Importance of microhabitat and acorn burial on Quercus ilex early recruitment: non-additive effects on multiple demographic processes

This study investigates the effect of microhabitat and seed burial on the main demographic processes operating during the early recruitment of Quercus ilex, such as postdispersal seed predation, seed germination, and seedling emergence, survival and growth. The effect of burial was positive over all the processes analysed in this study, since predation rate was lower (63.6% vs. 88%), whereas germination (53.1% vs. 21.8%) and emergence (32.0% vs. 5.5%) were higher for buried acorns. The quality of some microhabitats remained similar throughout the stages and processes studied. Thus, afforestation provided especially suitable microhabitats for oak establishment, since seed predation was lower, while germination, emergence and seedling survival were higher, than in any other microhabitat. By contrast, the quality of some microhabitats, such as open sites and Holm oaks, differed between recruitment stages. Acorns in open sites escaped predation and germinated easily, but most seedlings died due to summer drought. Similarly, although acorns under Holm oaks can germinate and survive drought, they cannot survive to postdispersal predators. This uncoupling results in a post-dispersal change in the spatial distribution of Q. ilex recruits. Furthermore, there were significant interactions between burial and microhabitat for some demographic processes. The recruitment was in afforestations high irrespective of burial, suggesting that burial is not as beneficial in high-quality habitats as it is in lower-quality ones. An accurate understanding of plant recruitment requires the determination not only of the direct effects of limiting factors but also the potential interactions occurring between them.     

Facilitation of Quercus ilex recruitment by shrubs in Mediterranean open woodlands

Question: Insufficient tree regeneration threatens the long‐term persistence of biodiverse Mediterranean open oak woodlands. Could shrubs, scarce due to decades of management (clearing and ploughing), facilitate holm oak recruitment at both acorn and seedling stages? Location: Open oak woodlands in Central Spain. Methods: Plots with four acorns were planted: (1) under the canopy of the spiny shrub Genista hirsuta, (2) in a small cage, protecting against ungulates, (3) in a shaded cage, protecting against ungulates and sun, and (4) in open grassland. Sets of these four treatments were spatially grouped according to a randomised block design, with 16 blocks near (< 10 m) and 16 away from (> 20 m) parent trees to test for distance‐related survival. Plots were regularly checked for seed removal. After emergence one seedling per plot (97 in total) was selected and its survival monitored. Results: Three months after sowing, 199 of 512 acorns were removed, predominantly by rodents. Acorn removal occurred at each treatment but was highest under shrubs. Eight months after sowing, seedling survival was highest under shrubs (50%), followed by shaded cages (16%), open grassland (4%) and cages (0%). Main mortality cause was drought (90%), killing most seedlings between June and July. No seedlings died from ungulate browsing. Conclusion: Shrubs demonstrated clear net facilitative effects for Quercus ilex recruitment, despite higher seed removal. Shading appears the crucial factor facilitating seedling survival. We therefore propose that lack of shrubs contributes largely to tree recruitment failure in Mediterranean open woodlands; management should aim at conserving shrubs.     

Post-fire regeneration strategies of Californian coastal sage shrubs

Summary Regeneration methods for coastal sage srub vegetation after fire were studied in the coastal Santa Monica Mountains of southern California. Six sites were sampled two years after a large fire of fall, 1978. The intensity of fire varied. Foliar cover and flowering incidence were recorded for individuals regenerating by resprouting or from seed. Resprouting plants contributed most to post-fire recovery, comprising 95% of the relative foliar shrub cover; 84% of resprout and 47% of seedling cover had flowered. An ANOVA of reproductive mode and fire intensity indicates that resprout total cover and individual size are significantly greater than those of seedlings, regardless of fire intensity. Among sites the average foliar cover of resprouts exceeded that of seedlings by factors ranging from 9 to 63. All coastal sage species examined resprout, although the potential vigor of resprouting appears to vary widely within genera (e.g. Encelia, Eriogonum, and Salvia) and even within species. In the second growing season following fire seedling density increased due to seeds shed by resprouted shrubs. Most of the cover on these stands of coastal sage scrub is destined to be either crown-sprouted individuals or their progeny.     

Effects of seed dispersal on Juniperus communis recruitment on a Mediterranean mountain

Abstract. The recruitment of the relict shrub Juniperus communis on a mountain in SE Spain was studied during the period 1994–1998. The main objective was to determine both the quantitative and qualitative effects of bird dispersal on seedling establishment. Seed removal by birds, seed rain, post‐dispersal seed predation, germination, and seedling emergence and survival were analysed in different microhabitats. Birds removed 53 ‐ 89% of the seeds produced by plants. Seed rain was spatially irregular as most seeds accumulated near stones used by birds as perches and below mother plants while a few seeds were dropped in wet meadows and open ground areas. Post‐dispersal seed predation by rodents affected < 10% of dispersed seeds but varied significantly among microhabitats. Only 3.6 ‐ 5.5% of dispersed seeds appeared viable, as many seeds had aborted or showed wasp damage. Seeds germinated in the second and third springs after sowing, reaching a germination percentage of 36%. Seedling emergence was concentrated in wet meadows. Seedling mortality was high (75–80%), but significantly lower in wet meadows, the only microhabitat where seedlings could escape from summer drought, the main mortality cause. Seed abortion, germination and seedling mortality proved to be the main regeneration constraints of J. communis on Mediterranean mountains. Birds exerted a strong demographic effect, although their qualitative effect was limited by abiotic factors which caused the pattern of seed rain to differ from the final pattern of recruitment between microhabitats.     

Spatial patterns of tree seedling establishment and their relationship to environmental variables in a cold-temperate deciduous forest of eastern North America

Patterns of seedling recruitment may have persistent effects on population and community processes. Assuming seed availability is not limiting, the environmental sieve (i.e., the suite of factors influencing seed germination and seedling emergence and survival) determines how many seedlings establish and, most importantly, where they do so. In this study, we identify the spatial structure of some resources and abiotic conditions known to be significant for tree seedling emergence and survival and determine how these environmental factors influence the establishment of Fagus grandifolia, Acer saccharum, Fraxinus americana, and Ostrya virginiana in a deciduous forest of southern Québec (Canada). We expect an increase from Fagus, through Acer and Fraxinus, to Ostrya in the control of environmental variables on seedling emergence and survival, because of differences in the seed size of these species. Density of newly-emerged seedlings of all four species showed positive spatial autocorrelation at distances of up to ca. 10 m. Environmental variables were also structured at the same spatial scale, except for soil moisture. Acer seedling emergence pattern was positively correlated to photosynthetic photon flux density (PPFD), and the pattern of Fraxinus to soil N and moisture. Seedling survival was not spatially autocorrelated for any of the four species, although it was positively density-dependent in Acer and Fagus. In only Ostrya was seedling survival correlated (positively) to one of the environmental variables studied, i.e., PPFD. Overall, environmental variables were spatially less heterogeneous than seedling emergence and survival. Either seed availability was not saturating or factors not considered here, such as competition and predation (the intensity of which often varies with resources and/or abiotic conditions), modified the influence that the physical environment had on patterns of seedling establishment. Our prediction of a greater environmental control on seedling emergence and survival in small-seed species was not totally confirmed.     

Does predation contribute to tree diversity?

Seed and seedling predation may differentially affect competitively superior tree species to increase the relative recruitment success of poor competitors and contribute to the coexistence of tree species. We examined the effect of seed and seedling predation on the seedling recruitment of three tree species, Acer rubrum (red maple), Liriodendron tulipifera (yellow poplar), and Quercus rubra (northern red oak), over three years by manipulating seed and seedling exposure to predators under contrasting microsite conditions of shrub cover, leaf litter, and overstory canopy. Species rankings of seedling emergence were constant across microsites, regardless of exposure to seed predators, but varied across years. A. rubrum had the highest emergence probabilities across microsites in 1997, but Q. rubra had the highest emergence probabilities in 1999. Predators decreased seedling survival uniformly across species, but did not affect relative growth rates (RGRs). Q. rubra had the highest seedling survivorship across microsites, while L. tulipifera had the highest RGRs. Our results suggest that annual variability in recruitment success contributes more to seedling diversity than differential predation across microsites. We synthesized our results from separate seedling emergence and survival experiments to project seedling bank composition. With equal fecundity assumed across species, Q. rubra dominated the seedling bank, capturing 90% of the regeneration sites on average, followed by A. rubrum (8% of sites) and L. tulipifera (2% of sites). When seed abundance was weighted by species-specific fecundity, seedling bank composition was more diverse; L. tulipifera captured 62% of the regeneration sites, followed by A. rubrum (21% of sites) and Q. rubra (17% of sites). Tradeoffs between seedling performance and fecundity may promote the diversity of seedling regeneration by increasing the probability of inferior competitors capturing regeneration sites.     

The effects of drought and fire on seed and seedling dynamics in a tropical seasonal forest in Thailand

Seed production and seedling dynamics were studied for 6 years(1992–1997) with particular emphasis on the effects of the drought andfire in a natural tropical seasonal forest at Mae Klong Watershed ResearchStation, Kantchanaburi, western Thailand. The number of seed produced andemerged seedlings varied greatly among years and species. Most tree speciesproduced very few seeds in 1994 and many in 1995 and 1996, particularly,Shorea siamensis Miq. which displayed the most seedinghabit. Some seedlings, Shorea, Dipterocarpus alatus Roxb.,and Dipterocarpus turbinatus Gaertn. f., emerged at theendof the dry season (late April, after the first rain but when soils still have alow soil moisture content), while the majority of seedlings emerged at thebeginning of the rainy season. After a fire disturbed the plot in April 1996,many species increased their seedling emergences, especially Berryaammonilla. Seedlings of most species less than a year old showed arelatively low mortality in the rainy season, but those ofPterocarpusmacrocarpus Kurz had high mortality. The mean survival rate for allseedling species in this forest was quite low (24.1% y^–1), anddifferent values existed for the rainy and the dry season (11.5, and 6.1%month^–1, respectively). This suggests that different specieshave adaptations related to the season of seedling emergence and resistance todrought in this tropical seasonal forest community. The traits of tree speciesare classified in terms of their traits in the early stages of regeneration;seed and/or seedling bank, resprouting, and drought resistance. Most specieshave adapted to fire and/or drought by resprouting, seed bank, and/or seedlingbank, although the few species which occur mainly in mesic evergreen forestshave less adapted to these environments. The demographic variations in seed andseedling stages may contribute to the coexistence of these species in thisspecies-rich dry tropical forest.     

Short‐term changes affecting regeneration of Fagus crenata after the simultaneous death of Sasa kurilensis

Abstract. Question: The aim of the present study is to determine whether seed/seedling predation will increase and Fagus survival will decline with the recovery of the Sasa cover. Methods: We examined Fagus crenata regeneration for seven years in an old‐growth Fagus‐Sasa forest near Lake Towada, northern Japan, by examining the effects of simultaneous death of Sasa, tree canopy gap formation, mast seeding of Fagus and seed and seedling predation by rodents on the survival of Fagus seeds and current year seedlings. We established four types of sites differing in forest canopy (closed or gap) and Sasa status (dead or alive) following the simultaneous flowering and death of Sasa kurilensis (dwarf bamboo) in 1995. Results: Fallen Fagus seed was abundant in 1997 and 2000 (mast years). In sites with alive Sasa, survival from the first growing season was low due to high seed and seedling predation. In sites with dead Sasa, seed survival under the canopy was high for both mast years, but in gaps it varied between years. Seedling survival was highest in canopy gaps with dead Sasa (gap‐dead) in 1998, because of higher light levels and lower predation by rodents. However, seedling survival in these plots was low in 2001, apparently because rapid Sasa recovery favoured rodent predation. In both mast years, Sasa die‐back had significant positive effects on seed and seedling survival under closed canopies because the seedlings there were more successful in escaping predation. Conclusion: The change in successful sites for the early stage of regeneration of Fagus appears to reflect the combined effects of canopy gap, seed/seedling predation and revegetation of Sasa.     

Species-mediated soil moisture availability and patchy establishment of Pseudotsuga menziesii in chaparral

The occurrence of mature individuals of Pseudotsuga menziesii in stands of Arctostaphylos species mark the initial stages of mixed evergreen forest invasion into chaparral in central coastal California. We planted two cohorts of P. menziesii seeds at three sites under stands of two Arctostaphylos species and Adenostoma fasciculatum in order to determine whether first-year seedling emergence and survival, particularly during the regular summer drought, underlie the spatial distribution of mature trees observed in chaparral. Regardless of the chaparral species they were planted under, P. menziesii seeds that were not protected from vertebrate predation displayed very little emergence and no survival. In contrast, emergence of P. menziesii that were protected from vertebrate predators was much higher but still did not significantly differ among the three chaparral species. However, survival of protected seedlings under Arctostaphylos glandulosa was much greater than under A. fasciculatum, with intermediate survival under Arctostaphylos montana. While mortality of protected seedlings due to insect herbivory, fungal infection, and disturbance displayed no consistent patterns, summer drought mortality appeared to drive the patterns of survival of P. menziesii under the different chaparral species. These emergence, mortality, and survival data suggest that spatial patterns of P. menziesii recruitment in chaparral are driven by first-year summer drought seedling mortality, but only in years when seeds and seedlings are released from vertebrate predation pressure. Because the first-year drought mortality and survival patterns of P. menziesii seedlings differed strongly depending on the chaparral species, we examined the additional hypothesis that these patterns are associated with differences in the availability of soil moisture under different chaparral species. Both higher survival and lower drought mortality of P. menziesii seedlings were associated with higher soil water potential under Arctostaphylos stands during the summer drought, especially in the subsurface soil. The data suggest that Arctostaphylos stands, particularly stands of A. glandulosa, ameliorate xeric summer conditions to a degree that facilitates first-year establishment of P. menziesii and strongly influences spatial distribution of mature trees. Received: 18 September 1998 / Accepted: 23 December 1998     

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.     

High Acorn Predation Prevents the Regeneration of Quercus liaotungensis in the Dongling Mountain Region of North China

The mature oak (Quercus liaotungensis Koidz) forests in the Dongling mountains of northern China have become degraded in recent years because regeneration has been limited. To determine whether or not seedling establishment of the oak is seed limited, microsite limited, or predator limited and to determine whether seedling establishment is affected by ground cover, we conducted field experiments during a mast year and investigated the fate of seeds and the soil seed bank dynamics of the oak. A large acorn crop (128.8 acorns/m²) was observed in the study period, and the peak density of acorns on the forest floor reached 46.5 acorns/m², suggesting that tree recruitment was not seed limited. Acorns in the soil seed bank were mainly lost through decay (principally after fungal attack), consumption in situ, and removal by animals. Predation (including consumption in situ and removal) accounted for 86.4% of acorn loss and was therefore likely to have been the most important factor influencing seed dynamics. More than 70% of acorns were found to have germinated, but no established seedling was observed on the forest floor. Using cages to exclude predators, it was estimated that 87% of acorns germinated and 49% became established as seedlings, indicating that the acorns on the forest floor could emerge and grow in the absence of predators. We conclude that the regeneration of the tree population is limited by predators rather than by the availability of microsites. The presence of ground cover increased the germination rate and increased the chance of seed survival in the early stage of the experiment, but at the end of the investigation, no established seedling was found in the quadrats both with and without ground cover, possibly because of high density of animal predators. On the basis of these results, we suggest that selective tree felling will increase the coverage of the herbaceous layer, which can further decrease the population density of the rodents, and thereby improve the regeneration of oak trees.     

Effect of Acorn Burying Depth on Germination,Seedling Emergence and Development of Quercus aliena var. acuteserrata

Acorns of Quercus aliena var. acuteserrata Maxim. are often predated by small mammals and birds in natural forests. These animals not only eat the acorns during the acorn ripening season, but also cache and hoard most of the remaining acorns on the forest floor in the soil for their future use. These buried acorns form the main seed resource for regeneration. Burying depth is potentially important for germination and for seedling development. The effects of burying depth on germination and seedling development in relation to acorn size were studied in an experiment, in which acorns were planted at 6 cm-, 12 cm- and 18 cm-depth. The experimental results showed that fewer acorns germinated as burying depth increased. From the deeply buried acorns fewer seedlings emerged at later time than from those acorns buried less deeply. They appeared to have more difficulties to emerge above-ground than the seedlings from shallowly buried acorns. The deeply buried acorns and their seedlings also appeared to be more susceptible to rot. Acorn size did not significantly affect germination and emergence of the seedlings. As early emerged seedlings had longer developmental periods in their first growing season, and therefore grew better than the late emerged seedlings, seedlings from the shallowly buried acorns took the advantage.     

Effects of increasing root carbon investment on the mortality and resprouting of Haloxylon ammodendron seedlings under drought

• Tree mortality induced by drought is one of the most complex processes in ecology. Although two mechanisms associated with water and carbon balance are proposed to explain tree mortality, outstanding problems still exist.
• Here, in order to test how the root system benefits survival and resprouting of Haloxylon ammodendron seedlings, we examined the various water‐ and carbon‐related physiological indicators (shoot water potential, photosynthesis, dark respiration, hydraulic conductance and non‐structural carbohydrates [NSC]) of H. ammodendron seedlings, which were grown in drought and control conditions throughout a grow season in greenhouse.
• The survival time of the seedling root system (died 70 days after drought) doubled the survival time of the shoot (died at 35 days). Difference in survival time between shoot and root resulted from sustained root respiration supported by increased NSC in roots under drought. Furthermore, investment into the root contributed to resprouting following drought. Based on these results, a death criterion is proposed for this species. The time sequence of major events indicated that drought shifted carbon allocation between shoot and root and altered the flux among different sinks (growth, respiration or storage). The interaction of water and carbon processes determined death or survival of droughted H. ammodendron seedlings.
• These findings revealed that the ‘root protection’ strategy is critical in determining survival and resprouting of this species, and provided insights into the effects of carbon and water dynamics on tree mortality.

     

锐齿槲栎橡子埋藏深度对发芽、幼苗出土和发育的影响

锐齿槲栎（Quercus aliena var.acuteserrata Maxim.)林内活动的许多型哺乳动物和鸟类不仅采食橡子,而且还常窖贮或在土壤大量的橡 子以便季后利用。这些埋藏的橡子构成了发育有效更新苗的主要种子来源。橡子被埋藏的深度和橡子的大小对幼苗的发育有潜在的影响。把橡子埋藏在6、12和18cm3个土层深度的实验结果显示：随着埋藏深度的增加,橡子的发芽率和发芽橡子的幼苗出土率都有明显下降,由深埋橡胶子发育的幼苗出土比较困难,所需时间较长,也更易腐烂,橡子的大小对发芽和幼苗出土没有明显的影响,早出土幼苗在第一个生长季内由于有较长的生长发育期,季末发育得较好,所以,由浅埋橡子发育的幼苗在更新成功方面有优势。     

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

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

Competitive effects of herbaceous vegetation on tree seedling emergence,growth and survival: Does gap size matter?

Question: What is the effect of gap size on the seedling emergence, growth and survival of four common tree species in wooded pastures? Location: A pasture in the Jura mountains, Switzerland. Methods: Seeds were sown in a complete three‐way factorial design with eight blocks in May 2003. Each block consisted of a competition treatment (four gap sizes including zero) and a mowing treatment (mown and unmown). Emergence, survival and total biomass of tree seedlings of three species (Picea abies, Acer pseudoplatanus and Fagus sylvatica) were measured. A fourth species (Abies alba) failed to germinate. Results: Gaps had a positive influence on the early stages of tree development for all species. Larger gaps favoured growth and survival more than small gaps. Seedling growth was higher when vegetation around the openings was mown. Mowing the vegetation at gap size zero enhanced both growth and survival compared to unmown vegetation. Mown gaps larger than zero had increased seedling desiccation but decreased seedling predation. Species showed similar trends in their emergence and growth responses to gap size and mowing treatments but for Picea emergence rate was higher and survivorship was lower than for Acer and Fagus. Conclusions: Gap size does matter for tree seedling success but even in more favourable large gaps only a small percentage of seedlings emerged and survived. The effects of gaps on tree seedling establishment are complex as a result of interactions between biotic and abiotic changes caused by gaps.     

Advantage of early seedling emergence in <Emphasis Type="Italic">Fagus crenata</Emphasis>: importance of cotyledon stage for predator escape and pathogen avoidance

To examine how early-emerging seedlings take advantage of establishment in a deciduous forest, we explored the relationships among the emergence date, growth stage and major causes of mortality (damping-off by fungi and predation by rodents) in seedlings of Fagus crenata. The emergence of current-year seedlings and their survival and damage were followed at short (3–7 days in early spring) intervals for a growing season. The growth stage was divided into two stages, with only cotyledons (CT stage) and with true leaves (TL stage). The survival rate was negatively correlated with the emergence date, indicating the advantage of early seedling emergence. This advantage was largely explained by the lower occurrence of damping-off. In contrast, seedling predation occurred independently of the emergence date, but depended strongly on the developmental stage. Rodents consumed a considerable number of seeds during the early period after emergence, and strongly preferred CT-stage over TL-stage seedlings throughout the growing season. Therefore, seedling predation was inferred to be concentrated in a relatively short period while the remaining seeds were depleted and CT-stage seedlings were abundant. The seedling stage synchronously shifted from the CT to TL stage. This growth-stage transition was independent of the emergence date and appeared to correspond with the timing of seed depletion. Delayed stage transition resulted in a disproportionately high risk of damping-off later in the season. Our results indicate that early seedling emergence is advantageous for F. crenata for resistance to pathogens and that the timing of growth-stage progression of seedlings appears important to escape predation because of the distinct food preference of predators.     

Effects of rodent gnawing on the survival of current-year seedlings of Quercus crispula

Emergence, survival and mortality patterns of current-year oak (Quercus crispula Blume) seedlings were investigated for 4 consecutive years in a secondary oak forest in Hokkaido, northern Japan. Despite the emergence of a considerable number of oak seedlings in the years following masting, few current-year seedlings survived until the end of the growing season. Almost all of the seedlings died from damage to their stems caused by the gnawing of rodents. Rodent gnawing on transplanted oak seedlings was also observed in the year following masting but not in the year following a bad crop year. Cuttings of dwarf bamboo, Sasa, did not reduce the seedling mortality caused by gnawing. However, transplanted oak seedlings were gnawed more quickly when they were placed on the forest floor with a thicker Sasa covering. All rodents trapped in the vicinity of the study area were Apodemus speciosus Temminck. These results suggest that rodents strongly influence the recruitment of oak trees not only through the predation and dispersal of acorns but also through gnawing seedlings.     

Multivariate path analysis of the relationships between seedling regeneration and environmental factors beneath a dwarf bamboo understory

Seedling emergence and establishment are fragile processes that determine the direction and structure of forest succession and regeneration. However, seedling emergence and establishment are easily affected by biotic and abiotic (environmental) factors. A dense and expanding understory of dwarf bamboo is one such important factor that can seriously hinder the seedling regeneration. We conducted a field experiment to investigate the emergence and establishment of canopy tree seedlings under artificially controlled densities of dwarf bamboo. We found that understory dwarf bamboo obstructed seedling emergence but reduced the death of seedlings. Although understory dwarf bamboo reduced the median retention time of seedlings, dense bamboo increased the mean survival time of seedlings. Our results suggest that understory dwarf bamboo has multiple selectivities for tree seedling emergence and establishment: high‐density dwarf bamboo was beneficial to evergreen species but lower‐density of bamboo was conducive to the survival of deciduous species, it means the dwarf bamboo potentially alters successional trajectories of forest communities. Path analysis revealed that the most important factors affecting tree seedling emergence and death were the abundance of seeds in the seed bank and the density of emerged seedlings, and that the soil temperature promoted seedling emergence but increased seedling death, the thickness of litter limited seedling emergence, and the leaf area index of the bamboo canopy limited seedling death. The present study suggests that dwarf bamboo can directly alter the microenvironment, significantly reducing light levels and soil temperature but increasing the thickness of litter and soil humus, thereby indirectly impacting the regeneration of tree seedlings. Our results indicate that various factors affected seedling emergence, and there were complex indirect relationships among these factors. In general, biological factors had a stronger influence on tree seedling regeneration than 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