Seed dispersal phenology and dispersal syndromes in a subtropical broad-leaved forest of China

This study describes the dispersal phenology and syndromes in Gutianshan 24 ha plot in a subtropical broad-leaved forest of China. The 130 0.5 m² seed traps collected 69,115 mature seeds, representing 27 species (belonging to 24 genera, and 15 families) in 12 months. One marked peak in the number of seeds and species during the year was found in dry season (November). Zoochory was the most common dispersal syndrome (70.4%), followed by anemochory (18.5%), ballistic dispersal (11.1%). Among fruit types, berry (33%), capsule (22%), nut (18%), and drupe (11%) were common in the subtropical evergreen forest. In fruit color, brown was the commonest (40%), followed by dark brown (30%), black (15%), red (11%), and yellow (4%). Overall, the community level seed rain study revealed that one marked peak in seed number occurred in the middle of dry season; zoochory was the principal dispersal mode of woody plants in subtropical forest, and dry seasons favor seed dispersal by animal and wind.     

Effects of bamboo stands on seed rain and seed limitation in a rainforest

Bamboos often negatively affect tree recruitment, survival, and growth, leading to arrested tree regeneration in forested habitats. Studies so far have focused on the effects of bamboos on the performance of seedlings and saplings, but the influence of bamboos on forest dynamics may start very early in the forest regeneration process by altering seed rain patterns. We tested the prediction that the density and composition of the seed rain are altered and seed limitation is higher in stands of Guadua tagoara (B or bamboo stands), a large-sized woody bamboo native from the Brazilian Atlantic Forest, compared to forest patches without bamboos (NB or non-bamboo stands). Forty 1 m² seed traps were set in B and NB stands, and the seed rain was monitored monthly for 1 year. The seed rain was not greatly altered by the presence of bamboos: rarefied seed species richness was higher for B stands, patterns of dominance and density of seeds were similar between stands, and differences in overall composition were slight. Seed limitation, however, was greater at B stands, likely as a resulted of reduced tree density. Despite such reduced density, the presence of trees growing amidst and over the bamboos seems to play a key role in keeping the seeds falling in B stands because they serve as food sources for frugivores or simply as perches for them. The loss of such trees may lead to enhanced seed limitation, contributing ultimately to the self-perpetuating bamboo disturbance cycle.     

Spatial Pattern of Bird-Dispersed Seed Rain ofDaphniphyllum macropodum in an Evergreen Broad-Leaved Forest

Abstract

We investigated how the locations of conspecific fruiting trees, other fruiting species, and canopy gaps affected spatial patterns of bird-dispersed seed rain for a common ornithochorous tree: Daphniphyllum macropodum.In a mature evergreen broad-leaved forest in southwestern Japan, seed rain was monitored with seed traps in a 1.2 ha plot on a site with a generally closed canopy. Many of the bird-dispersed seeds of D. macropodumwere concentrated near the conspecific fruiting trees, while other seeds were scatter-dispersed. The spatial distribution of scatter-dispersed seeds was independent of the distance from conspecific fruiting trees. The scatter-dispersed seed rain of D. macropodumwas biased by abundant and widely distributed fruiting trees of Cleyerajaponica.Other rare fruiting species and canopy gaps had no clear effect on the seed rain of D. macropodum.Results show that the spatial pattern of bird-dispersed seed rain in this forest is affected by the location of fruiting trees of the same and different ornithochorous species rather than by canopy gaps.     

动物取食和扩散森林植物种实研究进展

动物取食扩散森林植物种实作为森林生态系统中植物和动物联系的-个重要方面, 不仅是联系母株与其子代幼苗的重要桥梁, 也是植物生活史中的-个重要阶段。文中综述了近年来关于动物取食森林植物种实及其与这些植物种子散布、萌发、幼苗生长和种群扩展等方面关系的研究成果。动物在取食和破坏森林植物种实的同时也将这些植物的种子扩散到-个新的生境斑块, 其消化道机械的或化学的作用提高了这些森林植物的种子萌发率和萌发进程, 促进了其早期定居和种群扩大。动物对森林植物的这种作用具有长期性, 形成特定动植物间或多物种间的协同进化。近年来, 森林生态系统受到了严重的干扰和破坏, 可能会因为干扰了森林植物种实的动物散布者而影响森林植物种群的扩展。     

Seed dispersal networks in an urban novel ecosystem

The conflict resulting from the expansion of human activities into natural habitats affects the structure and functioning of ecosystems. Similarly, the anthropogenic redistribution of many species all over the world affects the composition of biological communities, possibly altering their capacity to sustain key ecological functions, such as seed dispersal. Urban parks are extreme examples of such novel ecosystems resulting from the anthropogenic redistribution of species in a new ecological framework. Here, we describe the avian seed dispersal network in an urban recreational woodland in central Portugal (c.79 ha). Four quantitative seed dispersal networks were assembled by identifying intact seeds in the droppings of mist-netted birds throughout the year. Overall, 1,244 seeds were identified, representing 33 links between 15 plant species and 11 bird species. Most birds dispersed alien seeds, but these represented a small proportion of the overall network (20 % of the seeds and 13 % of the droppings). Blackcap (Sylvia atricapilla) was the main disperser of native and alien seeds in all seasons, particularly those of the invasive Phytolacca americana. Fleshy fruits were more abundant in summer, but were more consumed in winter, presumably when other foods were difficult to find. Our study suggests that even in a highly managed urban park, seed dispersal networks can be temporally complex and variable and that a network approach can be an important monitoring tool to detect the status of crucial ecosystem functions in rapidly changing habitats such as urban parks.     

Intra and inter-annual variation in seed rain in a secondary dry tropical forest excluded from chronic disturbance

Domestic livestock grazing and firewood extraction are chronic and pervasive disturbances in dry secondary forest that reduce diversity and modify composition. Exclosure of such disturbances may allow the reestablishment of original diversity. To investigate potential vegetation recovery following 30 years of perturbation, seed rain was evaluated monthly during 3 years in 96 0.78 m² seed traps located in eight exclosures established in 2006 in a species-poor secondary dry forest in central Mexico. Twelve additional seed traps were located outside the exclosures in 2010 for comparison purposes. Baseline diversity of seed sources (trees, ?5 cm DBH) was recorded to compare standing tree composition with seed rain. Also, baseline diversity of potential primary (birds) and secondary (ants) dispersal vectors were registered to infer current plant-animal interactions. Total seed rain over 3 years within the exclosures comprised 180,375 seeds from 61 woody species, an average of 1.12 seeds/m²/month. Half of these species (31) and 95% of the seeds falling within the exclosures belonged to trees typical of perturbed sites. Sixty percent of all recorded seeds belonged to an early-successional, bird-dispersed shrub (Hamelia patens Jacq., Rubiaceae). Seed rain at one non-excluded plot was dominated by the wind-dispersed early-successional tree Heliocarpus sp. (Tiliaceae). Richness and seed density increased significantly in the second year of exclusion; however, by the third year, richness remained constant while seed density decreased. Seed rain was richer and more abundant during the dry season and was dominated by wind-dispersed species. Phenology, successional status and the dispersal modes of woody species explained most variance in richness and seed density of seed rain in early-successional dry forest. Seeds of early-successional species were dispersed in equal measure by abiotic and biotic vectors, whereas biotic vectors dispersed a higher density of late-successional seeds. The amount and timing of fall of abiotically dispersed seeds of the three most abundant species in the standing tree community may prolong their current domination of the landscape. However, even when richness in the seed rain within exclosures remained constant through time, composition was gradually shifting towards animal-dispersed species which may start a positive feedback producing an increase in animal visitation and a further increase of richness.     

缙云山常绿阔叶林种子雨组成及其时空动态

【目的】分析缙云山常绿阔叶林种子雨组成及其时空动态,为常绿阔叶林种子生态学大尺度和长时序的研究提供数据支持,并为受损常绿阔叶林近自然恢复过程中的种源收集和种子撒播提供参考信息。【方法】 2014-01—2016-12,收集缙云山常绿阔叶林1 hm 2标准地中172个种子收集框内的种子,鉴定物种并计数,风干后称质量。根据果实类型及种子特征并结合野外实地观察,确定每个物种种子(果实)的散布方式。【结果】 3年间共收集到成熟种子36 298粒,种子雨密度为140.69粒·m -2 a -1;种子分属于19科21属27种,占群落木本植物物种总数的36.49%;种子雨具有明显的季节动态,在秋季出现高峰,其中有17个物种在干季(1—4和10—12月)成熟并散布,6个物种在湿季(5—9月)成熟并散布,4个物种在干、湿季均有散布;种子雨密度年际变异系数为0.36~1.73,空间变异系数为1.54~15.20。【结论】缙云山种子雨产量在常绿阔叶林中处于较低水平;优势种对群落种子雨产量贡献较大,种子雨物种组成与群落物种组成相似性较低;群落水平种子雨具有明显的季节动态,且存在很大的年际变异和空间变异,反映出亚热带常绿阔叶林种子雨时空动态的典型特征。在区域植被近自然恢复中的种子采集和撒播环节,应当充分参照不同物种的种子雨掉落的时空动态。     

Negative effect of removing pulp from unripe fleshy fruits: seed germination pattern of Celtis sinensis in relation to the temporal context of fruit consumption

We explored the temporal effects of fruit consumption on the subsequent seed germination pattern of a fleshy-fruited tree, the Chinese hackberry (Celtis sinensis). Via nursery-based sowing experiments, we investigated (1) how pulp removal affected seed germination patterns at the different stages of fruit maturation, and (2) how the timing of seed dispersal (August, October, and December) affected the germination patterns of seeds from ripe fruits after the removal of pulp. We found that the removal of pulp from around the seeds of ripe fruit had no effect on the percentage and timing of germination. In contrast, the removal of pulp from seeds of unripe fruits largely reduced the germination percentage. The time of sowing did not affect the germination percentage, whereas the timing of germination was delayed for seeds that were sown later or under shaded environments.     

Grazing exclosure increases fruit and seed production but reduces seed viability in <Emphasis Type="Italic">Myrtus communis</Emphasis>

Plants have compensating mechanisms to alleviate the loss of vegetative and reproductive tissues by herbivores. In this study, we evaluated how reproductive traits (size and production of fruit and seed, as well as germination and viability of seeds) of Myrtus communis (myrtle) were affected by goat grazing exclosure of forest understory in the Doñana Natural Park. One hundred and fifty myrtles were selected in three sites with similar environmental characteristics, although with a different length of grazing exclusion: (1) exclosure for 2 years (GE2); (2) exclosure for 3 years (GE3); and (3) long ago grazed plots (GE15): non-accessible to goat for at least 15 years. Fruit production, fruit and seed morphology and seed viability and germination were evaluated. Length of exclosure to goats increased fruit production, mass and volume of the fruits and seed, and endocarp mass. However, seed viability was higher in more recently grazed plots (70–90%) than in GE15 plots (51%). The final balance of viable/germinable seeds produced by an average sized myrtle in each treatment showed that the recently grazed myrtles (GE2) produced nearly five times less seeds (1200) than the myrtles grazed 15 years ago (5500). However, myrtles recovered the ability to produce viable seeds quickly, almost tripling, each year of non-grazing the number of viable/germinable seeds. The results of this study indicated that goat grazing could affect regeneration of myrtle because it reduced reproduction output and the number of viable propagules. Future studies should evaluate other related ecological aspects, such as fruit consumption by potential frugivores and seed dispersal (distance and place of seed deposition).     

Colonization of fleshy-fruited plants beneath perch plant species that bear fleshy fruit

We evaluated the contribution of seed dispersal by birds to the colonization of fleshy-fruited plants beneath plants that bear fleshy fruits and were used as perches by birds. Specifically, we investigated the composition of seeds and saplings sampled beneath nine fleshy-fruited perch plant species in a Pinus thunbergii stand on a coastal sand dune in central Japan. Both the numbers and the species numbers of bird-dispersed (pulpless) seeds differed significantly among the perch plant species. Nevertheless, the quantitative compositions of bird-dispersed seeds were similar among the perch plant species because of the significant positive interspecific correlations of species rank order in numbers of bird-dispersed seeds per species. Similar relationships were also found for saplings. For each perch plant species, the numbers of fallen conspecific seeds were significantly higher than or not different from those of heterospecific seeds. Furthermore, the numbers of pulpy seeds that fell directly from the perch plant were significantly higher than those of bird-dispersed conspecific seeds for seven of nine perch plant species. However, the densities of conspecific saplings were significantly lower than those of heterospecific saplings. Most pulpy seeds, therefore, probably do not germinate beneath their mother plants, and conspecific seeds probably accumulate in seed banks or are consumed by herbivores.     

Long-lasting effects of unplanned logging on the seed rain of mixed conifer-hardwood forests in southern South America

Understanding the effects of disturbance on seed rain is critical to predict changes in forest species composition and diversity.Logging effects on seed rain in a mixed conifer-hardwood forest complex in southern Brazil were evaluated.One year of seed rain data were collected from a large-scale observational experiment in logged and protected forests and quantity and average seed size weighted by species abundance(CWM) were compared between old-growth and logged stands 55 years after logging activities.Using these data,variations in frequency of functional groups of species in the seed rain were examined to see if they could be attributed to logging.Results show that the number of seeds per trap was highly right-skewed,ranging from 13 to 12,788 seeds per trap in one year.Seed rain was affected by logging history,with seed traps in old-growth plots receiving significantly less seeds than traps in logged plots.All species included mean seed size weighted by species abundance were significantly smaller in logged than in old-growth forests.This difference persisted after the exclusion of Arawcaria a large-seeded pioneer which was intensively logged,although the difference of seed size between the two forest classes was greatly reduced.Species abundance in the seed rain differed significantly from the established tree community,between logged and oldgrowth stands.The composition of the seed rain was much more variable than the composition of the established tree community and its points more scattered over the ordination space than the points corresponding to the protected forests.The number of collected seeds across different functional groups of species significantly differed between logged and old-growth plots.The seed rain of logged forests reflects their arrested succes sion as indicated by reduced abundance of functional groups such as pioneers,large seeded pioneers and A raucaria,as well as reduced functional diversity.S eed rain differences between logged and old-growth stands reflect the intensive logging of A.angustifolia.     

A seed rain community in a reforested post-agricultural field and adjacent secondary forest of Mount Papandayan Nature Reserve,West Java,Indonesia

Understanding the mechanisms and barriers to the restoration of degraded land, especially post agriculture, will help provide protocols on effective ways of restoration into functional ecosystems. One of the barriers in early stages of forest restoration is the arrival and availability of propagules. Seed rain and factors affecting it(i.e., distance to the forest edge, species diversity and surrounding vegetation) were measured in a reforested post-agricultural field and in an adjacent secondary forest. Multivariate g lm analysis was used on the seed rain community data and univariate lm analysis on the most abundant seed captured(i.e., Schima wallichii(DC.) Korth.). After 8 months of seed rain collection, there was a total of 3596 seeds from eight tree species. Seeds were more abundant and more diverse in the secondary forest(74.9%, 8 species) compared to the reforested field(24.1%, 2 species). There was a limitation on seed dispersal in reforested field from the adjacent forest. The abundance of S. wallichii seeds determined by the dominance of adult trees(136 trees/ha in the forest and 115 trees/ha in the reforested field). Our study suggests, that after 7 years of planting, the reforested field has received limited seed rain and has not yet recovered.     

Exotic and invasive species compromise the seed bank and seed rain dynamics in forests undergoing restoration at urban regions

The control of exotic and invasive species in areas undergoing recovery is a challenge for ecological restoration and this problem may be even greater in urbanized areas. This study evaluated the seed bank and seed rain of a 5-year-old riparian forest located at the metropolitan region of Belo Horizonte in southeast Brazil, and verified changes over time in the abundance and species richness and whether planted species already were providing propagules to the site. At the forest, fifteen parcels of 100 m² were distributed randomly and soil samples were collected thrice (October 2011, March 2012 and July 2012) to evaluate the seed bank. Seed traps were installed in nine parcels to collect propagules over 1 year. Propagules were segregated and germinate at greenhouse. Species richness and abundance were estimated, and variance and similarity were quantified. In total, 86 species were recorded in the seed banks, 41.9% natives and 33.7% exotics, with the predominance of herbs (87.2%) and only 7% of trees. Species classified as weeds amounted to 52.3%. In the seed rain 642 seedlings germinated but they belonged to only 10 species, half of them were exotics. Regarding life forms, half of the species were arboreal and dominated the samples, highlighting the exotic species Melia azedarach. The species richness of the seed bank did not differ by season, unlike the seed rain which varied significantly by month. Seed bank and seed rain species had a very low floristic similarity with the list of planted species, suggesting that few planted trees had dispersed or were stocking propagules. Many non-planted species were recorded, mostly exotics and/or invasive that were probably derived from the surrounding urban matrix. These might interrupt the successional dynamics of the restoration forest and compromise the restoration process in the medium and long-terms. Our results highlight: (1) the importance of monitoring areas undergoing restoration; (2) that restoration methods and management actions directed to control exotic and invasive species are particularly important for sites in urban regions; and (3) the relevance of conserving forest fragments as sources of propagules for nearby areas.     

Seed supply and the regeneration potential for plantations and shrubland in southern China

Assessing the characteristics of seed supply will be vital to better understand the dynamics of forest regeneration. In this study, we surveyed the aboveground vegetation, the seed rain, the seed bank, and natural seedling emergence in four typical 24-year-old plantations (eucalyptus, mixed-native, mixed-legume, and mixed-conifer) and a naturally successioned shrubland in southern China. The dominant species in the understory were similar among the five plant communities. The seed rain and the seed bank were dominated by shrubs and herbs but indigenous tree species were rare. Species that were common to all five-plant communities represented a great proportion of the seeds in the seed rain and seed bank. The seed rain consisted mostly of seeds derived from the local plant community. Seed abundance was greater in the seed bank than in the seed rain, and species richness was greater in the seed bank and in the corresponding plant community than in the seed rain. Species composition similarity between the seed rain, the seed bank, and the aboveground vegetation was low, because the seed rain contained much fewer species, and the seed bank and aboveground vegetation contained many different species, respectively. These findings indicate that both the seed rain and the seed bank play important roles in providing seeds for plant recruitment in the understory, but the seed bank contributes more than the current seed rain to the diversity of recruited plants. The current plant community has little impact on the qualitative composition of the seed rain and seed bank. Based on these data, it appears that succession to the desired zonal, mature forest community is unlikely to result from seeds in the seed rain or seed bank. Lack of seed availability of desired zonal mature forest species is the main bottleneck currently limiting succession in the plantations. Reintroduction of late-successional species could facilitate the desired succession.     

Spatial variation of seed rain and seed banks in gaps of karst forest in the Maolan Nature Reserve, Guizhou Province

Based on an investigation on gaps and non-gap stands of the Maolan National Karst Forest Nature Reserve, Guizhou Province, quantitative characteristics and dynamic changes of seed rain and seed banks in gaps were analyzed. The results show that the total amount of seed rain was 117.4 ± 32.6 seeds/m² during the period of observation. The number of immature seeds was 56.3 ± 10.3 seeds/m², that of mature damaged seeds was 15.7 ± 4.7 seeds/m², and the number of mature germinated seeds was 45.4 ± 8.2 seeds/m². It is suggested that the seed number is rich for gap regeneration. Seed rain in gaps has spatial and temporal heterogeneities which deeply affect regeneration patterns of gap plants. Along a gradient from the gap center to a non-gap stand, seed density in the litter layer, the number of species, and the Shannon-Wiener diversity index were gradually reduced, but these indices increased in the soil. The seed density in the gap center was 2415 ± 639 seeds/m², near the gap center was 2218 ± 421 seeds/m² and at the gap border area 1815 ± 311 seeds/m². This shows that plants in gaps have good latent regeneration potential. In both gaps and non-gap stands, the Jaccard similarity index of seed in litter layer was the largest, second largest at 5–10 cm soil depth, and the least at the 0–5 cm soil layer the index. The Jaccard index between the soil seed bank and the present plant community was large in the litter layer, but decreased with soil depth both in gaps and non-gap stands. The results show that soil seed banks are the main source of gap regeneration in the karst forests of Maolan and contribute significantly to gap regeneration. __________ Translated from Acta Botanica Yunnanica, 2007, 29(3): 327–332 [译自: 云南植物研究]     

Seed production in Polylepis australis (Rosaceae) as influenced by tree size, livestock and interannual climate variations in the mountains of central Argentina

The expansion of tree species within a grassland-woodland matrix relies heavily on an abundant seed crop, which may be affected by many factors; among these, browsing by livestock has gone largely ignored. Here we explore how seed production is affected by tree size, grazing by large domestic herbivores and interannual climate fluctuations in a small-sized tree (Polylepis australis, Rosaceae), which dominates the remnant forests in the high mountains of central Argentina. Seed number and seed mass per tree were quantified in 100 different-sized trees located in areas with and without livestock. Using seed traps we also determined seed rain variation in 15 plots between 2005 and 2008. The estimated number of seeds per tree ranged from 0 to 224,000 and increased with tree height (power relation); for trees smaller than 354 cm in height, seed production was lower in areas with livestock and higher in areas without livestock. Mass per seed increased with tree height, with no significant livestock effect. The largest seed rain was recorded in 2008, probably due to growth of seed trees and not in response to climate. To promote seed production and forest expansion, livestock should be excluded from areas with small trees, such as forest edges.     

海南尖峰岭具有较高园林开发价值的5种野生植物介绍

从观赏特征、形态、生境、分布、应用特点等方面介绍海南尖峰岭的海南梧桐Firmiana hainanensis Kosterm、方枝蒲桃Syzygium tephrodes(Hance)Merr.&L.M.Perry、拟赤杨Alniphyllum fortunei(Hemsl.)Makino、显脉杜英Elaeocarpus dubius A.DC.和银珠Peltophorum tonkinensis(Pierre)Gagnep.5种适宜在南方园林中应用推广的野生植物,以期为乡土园林植物的开发利用提供一些新的资料,并引起相关工作者对乡土观赏植物的重视,促进乡土植物在南方园林中的应用和推广。     

Floristic and structural recovery of a laurel forest community after clear-cutting: A 60 years chronosequence on La Palma (Canary Islands)

We analyzed a post-clearcut chronosequence (0.5 to 60 years after harvesting) in the laurel forest of La Palma island (Canarian Archipelago) to determine the recovery of the stands with respect to species composition, richness, life strategies and structural parameters of the canopy. Multivariate analysis showed that exotic species, as well as annual ruderal species were confined to early-successional stages, while native perennials, typical of laurel forests, dominated the late-successional stages. Total species richness decreased significantly with time after clear-cutting. The relative fast recovery of understory native species may be due to low forest floor disturbance during harvesting. Shade-intolerant pioneer, pioneer-remnant and shade-tolerant late-successional species were the main life strategies of native tree species. Most structural parameters showed a continuous and monotonic increase (basal area, biomass) or decrease (density, percentage of photosynthetic biomass) during succession. Once clear-cutting, here performed with an interval of 8 years, is abandoned, the recovery of the laurel forest seems possible due to careful logging that protects the soil and a rapid asexual regeneration of native tree species, revealing this to be a sustainable management practice.     

浙江天童常绿阔叶林米槠种子雨和地表种子库动态研究

2008年在浙江天童山常绿阔叶林的林缘和林内2个米槠种群中各选择3株米槠(Castanopsis carlesii),采用种子筐及地表取样等方法,分析了米槠种子雨和地表种子库动态特征、种子存留及其命运。结果表明:米槠种子雨主要出现在10月中旬—11月,持续约50 d,11月3日—17日为高峰期;该地区米槠的种子产量较高,林缘种群高于林内种群,单株平均落果数分别为201.0、120.3个.m-2;种子雨中完好的坚果占52.2%,虫蛀的占29.3%,松鼠啃食的占10.2%,败育的占8.4%,腐烂的占5%;完好种子的长度、宽度分别为(9.01±0.89)mm、(9.25±1.00)mm,粒重为(0.47±0.13)g;在地表种子库中,林缘种群、林内种群完好种子的存留密度分别为1.5、1.0个.m-2,分别占种子库的62.1%、66.4%,动物捕食是种子受损的主要原因。米槠种群较高的种子产量、较低的地表种子存留率等环节限制了米槠种群的天然更新。     

Variation in seed rain from Widdringtonia whytei growing in different conditions on Mulanje Mountain in Malawi

Mulanje Mountain is one of the examples of tropical montane landscapes with plant populations varying in size from small fragments of less than 1 ha to riverine strips and stands of more than 100 ha. Seed availability is a potential limiting factor in vegetation recovery in such landscapes. Field observation on seed rain and seed limitation was conducted in 2008 and 2009 in three sizes of forest fragments at three sites on the mountain to investigate: (1) whether size of Widdringtonia whytei forest patches influences seed-rain density; (2) whether W. whytei seed rain occurs in a specific season; and (3) if seed limitation differs from within forest patches to the adjacent non-forest patch habitats. The results showed that there was a distinct difference in seed-rain density among fragments where large fragments (>20 trees) collected higher seed-rain density (10.4 ± 0.9 seeds m^?2 y^?1) than small fragments (1.1 ± 0.4 seeds m^?2 y^?1). The seed-rain density differed significantly between 2008 (2.1 ± 0.6 seeds m^?2 y^?1)and 2009 (6.1 ± 1.1 seeds m^?2 y^?1). Seed rain was recorded every month of the year but the densities were not significantly different among months. Seed-rain density was highest inside the forest fragments (12.04 ± 2.3 seeds m^?2 y^?1) and decreased drastically and very significantly at the edges (1.0 ± 0.8 seed m^?2 y^?1) and outside the forest fragments (0.11 ±0.1 seed m^?2 y^?1). Seed limitation was strong (0.98) on all sites; 0.95 in large fragments and 0.99 in medium and small fragments, respectively. Therefore, it can be concluded from this study that the presence and retention of more seed trees ensures the continued availability of W. whytei seed on the forest floor within the context of limited seed dispersal.