An interactive effect of simultaneous death of dwarf bamboo, canopy gap, and predatory rodents on beech regeneration

To clarify the interactive effect of the simultaneous death of dwarf bamboo (Sasa kurilensis), forest canopy gap formation, and seed predators on beech (Fagus crenata) regeneration, we analyzed beech demography from seed fall until the end of the first growing season of seedlings in an old-growth forest near Lake Towada, northern Japan. The simultaneous death of S. kurilensis took place in 1995. We established four types of sampling site differing in forest canopy conditions (closed or gap) and Sasa status (dead or alive). Beech seed survival and emergence ratio were both highest in gaps with dead Sasa (gap-dead), because rate of predation was lowest. Seedling survival during the first growing season was also highest in the gap-dead treatment, because of less predation and less damping off. As a result, even though density of seed fall was lowest in the gap-dead treatment, the living seedling density there was highest at the end of the first growing season. Predation, which caused the greatest mortality during the seed and seedling stages, was significantly lower at both sites in gaps and sites with dead Sasa. This was probably due to changes in the behavior of rodents in response to the structure of the forest canopy and undergrowth. Both the death of Sasa and canopy gap formation allowed seedlings to avoid damping off because of the high light availability. The indirect effect of the simultaneous death of Sasa and canopy gap formation in reducing predation contributed more to beech regeneration than their direct effect in increasing light for the seedlings.     

The effects of Sasa and canopy gap formation on tree regeneration in an old beech forest

Abstract. We examined the response of tree seedling emergence and survival to the dieback of Sasa and canopy gap formation in an old‐growth forest near Lake Towada, northern Japan. Synchronous death of Sasa occurred in 1995. We established four types of sampling sites differing in forest canopy conditions (Closed or Gap) and Sasa status (Dead or Live). Gap‐Dead sites had the highest light levels and the greatest fluctuation in soil temperatures. The death of Sasa alone facilitated the emergence (Acer japonicum, Fagus crenata, Fraxinus lanuginosa, and Tilia japonica) and survival (Acanthopanax sciadophylloides, F. crenata, F. lanuginosa, Kalopanax pictus, and Sorbus commixta) of species with a seedling bank strategy. Cercidiphyllum japonicum grew at all sites at a higher density than other species, but survived well only in Gap‐Dead sites. This behaviour was associated with a seed rain strategy. The additive effects of Sasa death and canopy gap formation promoted seedling emergence of pioneer tree species (Betula maximowicziana, Lindera umbellata, and Magnolia obovata), probably through break of dormancy by the large temperature fluctuation. In addition, the scarcity of advance regeneration in canopy gaps due to Sasa cover facilitates the regeneration of pioneer species. The dominance and dieback cycle of Sasa contributes to species diversity in this forest.     

Multiple stages of tree seedling recruitment are altered in tropical forests degraded by selective logging

Tropical forest degradation is a global environmental issue. In degraded forests, seedling recruitment of canopy trees is vital for forest regeneration and recovery. We investigated how selective logging, a pervasive driver of tropical forest degradation, impacts canopy tree seedling recruitment, focusing on an endemic dipterocarp Dryobalanops lanceolata in Sabah, Borneo. During a mast‐fruiting event in intensively logged and nearby unlogged forest, we examined four stages of the seedling recruitment process: seed production, seed predation, and negative density‐dependent germination and seedling survival. Our results suggest that each stage of the seedling recruitment process is altered in logged forest. The seed crop of D. lanceolata trees in logged forest was one‐third smaller than that produced by trees in unlogged forest. The functional role of vertebrates in seed predation increased in logged forest while that of non‐vertebrates declined. Seeds in logged forest were less likely to germinate than those in unlogged forest. Germination increased with local‐scale conspecific seed density in unlogged forest, but seedling survival tended to decline. However, both germination and seedling survival increased with local‐scale conspecific seed density in logged forest. Notably, seed crop size, germination, and seedling survival tended to increase for larger trees in both unlogged and logged forests, suggesting that sustainable timber extraction and silvicultural practices designed to minimize damage to the residual stand are important to prevent seedling recruitment failure. Overall, these impacts sustained by several aspects of seedling recruitment in a mast‐fruiting year suggest that intensive selective logging may affect long‐term population dynamics of D. lanceolata. It is necessary to establish if other dipterocarp species, many of which are threatened by the timber trade, are similarly affected in tropical forests degraded by intensive selective logging.     

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.     

Sugar maple (Acer saccharum Marsh.) establishment in boreal forest: results of a transplantation experiment

Aim To evaluate whether seedlings of sugar maple (Acer saccharum Marsh.) can establish beyond the species northern range limit in adjacent boreal forest. Location The hardwood–boreal forest transition zone on the north‐east shore of Lake Superior, Ontario, Canada. Methods Seed fall of sugar maple was monitored for 5 years in a stand of this species at its northern range limit, and seed from this stand was transplanted to five micro‐habitat types in an adjacent boreal forest. The establishment and survival of sugar maple seedlings there, and in the seed‐source stand, was monitored for the following 7–11 years. Soil‐surface light levels were measured in both forest types. Results Most seed fell in the final year of monitoring, when c. 250 seeds m^?2 were recorded. First‐year seedling establishment rates in the maple stand, deriving from this mast seed year, was approximately double that deriving from seed transplanted to the boreal forest sites; this is tentatively attributed to seed predator satiation in the maple stand. However, subsequent seedling survivorship in the boreal forest was greater than that in the maple stand, resulting in comparable seedling densities by the end of 6 years. This difference is tentatively attributed to better illumination in the boreal forest sites, and canopy‐opening disturbances appear to be especially facilitative of seedling survival. Main conclusions There is no fundamental impediment to sugar maple seedlings establishing in boreal forest communities if climate warming occurs and seed is available. If management intervention is needed to accelerate seed availability in a rapidly warming boreal forest, then diffuse seed application to disturbed boreal forest sites during mast years of local boreal tree species is recommended as the most effective way of avoiding seed predation and increasing seedling survival.     

Regeneration of beech (Fagus crenata) after the simultaneous death of undergrowing dwarf bamboo (Sasa kurilensis)

Seedling recruitment and survivorship of beech (Fagus crenata) were studied with special reference to the simultaneous death of undergrowing bamboo (Sasa kurilensis). The survival rate of beech seedlings on the floor whereSasa had withered was much higher than that on the floor whereSasa survived. Damping off caused the largest mortality among beech seedlings. However, the allocation pattern of matter to different parts of the seedlings indicated that their survival was greatly affected by production economy. The dense cover of dwarf bamboo prevented the establishment of beech seedling banks on the forest floor. The interval between the times when simultaneous death ofSasa occur and the length of its recovery period are thus important factors controlling the dynamics of beech forests in Japan.     

Seedling bank dynamics of Fagus grandifolia var. mexicana before and after a mast year in a Mexican cloud forest

Abstract. Fagus grandifolia var. mexicana seedling dynamics were studied in a relict forest inside the crater of a volcano where it is the only canopy tree species. The objectives of the study were (1) to determine changes in seedling density before and after a mast year, as well as to monitor seedling recruitment, mortality and growth rate in relation to the micro‐environment, and (2) to discuss the impact of masting on seedling dynamics of the population. Before masting, seedling density was 8.8 seedlings m^–2, afterwards seedling density peaked at 51.5 seedlings m^–2, then decreased by 50% in 3 months and by 85% after 8 months, down to levels of before the mast year (9.9 seedlings m^–2) after 16 months. Seedlings in the forest floor before the mast‐seeding event showed a low relative growth rate – 0.14 mm mm^–1 mo^–1 against 1.27 mm mm^–1 mo^–1 for seedlings that emerged immediately after masting. Seed germination was high (83%) and rapid (L₅₀= 7 days) following masting. Seedling growth was positively correlated with soil water content but not with temperatures and relative humidity. Data suggested that seed production in mast years is important in maintaining the seedling bank, and thus the viability of relict Fagus populations.     

A relict population of Fagus grandifolia var. mexicana at the Acatlan Volcano, Mexico: structure, litterfall, phenology and dendroecology

Aim Fagus grandifolia var. mexicana (Martinez) Little has an extraordinarily restricted distribution in the Mexican montane cloud forests. Isolated Fagus (beech) populations have been recorded in less than 10 small areas (2–40 ha) in the eastern Sierra Madre at altitudes from 1400 to 2000 m. The objectives were to determine tree and seedling age, forest structure, phenology, litterfall patterns and the relationship between mast and climatic variables. Location We report on three Fagus stands at the Acatlan Volcano, Veracruz, Mexico. Methods Changes in forest cover were determined using aerial photographs. Within each stand, basal area, density and tree species composition were determined in a 0.1‐ha band transect. Additionally, litterfall production was quantified and phenophases were recorded monthly over a 3‐year period, and 60 tree cores were collected to determine age distribution and tree‐ring growth. Results The forest was atypical in several respects. Fagus was the only dominant tree species in the crater stand, although in the rim and at the top of the volcano it was codominant with other tree species. Juveniles occurred only on the rim, but there was a seedling bank in the crater. Although forest cover in the area increased between 1968 and 1993, the Fagus stands did not change in size. Leaf production peaked in March and April, and leaf fall occurred from October through February. Litterfall production was the highest in November. During mast years, flowering started in February and between mast events there were no flowers or fruits. Minimum temperatures were highly correlated with Fagus litterfall and leaf fall. Seedlings ranged in age from 2 to 18 years and were 13–60 cm tall. Tree cores ranged from 76 to 120 years, but trees were older than the core samples. Main conclusions Although beech is considered a gap regeneration species that reaches the canopy after alternating periods of release and suppression, the trees in the crater were released when less than 1.5 m tall and have suffered few periods of suppression since. The results indicate that the crater stand was established after a severe disturbance destroyed the existing forest. We conclude that the relict beech population should be able to maintain itself, if not severely disturbed by humans or by climatic changes related to global warming.     

Seed survival on experimental dishes in a central European old‐growth mixed‐species forest – effects of predator guilds,tree masting and small mammal population dynamics

Predation of tree seeds can be a major factor structuring plant communities. We present a three year study on tree seed survival on experimental dishes in an old‐growth forest in central Europe in Austria. We addressed species specific, spatial and temporal aspects of post‐dispersal seed predation. Seeds of Norway spruce Picea abies, European beech Fagus sylvatica, and silver fir Abies alba were exposed on dishes in different types of exclosures which allowed access only to specific guilds of seed predators. Removal experiments were carried out in two old‐growth forests and a managed forest (macro‐sites), including micro‐sites with and without cover of ground vegetation. We conducted the experiment in three consecutive years with a mast year of beech and spruce before the first year of the study. The seed removal experiments were combined with live trapping of small mammals being potential seed predators. Our experiments showed a distinctly different impact of different predator guilds on seed survival on the dishes with highest removal rates of seeds from dishes accessible for small mammals. We observed differing preferences of small mammals for the different tree species. Seed survival in different macro‐ and micro‐habitats were highly variable with lower seed survival in old growth forests. In contrast to our assumption, and in contrast to the satiation hypothesis which assumes higher seed survival in and directly after mast years, seed survival was lower in the year following the mast year of beech when a population peak of small mammals occurred and higher in intermast periods when subsequently small mammal population crashed. This suggests a higher importance of sporadic masting shortly after mast years in intermast periods for establishment of forest trees provided that pollination efficiency is high enough in such years. Combined with the high seed mortality observed after the mast year, this corroborates the important role of seed predation for forest dynamics. An altered synchrony or asynchrony of masting of different tree species and changed masting frequencies through climate change may thus lead to strong and non‐linear effects on forest dynamics.     

Neighbourhood interactions and environmental factors influencing old‐pasture succession in the Central Pyrenees

Abstract. The shrub Buxus sempervirens and the trees Abies alba and Fagus sylvatica have recently recolonized old‐pastures in the Central Pyrenees. We mapped all live and dead individuals (> 1.3 m tall) in a large forest plot in Ordesa Valley to examine the importance of density‐dependent processes during recolonization. Biotic interactions were inferred from changes in horizontal structure and the influences of neighbours on tree survival. Buxus differentially influenced establishment and survival of tree species, thereby controlling future canopy composition and spatial structure. The rapidly invading Abies formed denser patches on elevated sites less occupied by Buxus, whereas Fagus preferentially established within shrubs. Abies reached densities which led to intense intraspecific competition and high mortality rates among saplings. Self‐thinning in Abies led to smaller numbers of regularly spaced survivors, and greater relative dominance of Fagus. Disregarding intraspecific effects and abiotic environment, Abies survival was significantly lower under Buxus shrubs, which suggests that the spatial location and abundance of Abies was constrained by the location of Buxus. Fagus survival was not related to Buxus density, but remained significantly lower in denser Abies patches. The higher mortality of Fagus in denser Abies patches, and the resulting spatial segregation of the species, reflects asymmetric interspecific competition. Inhibition from Buxus shrubs and intraspecific competition prevent invading Abies from dominating and may therefore help in maintaining a mixed Abies‐Fagus stand.     

Regeneration process of coniferous forests in northern Hokkaido I.Abies sachalinensis forest andPicea glehnii forest

Regeneration of natural forests was studied in the Nakagawa Experiment Forest of Hokkaido University using age distribution surveys made by the clear felling method. In Plot 1 (30 m × 65 m),Abies sachalinensis dominated the canopy layer but there were also a fewBetula ermanii trees.Sasa senanensis densely covered the forest floor. Most of the canopy trees were from 122 to 195 years old. Seedlings younger than 50 years old ofA. sachalinensis were found on fallen logs and root bases. There were, however, few trees from 50 to 120 years old. The present canopy trees seemed to have regenerated after competitive pressure from old canopy andSasa disappeared 180 years ago. Plot 2 (50 m × 100 m) on serpentinite soil was dominated byPicea glehnii. Sasa kulirensis covered the floor but not as densely asS. senanensis in Plot 1. The ages ofP. glehnii ranged from 1 to 586 years old, and the age distribution ofA. sachalinensis was L-shaped. A small gap in the canopy formed about 290 years ago, and it gradually extended. Conifers regenerated continuously in the extending gap butB. ermanii did not. One hundred thirty years ago, part of Plot 2 was again destroyed andA. sachalinensis andB. ermanii regenerated. Thus, two types of regeneration were found. One regenerated both conifers andBetula after a sudden disturbance of canopy layer or death ofSasa, and the other, under an extending gap, regenerated only conifers.     

Gnawing damage by rodents to the seedlings ofFagus crenata andQuercus mongolica var.grosseserrata in a temperateSasa grassland-deciduous forest series in southwestern Japan

The effects of dwarf bamboo,Sasa, cover on the initial morrality of hardwood seedlings were investigated by transplanting 1-year-old beech (Fagus crenata) and current-year oak (Quercus mongolica var.grosseserrata) seedling to three different stands; old-growth beech and secondary oak forests withSasa undergrowth, and aSasa grassland in a grassland-forest series near the top of Mt Jippo, southwestern Japan. The most frequent cause of seedling morrality was gnawing of the stems by rodents. In the beech forest, the gnawing was more likely to occur underSasa cover, suggesting that it provides a good habitat for rodents on the beech forest floor. TheSasa under growth may thus play an imporrant role in regeneration of beech forest. In the oak floor, mortality of both species was low and only a little gnawing occurred during a year. However, no natural oak seedling were found in the forest even after a mast year. This may be because most of the acorns disappeated before establishment. The early-stage demography of hardwood seedling as oak may thus play an imporrant role in regeneration of oak forest. In theSasa grassland where the seed supply is small, almost all of the seedlings died fromo gnawing regardless of the presence ofSasa cover. These factors prevent the recruitment of a sizable seedling bank. Rodents may thus play an imporrant role in maintenance of theSasa grassland.     

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.     

Seedling establishment in a masting desert shrub parallels the pattern for forest trees

The masting phenomenon along with its accompanying suite of seedling adaptive traits has been well studied in forest trees but has rarely been examined in desert shrubs. Blackbrush (Coleogyne ramosissima) is a regionally dominant North American desert shrub whose seeds are produced in mast events and scatter-hoarded by rodents. We followed the fate of seedlings in intact stands vs. small-scale disturbances at four contrasting sites for nine growing seasons following emergence after a mast year. The primary cause of first-year mortality was post-emergence cache excavation and seedling predation, with contrasting impacts at sites with different heteromyid rodent seed predators. Long-term establishment patterns were strongly affected by rodent activity in the weeks following emergence. Survivorship curves generally showed decreased mortality risk with age but differed among sites even after the first year. There were no detectable effects of inter-annual precipitation variability or site climatic differences on survival. Intraspecific competition from conspecific adults had strong impacts on survival and growth, both of which were higher on small-scale disturbances, but similar in openings and under shrub crowns in intact stands. This suggests that adult plants preempted soil resources in the interspaces. Aside from effects on seedling predation, there was little evidence for facilitation or interference beneath adult plant crowns. Plants in intact stands were still small and clearly juvenile after nine years, showing that blackbrush forms cohorts of suppressed plants similar to the seedling banks of closed forests. Seedling banks function in the absence of a persistent seed bank in replacement after adult plant death (gap formation), which is temporally uncoupled from masting and associated recruitment events. This study demonstrates that the seedling establishment syndrome associated with masting has evolved in desert shrublands as well as in forests.     

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.     

Early post-fire vegetation regeneration in <Emphasis Type="Italic">Larix kaempferi</Emphasis> artificial forests with an undergrowth of <Emphasis Type="Italic">Sasa senanensis</Emphasis>

Post-fire vegetation regeneration was studied for a 6-year period in a 13-year-old-artificial forest consisting of Larix kaempferi with a dense undergrowth of Sasa senanensis. The study site was classified into three fire severity categories according to the degree of Sasa senanensis scorching, that is, a high-severity category, a mid-severity category, and a low-severity category. Study plots were established in areas which fitted the criteria for each category, and in nearby unburned sites. A total of 41 woody species were newly emerged during the 6-year study period in the burned and unburned plots. Only a few seedlings and resprouts emerged in the unburned plots, while many seedlings emerged in the high-severity plots from the first year after fire onward. A high-severity fire that burns the rhizomes of Sasa is necessary for the vegetation recovery by germination of seed. Whereas the establishment of seedlings was restricted to a few years after fire, the regeneration through resprouting continued into the last year of observation. The survival time of resprouts was longer than that of seedlings, and the survival time of shade-tolerant species was longer than that of shade-intolerant species. In contrast, shade-intolerant species grew more rapidly than shade-tolerant species. The plants ability to exceed the maximum height of the Sasa before the bamboo recovers can be critical to the survival of shade-intolerant species. Because resprouts have a stronger resistance to the shade of Sasa than seedlings, the resprouts of shade-tolerant species play a major role in the re-establishment of woody species after fire in sites with considerable Sasa ground-cover.     

Factors Affecting Seed Germination and Seedling Establishment of Fen-Meadow Species

Availability of seeds and provision of “safe sites” for seedling recruitment are essential for successful restoration of seminatural grassland communities. Inability to provide species‐specific conditions for seedling recruitment appears to be a major factor limiting establishment of fen‐meadow species on restoration sites. This contention was tested in the field for both germination and establishment conditions for a selection of fen‐meadow species. A Cirsio‐Molinietum fen meadow and an agriculturally semi‐improved species‐poor grass dominated rush pasture were used. Seeds of Carex ovalis, Cirsium dissectum, Molinia caerulea, Succisa pratensis, and Holcus lanatus were sown onto treatments comprising either irrigation or no irrigation, presence or absence of existing vegetation canopy, and presence or absence of soil disturbance. Germination of all except H. lanatus was higher in the fen meadow than in the rush pasture. The fen‐meadow site was less susceptible to drought, provided more light to the seed environment, and showed a stronger day–night variation in relative humidity compared with the rush pasture. All the fen‐meadow species responded strongly to the experimental treatments, whereas H. lanatus showed only a small response. Soil disturbance was the major factor that increased germination. Removal of the vegetation canopy improved germination only in S. pratensis. Conditions affecting survival of seedlings were different from those affecting seed germination. Seedling survival was greater on the fen‐meadow site than on the rush pasture. Canopy presence was the major factor that reduced seedling survival. Few seedlings survived in the presence of the rush pasture canopy. Irrigation and soil disturbance were of minor importance for seedling survival on both sites. Safe sites for seed germination and seedling establishment of fen‐meadow species existed on the fen meadow even without soil disturbance and gap creation. Safe sites for seedling recruitment were not present in the rush pasture. The need for species‐specific definition of safe site characteristics at the two stages of seedling recruitment (i.e., for seed germination and for seedling survival) was demonstrated. The implications of these findings for restoration of seminatural grasslands are discussed.     

Reproductive potential and seedling establishment of the invasive alien tree Schinus molle (Anacardiaceae) in South Africa

Schinus molle (Peruvian pepper tree) was introduced to South Africa more than 150 years ago and was widely planted, mainly along roads. Only in the last two decades has the species become naturalized and invasive in some parts of its new range, notably in semi‐arid savannas. Research is being undertaken to predict its potential for further invasion in South Africa. We studied production, dispersal and predation of seeds, seed banks, and seedling establishment in relation to land uses at three sites, namely ungrazed savanna once used as a military training ground; a savanna grazed by native game; and an ungrazed mine dump. We found that seed production and seed rain density of S. molle varied greatly between study sites, but was high at all sites (384 864–1 233 690 seeds per tree per year; 3877–9477 seeds per square metre per year). We found seeds dispersed to distances of up to 320 m from female trees, and most seeds were deposited within 50 m of putative source trees. Annual seed rain density below canopies of Acacia tortillis, the dominant native tree at all sites, was significantly lower in grazed savanna. The quality of seed rain was much reduced by endophagous predators. Seed survival in the soil was low, with no survival recorded beyond 1 year. Propagule pressure to drive the rate of recruitment: densities of seedlings and sapling densities were higher in ungrazed savanna and the ungrazed mine dump than in grazed savanna, as reflected by large numbers of young individuals, but adult : seedling ratios did not differ between savanna sites. Frequent and abundant seed production, together with effective dispersal of viable S. molle seed by birds to suitable establishment sites below trees of other species to overcome predation effects, facilitates invasion. Disturbance enhances invasion, probably by reducing competition from native plants.     

六盘山区辽东栎的实生苗更新及其影响因子

研究了六盘山区辽东栎(Quercus liaotungensis)灌丛的种子产量、土壤种子库组成、湿沙和风干贮藏对种子寿命的影响以及动物取食子叶对种子萌发和幼苗建立的影响。结果表明, 辽东栎灌丛种子的完好率为27.51%, 被动物取食或搬运种子的比例(41.51%)显著高于其他类型种子(p < 0.01); 辽东栎次生林土壤种子库中萌发和虫蛀种子分别占35.16%和38.29%, 完好种子仅占13.65%, 捕食动物主要通过贮藏或搬运而影响土壤种子库中的种子密度。湿沙贮藏60天的辽东栎种子自动萌发率高达96.67%, 短期贮藏可加快种子的萌发进程, 提高萌发率和萌发指数, 但随着贮藏时间的延长, 种子萌发进程延迟, 萌发率、萌发指数和活力指数均不同程度地降低。排除动物取食处理的幼苗在林窗和林下生境的存活率分别为80%和83%, 而不排除动物取食幼苗在2种生境中分别仅有25%和31%能够存活, 表明子叶在幼苗建立中具有重要作用。林窗中幼苗子叶的动物取食率(85.00%)高于林下(71.00%), 子叶留存的幼苗在林窗中的存活率(6.00%)低于林下(15.50%), 而子叶被取食幼苗的存活率在两种环境中基本相等(分别为18.50%和18.00%)。     

Regeneration of<Emphasis Type="Italic"> Rhizophora mangle</Emphasis> in a Caribbean mangrove forest: interacting effects of canopy disturbance and a stem-boring beetle

Current theory predicts that in low-density, seed-limited plant populations, seed predation will be more important than competition in determining the number of individuals that reach maturity. However, when plant density is high, competition for microsites suitable for establishment and growth is expected to have a relatively greater effect. This dichotomous perspective does not account for situations in which the risk of seed predation differs inside versus outside recruitment microsites. We report the results of a field experiment and sampling studies that demonstrate such an interaction between microsite quality and the risk of propagule predation in mangrove forests on the Caribbean coast of Panama, where it appears to play a key role in shaping the demography and dynamics of the mangrove, Rhizophora mangle. Rhizophora's water-borne propagules establish wherever they strand, but long-term sampling revealed that only those that do so in or near lightning-created canopy gaps survive and grow to maturity. These microsites afford better growth conditions than the surrounding understory and, as importantly, provide a refuge from predation by the scolytid beetle, Coccotrypes rhizophorae. This refuge effect was confirmed with a field experiment in which Rhizophora seedlings were planted at different positions relative to gap edges, from 5 m inside to 20 m outside the gap. Mortality due to beetle attack increased linearly from an average of 10% inside a gap to 72% at 20 m into the forest. The interaction between canopy disturbance and propagule predation may be having a large impact on the composition of our study forests. Being shade-tolerant, Rhizophora seedlings that escape or survive beetle attack can persist in the understory for years. However, the high rate of beetle-induced mortality effectively eliminates the contribution of advance regeneration by Rhizophora saplings to gap succession. This may explain why the shade-intolerant mangrove, Laguncularia racemosa, is able to co-dominate the canopy in low intertidal forests at our study sites.