋落物覆盖厚度的增加而不断减小.与对照相比,凋落物覆盖下幼苗光合与非光合组织生物量比均有所增加.相同覆盖厚度下,自然和塑料凋落物对杉木幼苗出土和早期生长的影响均无显著差异,表明短期内凋落物覆盖对幼苗出土和早期生长的影响主要是物理作用.随着凋落物覆盖厚度的增加,杉木幼苗出土和早期生长表现为先促进后抑制,且杉木幼苗为了穿过厚厚的凋落物层倾向于把资源分配给地上部分.本研究结果为凋落物是影响杉木幼苗建植和天然更新的一个重要生态因子提供了试验证据.