植物种群更新限制——从种子生产到幼树建成

更新限制是指种子由于各种原因,不能够萌发并生长成幼树。它作为解释生物多样性的理论,一直受到国内外群落生态学家关注。从种源限制、传播限制和建成限制3个角度,对更新限制机制研究进展进行了综述。从种源限制而言,时空因素是影响植物种群更新限制的重要因素,因为植物结实量存在明显时空变化,造成植物更新个体出现明显的时空规律。从传播限制而言,传播数量、距离和食果动物行为均限制植物种群更新。数量上,缺乏有限传播者势必减少传播数量,但如果种子拥有较高质量,则能逃脱数量限制;距离上,植物更新个体显示出明显的Janzen-Connell格局,但传播距离趋向稳定,形成植物种群的进化稳定对策;食果动物行为上,不同传播者对更新贡献存在差异,捕食者直接降低更新,融入两类动物行为的模型更能反映食果动物对更新的限制。从建成限制而言,环境因子制约植物生长。小尺度下,微生境的好坏对于植物幼苗建成至关重要;大尺度下,植物提供较好的广告效应则能摆脱生境限制。将传播者行为、捕食者行为与幼苗的空间分布格局、种子传播机理模型等结合,建立植物更新限制机理模型应是更新限制未来的研究热点。选择稀有种和古老种为主题的长期更新限制研究,为种群恢复提供指导,也是未来重要研究方向。     

Hypericum cumulicola demography in unoccupied and occupied Florida scrub patches with different time‐since‐fire

1 Metapopulation models predict that unoccupied, but suitable, patches will exist for species subject to extinction and colonization dynamics. We compared the demographic responses of Hypericum cumulicola , a rare herbaceous species almost entirely restricted to Florida rosemary scrub, when transplanted to occupied or unoccupied patches.
2 Seedlings were transplanted and seeds buried into Florida rosemary scrub patches differing in time since last fire, and in the presence or absence of H. cumulicola. We used a replicated, factorial design to place the transplants and seeds in the field, and monitored their performance for 18 months.
3 Neither time‐since‐fire nor prior H. cumulicola site occupancy affected survival of transplants. Only time‐since‐fire affected growth. Time‐since‐fire, H. cumulicola occupancy, and their interaction affected reproductive effort, but these effects were not consistent between years.
4 Flowering and seed production led to subsequent seedling recruitment near transplants, mainly in recently burned sites. Genetic screening of transplants and seedlings showed that transplants in occupied sites could have crossed with nearby resident plants, but that offspring in sites previously unoccupied were likely to have been parented only by nearby transplants.
5 Seeds buried, and later exhumed, germinated after 1 or 2 years of burial, demonstrating a persistent soil seed bank from which populations could recover after fire. Neither time‐since‐fire nor H. cumulicola occupancy affected seed dormancy or germination.
6 Similar demography in unoccupied and occupied patches suggests that the patchy pattern of site occupancy by H. cumulicola is probably due to limited dispersal and periodic extinction, especially associated with long fire‐free intervals. Conservation measures need to protect unoccupied patches to allow metapopulation dynamics and persistence.     

Seed dispersal and seedling recruitment of trees at different successional stages in a temperate forest in northeastern China

Aims Spatial distribution of adult trees in a forest community is determined by patterns of both seed dispersal and seedling recruitment. The objectives of our study were to understand the processes of seed dispersal and seedling recruitment of dominant tree species in a temperate forest of northeastern China and to identify the factors constraining seed dispersal and seedling establishment at different stages of forest succession.Methods During three summer and autumn sessions between 2006 and 2008, altogether 113080 seeds from 22 different tree species were collected in three large field plots representing different forest types in the Changbai Mountain region of northeastern China. The spatial distribution of seed abundance was analyzed using a Syrjala test. Regeneration success of nine major tree species was assessed using variables defining 'limitations' in 'seeds' and 'seedling establishment'.Important findings We found that seed production fluctuated between years and varied greatly with forest types. Four tree species, Acer spp., Fraxinus mandshurica, Tilia amurensis and Betula spp., had the greatest seed production and the widest range of seed dispersal, whereas Quercus mongolica showed the most sustained seed production pattern. The spatial patterns of seed abundance differed significantly among forest types and years. The tree species investigated in this study differed in the degree of seed limitation, as well as in limitation of seedling establishment. There were both negative and positive correlations between seed density and seedling density, depending on site and parental tree density. Seeds of 16 tree species were found in the Populus davidiana–Betula platyphylla forest (PBF) plot, 11 in the conifer and broad-leaved mixed forest (CBF) plot but only 8 in the broad-leaved-Korean pine mixed forest (BKF) plot. The number of seed-contributing species was not only greater in the secondary forests (CBF and PBF plots) than in the primary forest (BKF plot) but was also more variable during the 3 years of assessment. Results from the correlations between seed density and seedling occurrence and that between parental tree density or seed weight and dispersal limitation confirm our intuitive expectations, i.e. heavy seeds had greater dispersal limitation but higher establishment success than light seeds.     

Seed to seedling transitions in successional habitats across a tropical landscape

Recognition that tree recruitment depends on the balance between seed arrival and seedling survival has led to a surge of interest in seed‐dispersal limitation and seedling‐establishment limitation in primary forests. Virtually unaddressed are comparisons of this balance in mature and early successional habitats. We assessed seed rain and seedling recruitment dynamics of tree species in primary forest, secondary forest and pasture released from grazing in a tropical agricultural landscape. Seed to seedling ratios (seed effectiveness; Φ_i) for 43 species in southern Mexico determined differences in the extent to which seeds produced seedlings by habitat, life history, and dispersal mode. Reproductive potential as estimated by the transition from seed rain to seedling recruitment, differed by habitats, and varied dramatically by life history and dispersal mode. Expected recruit densities (E_it) were higher for animal‐dispersed than wind‐dispersed species, and for non‐pioneer than pioneer species. Non‐pioneers and animal‐dispersed species had higher expected relative recruit abundance (ε_it) in primary forest (median of 4 seeds recruit^?1) whereas in secondary forest wind‐dispersed pioneers had the highest expected relative recruit abundance (median of 16 seeds per recruit). In pastures, wind‐dispersed pioneer species were most successful with many more seeds per recruit (median of 291) than both forest habitats. Seeds per recruit (Φ_i) appeared to decrease with increase in seed mass for 43 species for which data were available (r = –0.55, P < 0.001). This was associated with a negative correlation of Φ_i with seed size in primary forest (r = –0.50, P = 0.08 for 13 species); Φ_i was not correlated with seed size in secondary forest (n = 16) or pasture (n = 14). Metrics of seeds per recruit, expected recruit density and expected relative recruit abundance dramatically illustrate differences in barriers to recruitment in successional habitats.     

Colonization of experimentally created gaps along an alpine successional gradient

The colonization of artificially created gaps was analyzed along an alpine successional gradient from pioneer to early, late, and old successional stages. The presence/absence of species and the abundances of seedlings and adults in the gaps were recorded and compared with those of the surrounding areas. We hypothesized that in the older successional stages, the gaps were likely to be colonized by clonal ingrowth of the surrounding species. In the younger stages, we expected to find a high presence of seedlings and adults recruited by seeds. Micro-succession in the gaps occurred at each successional stage, with all life forms among the colonizers. The abundance of seedlings was significantly higher in the gaps compared with the surrounding area. At the early and late successional stages, the surrounding areas provided safe sites for seedling establishment, with the abundance of adults recruited by seeds higher at the gap edges than in the gap centers. We can confirm the first hypothesis of a higher clonal ingrowth in the old successional stage. Clonal ingrowth also occurred in the younger successional stages. Despite the lower species richness in the gaps, a positive correlation was found between gap and surrounding species frequencies, which were the highest in the pioneer and the lowest in the old successional stage. We conclude that gaps are relevant for seedling recruitment along the entire primary succession gradient. New species invasions from greater distances were not observed in the gaps. The dominant species on each site were identified to be successful gap colonizers.     

Niches, rather than neutrality, structure a grassland pioneer guild

Pioneer species are fast-growing, short-lived gap exploiters. They are prime candidates for neutral dynamics because they contain ecologically similar species whose low adult density is likely to cause widespread recruitment limitation, which slows competitive dynamics. However, many pioneer guilds appear to be differentiated according to seed size. In this paper, we compare predictions from a neutral model of community structure with three niche-based models in which trade-offs involving seed size form the basis of niche differentiation. We test these predictions using sowing experiments with a guild of seven pioneer species from chalk grassland. We find strong evidence for niche structure based on seed size: specifically large-seeded species produce fewer seeds but have a greater chance of establishing on a per-seed basis. Their advantage in establishment arises because there are more microsites suitable for their germination and early establishment and not directly through competition with other seedlings. In fact, seedling densities of all species were equally suppressed by the addition of competitors' seeds. By the adult stage, despite using very high sowing densities, there were no detectable effects of interspecific competition on any species. The lack of interspecific effects indicates that niche differentiation, rather than neutrality, prevails.     

Giant Sequoia (Sequoiadendron giganteum) Regeneration in Experimental Canopy Gaps

Restoration of giant sequoia populations is a high priority for managers, but few experimental studies have examined the efficacy of restoration treatments. To inform giant sequoia restoration treatment options, we assessed the response of giant sequoia regeneration (germination, mortality, and growth) to experimental gaps within a native giant sequoia grove. We created experimental gaps, ranging in size from 0.05 to 0.4 ha. Following gap creation, we sowed seeds and planted seedlings along north‐south transects across gaps. Transects were planted on paired burned and unburned substrates. The seed‐sowing treatment did not result in a cohort of established seedlings, although the amount of seeds sowed was far short of the potential amount released during intense fires. Mortality of planted seedlings did not vary with gap size (average 25% mortality). However, there was a distinct relationship between gap size and second year seedling growth. The relationship was best modeled with an asymptotic curve for both burned and unburned substrates. Relative seedling growth more than doubled as gap size increased from 0.05 to 0.2 ha, then increases in growth diminished. Growth rates of giant sequoia seedlings saturated above 70% light availability while increasing linearly with belowground resource availability. Long‐lived pioneer species such as giant sequoia require restoration treatments that involve relatively severe disturbances to facilitate cohort establishment and recruitment.     

On the adaptive value of large seeds for tropical moist forest trees: A review and synthesis

In moist temperate and tropical environments species that typically become established in closed, shaded habitats tend to have larger seeds than those that regenerate in open, secondary habitats. Despite this common pattern and the frequency with which benefits of small seed size for early successional species (large number, enhanced dispersal potential) have been discussed, little attention has focused on the advantages of large seeds for species that regenerate in closed, late successional associations. It is generally considered that large seeds enhance seedling survivorship at low light intensities. However, light intensity is only one of several factors that differ between shaded and sunlit habitats. This review examines microclimatic and biotic differences between shaded subcanopy habitats in mature tropical forests and those in sunlit, light gap habitats in which the early stages of tropical forest succession occur. Each factor is examined as a possible selective agent responsible for maintaining seed size differences between two guilds of tropical rainforest trees; the pioneer species that have small seeds and typically become established in large, sunlit gaps in the forest canopy and the persistent, relatively shade-tolerant species that have larger seeds and produce seedlings that survive for variable periods of time in the shade beneath the forest canopy. Three microclimatic factors that differ in subcanopy and gap habitats are examined; temperature, moisture, and light intensity. It is unlikely that temperature has been an important selective agent in maintaining the differences in seed sizes observed between the pioneer and persistent tree guilds. However, greater desiccation stress in light gaps might prevent successful regeneration of larger seeds in this habitat and thus might impose the smaller mean seed sizes of pioneer species. Reduced light intensities in subcanopy habitats also could favor larger seeds in the persistent guild because large seed reserves might 1) enhance the abilities of seeds to persist until suitable light (or moisture) conditions arise by providing for metabolic requirements of seeds during quiescent periods, 2) provide secondary compounds for defense of persistent seedlings against pathogens and predators during periods of low energy availability, 3) provide energy for construction of large amounts of photosynthetic tissue needed to maintain a positive net energy balance when light conditions are just above the leaf light compensation point of the plant, 4) provide energy for growth into higher light intensity strata, and 5) provide nutrients for replacement of lost or damaged tissues in persistent seedlings. Differences between soils in light gaps and subcanopy habitats are considered briefly. It is concluded that too little is known for predictions to be made regarding the probable effects of soil differences on the sizes of seeds able to survive in each habitat. Finally, differences between the two habitats in four biotic factors (competition, predation, pathogens, and mycorrhizal availability) are considered. Of these, greater competition for nutrients in the subcanopy habitat, and competition among co-germinating seedlings for light could have been important in favoring large seeds in the guild of persistent species. Pathogens are known to be more effective in shaded habitats, but data on seedling resistance to pathogens do not provide support for a role of seed size in enhancing resistance. Although differences in predation intensity and in mycorrhizal abundance in the two habitats have not been evaluated in the field, potential roles of these two factors in maintaining the seed size differential between these two guilds of forest trees are discussed. Despite the existence of numerous potential benefits of large seed reserves, seed sizes often must reflect compromises between conflicting selective pressures. Environmental conditions (e.g., moisture availability) can impose upper limits on seed size. Enhanced dispersal potential and greater total propagule numbers from maternal energy reserves are benefits of small seed size that can counterbalance selection for large seed reserves. The interactions between selective forces in molding seed sizes are discussed in a final section.     

广东南澳岛不同森林群落的林窗环境对3种移植树苗生长的影响

 先锋种和非先锋种可能对林窗产生不同的反应。在广东南澳岛4个不同的森林群落样地里移植了大叶相思（Acacia auriculaeformis A. Cunn.）、尾叶桉（Eucalyptus urophylla S. T. Blake）和荷木（Schima superba Gardn. et Champ）3种树苗,其中大叶相思和尾叶桉是先锋种,而荷木是非先锋种,测定了林窗和非林窗生境中的透光率和幼苗生长高度。结果表明,林窗的透光率大于非林窗的;各群落的透光率有季节性的变化,变化幅度各不同。3种幼苗在林窗内比     

Restoration of Araucaria Forest: The Role of Perches, Pioneer Vegetation, and Soil Fertility

The Araucaria forest is Brazil's highly threatened subtropical forest ecosystem that has been disappearing in recent decades. Restoration programs involving this forest type are scarce, and there is a lack of scientific information on how ecological processes such as competition, facilitation, and seed dispersal influence natural forest restoration. This work aims to investigate how use of perches to attract seed dispersers and the influence of pioneer vegetation and soil fertilization could affect the colonization of woody species in a degraded area. An experiment was conducted in an abandoned field where the natural establishment of seeds and seedlings of woody species was monitored under factorial combinations of the following treatments: (1) pioneer vegetation (presence and absence); (2) soil fertility (addition of NPK and control); and (3) perches (presence and absence). Seed and seedling abundance, seed and seedling species richness, and seedling mortality were recorded monthly during 12 months. Seed abundance and species richness were significantly greater in places with perches than in control plots. These results were consistent over the year and more pronounced when the surrounding forest produced a higher amount of fruit. Species richness and abundance of seedlings were significantly greater in places with perches than in control plots, and in places with vegetation than without. Soil fertility did not influence seedling establishment. Facilitation and seed dispersal are important factors affecting the colonization of woody species in this subtropical area. Nutrient availability neither regulates the facilitation process nor influences species replacement during the early stages of Araucaria forest succession.     

Fine-scale spatial heterogeneity and seed size determine early seedling survival in a desert perennial shrub (<Emphasis Type="Italic">Ericameria nauseosa:</Emphasis> Asteraceae)

In heterogenous landscapes, seeds settle in some microsites more readily than others, independently of whether those microsites are suitable for germination and subsequent growth, and survival of seedlings. Wild seeds and seedlings of Ericameria nauseosa var. oreophila in the sand-dune ecosystem near Mono Lake, California show both concordance and conflict in where seeds are retained and where seedlings flourish. Using a field experiment, we followed performance of seeds and seedlings (i.e., the rate of germination and successful emergence, initial seedling size, seedling growth and survival over first growing season), by planting seeds of known sizes in pre-identified microsites (windward and lee aspects of interspaces between shrubs, under shrub canopies, and under snag canopies, respectively). Heavier seeds were more likely to germinate and emerge successfully and resulted in initially larger seedlings than lighter seeds. These initially larger seedlings subsequently remained larger over the growing season and lived longer than initially smaller seedlings. Independent of seed and initial seedling size, seedlings that germinated under the canopy of adult shrubs grew larger and survived longer than seedlings that germinated in interspaces (open space with little or no vegetation) or under snags (dead adult shrubs). Seedlings grown under the canopy of adult shrubs experienced significantly less solar radiation and wind-deposited sand than seedlings grown in interspace or snag microsites. Sand burial more than 1 cm was lethal for seedlings. The two variables of seed size and microsite type had by far greater impact on eventual growth and survival of seedlings than did aspect, and the effects of the former two variables were independent of each other. This study amplifies the body of work on E. nauseosa at this site demonstrating that the earliest events in the life history of this long-lived perennial shrub appear to persist through time despite the harsh and variable environment.     

Seed regeneration potential of canopy gaps at early formation stage in temperate secondary forests, Northeast China

Promoting the seed regeneration potential of secondary forests undergoing gap disturbances is an important approach for achieving forest restoration and sustainable management. Seedling recruitment from seed banks strongly determines the seed regeneration potential, but the process is poorly understood in the gaps of secondary forests. The objectives of the present study were to evaluate the effects of gap size, seed availability, and environmental conditions on the seed regeneration potential in temperate secondary forests. It was found that gap formation could favor the invasion of more varieties of species in seed banks, but it also could speed up the turnover rate of seed banks leading to lower seed densities. Seeds of the dominant species, Fraxinus rhynchophylla, were transient in soil and there was a minor and discontinuous contribution of the seed bank to its seedling emergence. For Quercus mongolica, emerging seedling number was positively correlated with seed density in gaps (R?=?0.32, P<0.01), especially in medium and small gaps (<500 m(2)). Furthermore, under canopies, there was a positive correlation between seedling number and seed density of Acer mono (R?=?0.43, P<0.01). Gap formation could promote seedling emergence of two gap-dependent species (i.e., Q. mongolica and A. mono), but the contribution of seed banks to seedlings was below 10% after gap creation. Soil moisture and temperature were the restrictive factors controlling the seedling emergence from seeds in gaps and under canopies, respectively. Thus, the regeneration potential from seed banks is limited after gap formation.     

Seedling growth response to added nutrients depends on seed size in three woody genera

1 We tested whether seedlings of small‐seeded species were more reliant on soil nutrients than large‐seeded ones by growing 21 species from three woody genera ( Eucalyptus, Hakea and Banksia ) along a gradient of nutrient availability.
2 At very low nutrient availability, larger seeds produced larger seedlings. This was seen especially among the eucalypts, but the difference was eliminated at optimal soil nutrient levels. Hakea species with large seed mass, and all Banksia species, appeared unable to exploit additional soil nutrients for growth, whatever the level supplied.
3 Larger seeds tended to have proportionately higher contents of N, P and K and, under nutrient‐poor conditions, supplied more of these to their seedlings, although at a diminishing rate.
4 We suggest that large‐seededness could be an adaptation to the high‐light, nutrient‐impoverished habitats in which these species occur by providing the seedling with the mineral nutrients, rather than carbon‐based metabolites, needed for maximizing initial root growth. Reaching reliable moisture before summer (drought avoidance) is an alternative strategy to physiological tolerance of drought.     

Seed- vs. microsite-limited recruitment in a myrmecochorous herb

Dispersal in myrmecochorous herbs is frequently short-distance and this might limit patch occupancy. Here we ask whether recruitment and patch occupancy in the myrmecochere Helleborus foetidus are seed- or microsite-limited and what role dispersal plays in recruitment limitation. In pine forests from two widely separated mountain systems in the Iberian Peninsula, we performed two seed addition experiments to explore the nature of recruitment limitation in this perennial herb. In an introduction experiment, we added seeds in patches far from reproductive individuals, which rarely receive any seed. In an augmentation experiment, we added seeds in patches beneath reproductive plants. Each experiment considered three main factors: seed addition (added or not added), rodent exclusion (excluded or not excluded), and microhabitat (scrub-covered or not scrub-covered). The augmentation experiment tests whether recruitment is limited by seed availability (exploring also the possible role of rodent granivory in such limitation) while the introduction experiment tests whether the empty patches occur either because they are unsuitable for the species (microsite limitation) or due to a low probability of seed arrival (dispersal limitation). Adding seeds almost always increased seedling emergence and recruitment, demonstrating that recruitment in this species is seed-limited and refuting the possibility of microsite limitation. The comparison of the introduction and augmentation experiments indicated that Helleborus recruitment dynamics and patch occupancy are constrained by additive effects of fecundity and dispersal limitation.     

An experimental test of limits to tree establishment in Arctic tundra

1 Five treeline species had low seed germination rates and low survivorship and growth of seedlings when transplanted into Alaskan tundra. Seed germination of all species increased with experimental warming, suggesting that the present treeline may in part result from unsuccessful recruitment under cold conditions.
2 Growth, biomass and survivorship of seedlings of treeline species transplanted into tundra were largely unaffected by experimental warming. However, transplanted seedlings of three species ( Betula papyrifera , Picea glauca and Populus tremuloides ) grew more when below‐ground competition with the extant community was reduced. All three measures of transplant performance were greater in shrub tundra than in the less productive tussock or heath tundra. Establishment of trees in tundra may thus be prevented by low resource availability and competition.
3 Two species ( Alnus crispa and Populus balsamifera ) had low seed germination and survivorship of germinated seeds; transplants of these species did not respond to the manipulations and lost biomass following transplanting into tundra. Isolated populations of these two species north of the present treeline in arctic Alaska probably became established during mid‐Holocene warming rather than in recent times.
4 Of all the species studied here, Picea glauca was the most likely to invade intact upland tundra. Its seeds had the highest germination rates and it was the only species whose seedlings survived subsequently. Furthermore, transplanted seedlings of Picea glauca had relatively high survivorship and positive growth in tundra, especially in treatments that increased air temperature or nutrient availability, two factors likely to increase with climate warming.     

山西文峪河上游河岸林的土壤种子库与树种更新特征

以山西文峪河上游13种典型的河岸林为研究对象,通过土壤种子库和树种更新研究,分析群落种子库与林下更新随演替进展的变化趋势,以及该区河岸树种的繁殖对策。结果表明：13种群落的土壤种子库密度间于1290±103～3950±154粒/m2,63.5%的种子留存于0～5 cm的层次;种子库包含49种植物,以多年生草本为主,存在耐干扰种和湿地植物的种子;处于相同或相邻演替阶段的群落,种子库相似性较高;随演替进展,种子库密度、丰度、Shannon-Wiener指数及种子库与地上植被的相似性均呈降低趋势;处于演替后期的青杄Picea wilsonii林存在丰富的＂青杄幼苗库＂;先锋种白桦Betula platyphylla的种子存在于演替各阶段的群落中,储量丰富,其更新主要依赖于风媒种子,并存在少量萌蘖;青杄、白杄P.meyeri、华北落叶松Larix principis-rupprechtii、油松Pi-nus tabulaeformis和辽东栎Quercus liaotongensis的种子库损耗严重,没有或仅存少量种子,其中云杉和油松的更新幼苗幼树多,属持久幼苗库更新;华北落叶松幼苗幼树少,且仅出现于林缘或林窗等开阔地,属植被空隙中季节性更新;辽东栎主要依赖丰富的幼苗库进行更新,同时存在一定的萌蘖;青杨Populus cathayana以大量风媒种子更新结合营养扩展。     

Predispersal seed predation on five Piper species in tropical rainforest

Absolute number of seeds lost to predispersal seed predators and proportion of total seeds lost per infructescence were compared among five Costa Rican Piper species of different annual fecundities. Mean seed number and mean seed size in the five species were negatively correlated. The impact of predation on these species was inversely related to the number of seeds they produced. The two early successional species had very high fecundities, a combination of many seeds per infructescence, many infructescences per plant, and, in one species, year-round reproduction. Although seed predators destroyed as many or more seeds of these early successional species than they did of the less fecund, late successional species, this loss accounted for a relatively minor proportion (9 and 12%) of the seeds of the early successional species. In contrast, late successional species produced fewer, larger seeds in a smaller number of infructescences and were not continually in fruit. One of these species, which produced intermediate numbers of intermediately sized seeds, lost 30% of the seeds in each infructescence on average. Seed predators destroyed a larger proportion (65 and 76%) of the seeds per infructescence in the two species with fewest seeds per infructescence. High levels of insect damage in these late successional species caused many of their infructescences to abort prematurely. Taken together these factors resulted in annual fecundities several orders of magnitude smaller in shade-tolerant Piper species than the annual fecundities of shade-intolerant, early successional species. Seedlings of the two early successional species were common in large gaps and other sunny clearings and seedlings of the species with 30% seed loss were occasional, whereas no seedlings were seen of the two species with the highest proportional seed loss, suggesting that seed predation on the latter species may limit seedling recruitment.     

Impacts of secondary seed dispersal and herbivory on seedling survival in Aesculus turbinata

Abstract. Aesculus turbinata is a tree species with large seeds (6.2 g mean dry weight). We studied the demography of its seeds and seedlings in a temperate deciduous forest in northern Japan to elucidate the ecological significance of large seeds with special reference to herbivory and secondary dispersal. Both seed and seedling stages suffered greatly from herbivores. Seedling herbivory was important judged from experiments with shoot clipping and hypogeal cotyledon removal. However, some seedlings survived through re-sprouting after herbivory. Survival rate and percentage resprouting seedlings were lower than those with remaining cotyledons, though seedling size was not affected. This suggests that stored resources in hypogeal cotyledons are working as a kind of ‘risk hedge’ against severe aboveground shoot clipping experienced by A. turbinata. The spatial distribution of seedlings was expanded via seed scatter-hoarding by rodents. Seedling survival rate was higher within canopy gaps than under closed canopy, indicating that canopy gaps are safe sites for establishment, and was negatively correlated with seedling density. Therefore, secondary seed dispersal in this species seems to be effective in ‘finding’ safe sites and in ‘escaping’ density-dependent mortality. The large seeds and seedlings of A. turbinata are attractive to herbivores, but the high resistance of seedlings to herbivory due to large reserves and the effective secondary dispersal appear to mitigate these disadvantages.     

Effects of Sward Height, Gap Size, and Slug Grazing on Emergence and Establishment of Trollius europaeus (Globeflower)

Abstract To determine the effects of sward height and gap size on the emergence and subsequent development of Trollius europaeus, seeds were introduced into artificially created gaps in mown and unmown wet meadow grassland in Ayr, Scotland. Emergence and establishment of nondormant seed (previously leached in running tap water and soaked in gibberellic acid) placed in a range of circular gaps (25 mm, 50 mm with and without root barrier, 100 and 200 mm diameter) were monitored over 305 days. Trollius demonstrated high field emergence (mean, 46% of viable seed sown) in this experiment. Emergence was not significantly affected by either sward height or gap size, although emergence was greater in mown swards. The critical factor determining subsequent survival of seedlings was slug predation rather than sward height or gap size. However, losses due to slug predation were significantly greater in unmown swards, resulting in more seedlings in mown swards. One hundred eleven days after sowing (September) seedling numbers had declined substantially in both unmown and mown swards, and by day 305 (April) seedlings were extinct in both sward types. The implications of the research for the restoration and management of T. europaeus are discussed.     

Seedling recruitment of Succisella inflexa in fen meadows: Importance of seed and microsite availability

Abstract. The effects of different forms of land use on germination and establishment of the rare fen species Succisella inflexa were investigated in seed introduction experiments in a mown and an abandoned fen meadow in SE Germany. Treatments included abandonment, mowing in fall and mowing with creation of gaps in the moss and litter layer. Floating capacity of seeds was tested in order to estimate potential dispersal by water. On the mown meadow, gaps had a slightly positive effect on germination rates, while greatly increasing seedling survival until the next spring. At the abandoned site, litter inhibited germination, whereas mosses had a negative effect on germination and a positive effect on survival rates during the first year after germination. Both germination and seedling establishment were negatively affected by the presence of slug herbivores. On the abandoned site, no seedlings at all survived until the next spring. Even though seeds of Succisella inflexa were capable to float for several weeks and to germinate thereafter, the situation at the field sites indicates that longdistance dispersal is highly unlikely. Our results showed that not only direct effects of abandonment, such as accumulation of litter, may have led to poor germination and poor seedling establishment of the species. Additionally, indirect consequences of changes in land use, such as higher seedling herbivory by slugs and successional vegetation changes due to abandonment, were important in determining habitat quality and availability of microsites for seedling recruitment. Furthermore, early mowing imposed seed limitation on plant populations.