胡杨种子散布的时空分布格局

以额济纳胡杨为研究对象,对种子雨的散布时间、强度、散布距离以及种子雨和空气湿度、风之间的关系进行了研究。胡杨种子雨可以分为初始期、高峰期和消退期3个阶段,大部分的种子集中在高峰期落下。种子的散布主要受湿度和风的影响。湿度对种子雨的强度起主要作用,在一天之中,种子在湿度较低的中午和下午集中散落。应用一元线性回归模型对种子雨强度和相对湿度进行分析后表明二者之间存在显著的负相关关系。对种子的散播距离进行研究后发现,大部分种子落在母树附近,少部分种子能够进行长距离传播。风对种子的传播的方向和距离起决定性的作用,不同方向上的种子传播距离和强度相差很大。在顺风方向上,种子的传播距离最远,所有的长距离传播现象几乎都发生在这一方向上;而在主风向的垂直方向和逆风方向上,种子的散布距离较小,很少有种子能够进行长距离传播。对风的观测表明中午后和下午初的风力较强,而此时种子雨强度又最大,有利风力条件和高种子雨强度出现的同步性可能是促进胡杨种子进行长距离传播最有效的生物控制机制。由于胡杨种子在自然条件下的存活时间非常短暂,所以研究中不同胡杨母树林间种子散播时间的差异可能是胡杨种群内部为适应不同洪水期所表现出的风险分摊机制所造成的。     

呼伦贝尔沙地天然樟子松种子扩散机制研究

通过对沙地樟子松母树种子落种时空格局的观测,发现沙地樟子松种子大部分在高峰期集中飞散在母树周边,与母树距离的增加种子数量减少,在距离母树4m左右处,落种密度达到最大,且母树北侧和东侧的落种密度要高于南侧和西侧.此外文章还探讨了落种密度和距离与母树本身的关系,如树高和冠幅等,推算出了树高和扩散距离之间的量化关系,计算得出在已知风速的情况下沙地樟子松种子的最大扩散距离公式.最后将沙地樟子松种子更新过程分为母树、种子和幼苗阶段3个阶段和母树的结实与生产、种实扩散、种子萌发和幼树生长4个时期,从环境状况和母树自身特征角度分别探讨了沙地樟子松更新的影响因素,并在此基础上建立了沙地樟子松种子更新的时空模型.     

松嫩平原碱化草甸朝鲜碱茅种子散布机制的分析

采用顺序远离母株取样的方法,定量分析了松嫩平原碱化草甸朝鲜碱茅（Ｐｕｃｃｉｎｅｌｌｉａｃｈｉｎａｍ ｐｏｅｎｓｉｓ Ｏｈｗ ｉ）的种子散布机制。结果表明,在８个方向中仅西南方向表现为近母株散布距离,其它７个方向均表现为远母株散布距离的种子散布机制。经相关性分析,朝鲜碱茅向不同的方向散布的种子数量与种子散布时期顺风向的风速和风向频率呈正相关,反映了朝鲜碱茅是借助于风媒介实现了远母株距离扩展潜在生态位空间的种群对策。朝鲜碱茅向母株周围散布的种子呈连续分布格局的散布半径平均为１７５ ｃｍ 。     

神农架巴山冷杉种子雨的时空格局

巴山冷杉(Abies fargesii)是我国特有树种, 其分布区以秦巴山地为中心。在湖北神农架自然保护区选择不同的巴山冷杉群落设置样地, 通过布设种子收集器结合室内实验分析, 对巴山冷杉的种子雨进行研究。各样地巴山冷杉种子雨开始和持续时间大致相同, 始于10月初, 持续超过2个月, 但是不同的巴山冷杉种群表现出不同的种子雨时空格局。巴山冷杉－箭竹(Fargesia nitida) 群落种子雨的平均强度为167.93±111.14粒/m2, 有活力种子比例占22.31%, 落种高峰期集中于10月27日到11月2日间, 种子雨呈现聚集分布。巴山冷杉－茵芋(Skimmia reevesiana)群落的种子雨强度在3个样地中最小, 只有16.41±14.41粒/m2, 有活力种子仅占3.05%。对巴山冷杉－陕甘花楸(Sorbus koehneana)群落, 沿等高线方向和垂直等高线方向布置收集器带, 收集器自群落内中心母树延伸至林冠范围之外(距中心母树的距离>18 m)。收集结果表明种子雨落种高峰为10月15–21日, 种子雨主要集中在林冠范围内, 林冠内种子雨数量占到了收集总数的87.95%。方差分析结果显示各方向的种子雨数量不存在显著差异, 按照远离中心母树的距离进行分段比较, 0–6 m区段内的收集数量和其他各区段间(6–12 m, 12–18 m, >18 m)存在显著差异, 扩散的种子数量与离开中心母树的距离间的关系近似正态分布。研究结果说明巴山冷杉种子雨的强度大但种子质量不高, 而且种子雨的扩散距离有限。这些特征将影响到巴山冷杉种群的分布格局和种群更新。     

湖北三峡大老岭自然保护区光叶水青冈群落种子雨10年观测: 种子雨密度、物种构成及其与群落的关系

种子雨是植物种子扩散的起点, 对群落更新及种群动态起着关键作用。该文以三峡大老岭自然保护区内一片面积为1.3 hm ²的光叶水青冈(Fagus lucida)群落固定样地为研究对象, 运用分层随机设计, 在10个不同的地形部位放置了100个种子雨收集框, 自2001年起进行种子雨观测, 对该群落种子雨的数量与物种多样性的年际动态、种子雨和群落物种构成的关系等进行了统计分析。结果表明: 1)过去10年间, 共收集到来自48种木本植物的60 926粒种子, 种子雨的多年平均密度为(82.9 ± 61.5) seeds·m^-2·a^-1(mean ± SD), 平均物种丰富度为(16.7 ± 5.5) species·a^-1(mean ± SD)。2)种子生产的种间差异极为显著, 种子量排名前三的植物贡献了累计种子雨总量的70%。3)群落种子雨的密度和物种丰富度在10年中基本同步, 均呈现显著的周期性波动, 并出现了3个大年。乔木和灌木种子雨密度的年际波动无显著相关性, 但物种数变化显著正相关; 4)种子雨与样地群落共有种为23种, 分别占种子雨和群落中木本植物种数的47.92%和54.76%, 但这些共有种贡献了种子总量的96.22%, 表明扩散限制在研究群落中十分显著。与国内其他森林群落种子雨研究结果相比, 该研究群落的种子雨密度明显较低。     

种子雨研究进展

种子雨是指在特定的时间和特定的空间从母株上散落的种子量。种子雨的组成和大小具有时空异质性。种子雨的空间异质性表现在种子雨的组成和大小因群落而异,种群间的种子雨因种群而异,种群内部的种子雨因个体而异;种子雨的时间异质性表现在不管是群落、种群还是种群内部的个体,其种子雨既具有季节动态,又具有年际变化。种子雨、种子库、幼苗库和地上植被相互联系、相互作用。种子雨的研究对更好地了解种群和群落动态等具有重要意义。应用现代的分子遗传标记技术、同位素标记法和荧光染料法等研究种子雨的散布过程和种子命运将是未来种子雨研究的热点,种子雨和种子库的结合研究及其与动植物关系的研究尚需加强。     

木麻黄海防林种子雨的时空动态

为了解木麻黄(Casuarina equisetifolia)种子雨的时空间分布格局和萌发性能对其天然更新的影响,对海口木麻黄海防林种子雨的时空分布特征和种子雨的萌发动态进行了研究。结果表明,木麻黄种子落雨时间始于7月中旬,结束于次年4月中旬,持续长达9个月,期间种子雨年际平均密度为1 764.63 grain m~(-2)a~(-1);种子雨高峰期在12月上中旬,落种量占总散落量的43.38%;种子雨呈聚集分布,且种子雨密度与附近球果数量呈极显著正相关;种子雨萌发率较低,仅为11.31%,但由于种子雨密度较大,可萌发的种子雨密度仍然较大,为199.58grainm~(-2)a~(-1)。因此,种子数量和质量均不是制约木麻黄天然更新困难的主要障碍因子,但聚集分布特征使木麻黄种子在林内扩散能力十分有限。     

灵山湾国家森林公园刺槐林下垂序商陆种子雨时空动态

垂序商陆(Phytolacca americana)的入侵给山东省沿海防护林的生物多样性及乔木更新造成了严重的危害, 为揭示其繁殖特性和扩散机制, 在灵山湾国家森林公园刺槐(Robinia pseudoacacia)林下由林缘至林内60 m设置5个梯度, 采用机械布点法对垂序商陆的结实量、种子雨分布和扩散格局进行了调查。结果表明: 1)垂序商陆的结实量与种子雨量由林缘至林内逐渐减小, 分别为2 273-1 846和1 382-621粒·m^-2, 不同空间种子雨密度占结实密度的60.80%-33.64%, 平均为49.52%, 人为搂割及动物取食是种子雨量损失的主要原因; 2)种子雨始于9月中旬, 林缘种子雨的发生早于林内, 种子雨的平均强度由林缘至林内逐渐减小, 在10月下旬和12月中旬各有一次种子雨高峰, 最大强度分别为32-4和59-5粒·m^-2·d^-1; 3)种子雨由林缘至林内散布范围逐渐减小, 同时随距母株距离的增加而减小, 其中最大散布距离与加权平均距离分别为210-180 cm和94.32-63.03 cm, 整体加权平均距离为81.00 cm。表明垂序商陆种子雨具有明显的时空异质性, 对预测其扩散潜力及群落结构组成趋势有重要的意义。     

香果树(Emmenopterys henryi)种群种子雨、种子库及 实生苗数量的海拔梯度变化

香果树为我国Ⅱ级野生保护植物,由于原生境中香果树种群的数量迅速减少,目前已濒临灭绝,种群亟待恢复。研究了武夷山不同海拔的香果树种群种子雨、种子库及其幼苗存活现状,目的是阐明不同海拔地区的香果树种群自然更新的动态过程,寻找其更新脆弱的环节及其影响因素,为香果树自然种群的恢复提供理论依据。以分布于不同海拔高度（819、980、1 140及1 301 m）的香果树母树为中心,在其周围（东、南、西、北四个方位）布设种子雨收集框和土壤种子库样方,连续观测种子雨和种子库的动态,并对不同地表覆盖物中种子萌发及幼苗数量进行观测和记录。结果表明,武夷山香果树种群种子雨持续时间近2个月,尤其是11月底至12月中旬为种子雨高峰期;随着海拔的上升,香果树种子总密度以及虫蛀种子密度均显著降低,但千粒重以及饱满种子密度则有所增加;香果树土壤种子库为瞬时种子库,其水平分布以母树南侧和西侧种子密度较大,母树北侧种子密度最小。香果树种子的垂直分布主要集中于枯落物和苔藓层;3月时香果树土壤种子库中存留的饱满种子仅占种子散布后1.80%,而1年实生苗存活率为6.18%,土壤种子库的损耗是其种群自然更新困难的主要原因;不同地表覆盖物对香果树幼苗的存活产生显著影响,枯落物及苔藓层的幼苗死亡率显著高于土壤表面。建议清理林内枯落物及苔藓,增加林内裸露土壤的面积,减小地上草本的盖度,以减少种子霉烂、虫蛀的几率,增加香果树实生苗胚根与土壤接触的几率以及改善光照条件,促进香果树自然更新。     

山地常绿落叶阔叶混交林种子雨的地形格局

种子雨是森林群落更新繁殖体的主要来源。而地形对植被空间格局异质性的影响机制之一 ,就是作用于种子雨的空间分布。为了在亚热带山地常绿落叶阔叶混交林群落中检验这一假设 ,在湖北宜昌市大老岭国家森林公园内、海拔 130 0～ 14 95 m之间的一片天然次生林内进行野外比较观测实验。选择 10个不同的地形部位 ,在每一点设置重复 (5个 )的种子雨收集器 ,在种子雨期间定期收集并记录种子雨的种类及数量。 2 0 0 1、2 0 0 2年的观测数据分析表明 :(1)种子雨密度和物种丰富度在不同地形坡位、坡形上差异显著 ,都沿山脊 -山坡 -山谷梯度和凸坡 -平坡 -凹坡梯度而减小 ;(2 )种子雨的密度和物种丰富度受坡向和坡度的影响不显著 ;(3)种子雨和乔木层物种构成的相似性与坡位和坡形呈显著的正相关 ;与坡度呈不显著的负相关 ,与坡向值间存在非线性关系 ;(4 )地形影响种子雨扩散的可能机制包括 ,影响不同种类母树的分布及其密度格局 ,影响不同坡位或坡形上分布的母树种子生产的强度和节律 ,影响风力的方向和大小的分布 ,从而形成水平方向种子流的源 -汇分化。     

Effect of Bat Exclusion on Patterns of Seed Rain in Tropical Rain Forest in French Guiana

To assess the impact of bats on seed dispersal in a tropical mature forest (Nouragues, French Guiana), we conducted a bat exclusion experiment and tested the hypotheses that an artificial reduction in the abundance of bats would result in: (1) a decrease in seed species diversity, at the community level; and (2) an increase in seed limitation (a failure of seeds to reach all suitable sites for germination) at the species level. Seed rain was sampled in two contiguous forest plots for a total of 120 d, using 49 seed traps (1 m²) arranged in 7 × 7 grids and spaced at 5-m intervals. Using mist nets, bat activity was reduced in one forest plot for a total of 60 nights. Thirty-nine plant species, or species groups, likely to be consumed and dispersed by bats, were collected within a total sample of 50,063 seeds. The overall seed rain was dominated by epiphytic Araceae and Cyclanthaceae (83.3%) and tree species within the genera Cecropia and Ficus (16.0%). Seeds from bat-dispersed shrubs and treelets ( Piper , Solanum , and Vismia ) were relatively rare (0.7%). The bat exclusion resulted in a 30.5 percent reduction in seed species richness and increased seed limitation for most of the species sampled. Seed limitation was caused mainly by a reduced seed rain (seed source limitation) rather than a decrease in seed dispersal uniformity (seed dispersal limitation). Therefore, bat-dispersed plants with low seed production are likely to be particularly affected by a decline in bat abundance, as a result of anthropogenic change.     

Ficus seed shadows in a Bornean rain forest

Due to their copious seed production and numerous dispersers, rain forest fig trees have been assumed to produce extensive and dense seed shadows. To test this idea, patterns of seed dispersal of two species of large hemiepiphytic fig tree were measured in a Bornean rain forest. The sample included four Ficus stupenda and three F. subtecta trees with crop sizes ranging from 2,000 to 40,000 figs (400,000 to 13,000,000 seeds). Seed rain out to a distance of 60 m from each study tree was quantified using arrays of seed traps deployed in the understory. These trees showed a strongly leptokurtic pattern of dispersal, as expected, but all individuals had measurable seed rain at 60 m, ranging from 0.2 to 5.0 seeds/m². A regression of In-transformed seed rain density against distance gave a significant fit to all seven trees' dispersal patterns, indicating that the data could be fitted to the negative exponential distribution most commonly fitted to seed shadows. However, for six of seven trees, an improved fit was obtained for regressions in which distance was also In-transformed. This transformation corresponds to an inverse power distribution, indicating that for vertebrate-dispersed Ficus seeds, the tail of the seed rain distribution does not drop off as rapidly as in the exponential distribution typically associated with wind dispersed seed shadows. Over 50% of the seed crop was estimated to fall below each fig tree's crown. Up to 22% of the seed crop was dispersed beyond the crown edge, but within 60 m of the tree. Estimates of the maximum numbers of seeds which could have been transported beyond 60 m were 45% for the two largest crops of figs, but were under 24% for the trees with smaller crops. Seed traps positioned where they had an upper canopy layer above them were associated with higher probabilities of being hit by seeds, suggesting that vertebrate dispersal agents are likely to perch or travel through forest layers at the same level as the fig crown and could concentrate seeds in such areas to some degree. The probability of a safe site at 60 m from the fig tree being hit by seeds is calculated to be on the order of 0.01 per fruiting episode. Fig trees do not appear to saturate safe sites with seeds despite their large seed crops. If we in addition consider the rarity of quality establishment sites and post-dispersal factors reducing successful seedling establishment, hemiepiphytic fig trees appear to face severe obstacles to seedling recruitment.     

Rainforest seed rain into abandoned tropical Australian pasture is dependent on adjacent rainforest structure and extent

It is well known that the recovery of abandoned tropical pastures to secondary rainforest benefits from the arrival of seeds from adjacent rainforest patches. Less is known, however, about how the structural attributes of adjacent rainforest (e.g. tree density, canopy cover and tree height) impact seed rain patterns into abandoned pastures. Between 2011 and 2013, we used seed traps and ground seed surveys to track the richness and abundance of rainforest seeds entering abandoned pastures in Australia's wet tropics. We also tested how seed rain diversity is related to the distance from forest, the proportion of forest cover in the landscape and several structural attributes of adjacent forest patches, specifically average tree height, canopy cover, tree species richness and density. Almost no seeds were captured in elevated pasture seed traps, even near forest remnants. Abundant forest seeds were found in ground surveys but only within 10 m of forest edges. In ground surveys, seeds from wind‐dispersed species were more abundant, but less species rich, than animal‐dispersed species. A survey of pasture seedling recruits suggested that some forest seeds must be dispersing more than 10 m into pasture at very low frequencies, but only a few species are establishing there. Recruits were predominantly animal‐dispersed not wind‐dispersed species. In addition to distance from forest and the proportion of forest within a 100‐ to 200‐m radius of sampling sites, the richness and density of adjacent forest trees were the most important factors for explaining the probability of seed occurrence in abandoned pastures. Results suggest that without some restoration assistance, the recovery of abandoned pastures into secondary rainforest in Australia's tropical rainforests will likely be limited, at least in part, by a very low rate of seed dispersal away from forest edges and by the diversity and density of trees in adjacent remnant forests.     

Factors Limiting Tropical Rain Forest Regeneration in Abandoned Pasture: Seed Rain,Seed Germination,Microclimate, and Soil1

Vegetation, seed rain, seed germination, microclimate, and soil physical and chemical parameters were measured in a recently abandoned pasture and adjacent primary rain forest in southern Costa Rica. The goal of this study was to assess the importance of these factors in limiting forest regeneration in abandoned pastures. Seed rain of animal dispersed species decreased dramatically in the pasture >5 m from the forest/pasture edge; fewer wind dispersed seeds fell in the pasture than in the forest, but the difference was much less than for animal dispersed seeds. Percent seed germination of most species studied was similar in the forest and in pasture with grasses; seed germination was lower during the dry season in areas of pasture cleared of grasses. Air temperature, vapor pressure deficit (VPD), and photon flux density (PFD) were much higher in the pasture than in the forest at 1 m above the ground. VPD and PFD at ground level and soil temperature were similar in the pasture and the forest, indicating that pasture grasses strongly modify microclimatic conditions near the soil surface. The lowest gravimetric water content recorded in the pasture during the dry season was 0.5 and leaf relative water contents of the two species measured in the forest and pasture were identical, suggesting that plants in the pasture were not water stressed. Levels of most soil nutrients were lower in the pasture as compared to the forest; however, aboveground and root biomass for seedlings grown in pasture and forest soils did not differ significantly. Although a number of factors impede forest recovery in abandoned pastures, these results suggest that the most imporrant limitation is lack of seed dispersal.     

浑善达克沙地沙地榆种子雨的扩散规律

于2010年采用定点连续观测的方法,研究了浑善达克沙地不同树龄(23年生、40年生、56年生)沙地榆孤立木种子雨的数量、组成及散布过程,分析沙地榆种子雨的扩散规律及其与外界环境干扰因子的关系。结果表明:(1)孤立木种子雨的总量随其年龄增长而增加,其中56年生孤立木>40年生孤立木>23年生孤立木,其种子雨的总量分别为(7744±16.26)粒/m2、(5236±18.87)粒/m2、(2456±12.67)粒/m2;(2)40年生孤立木的种子质量较好,成熟种子的比例最高,未成熟、空粒和虫害种子的比例较低,56年生孤立木次之,23年生孤立木质量最差,56年生沙地榆和40年生沙地榆是自然更新种子的主要来源;(3)榆树种子于当年5月下旬开始成熟并于6月初开始脱落,6月中旬达到降落高峰期,落种持续到6月下旬结束,落种期持续近1个月;(4)种子雨的前扩散过程中种子集中降落在主风向的下风向50 m和树冠周围20 m的范围内;(5)在种子的后扩散过程中,种子沿下风向继续运动,散落在下风向50 m范围或更远处,后扩散对前扩散表现为"削锋"作用。     

千岛湖片段化生境中木本植物种子雨基本特征及其影响因子

为探究片段化生境中木本植物种子雨的基本特征,该研究根据2015—2020年(研究期间)在千岛湖样岛上的植物群落长期监测样地内每月收集的种子雨数据,采用Kruskal-Wallis检验对木本植物的种子雨密度进行年际差异分析,对不同传播方式物种的种子雨密度进行月份间差异性分析,并利用线性混合效应模型,探究岛屿空间特征(岛屿面积、距最近岛屿的距离、距大陆的距离)以及气候因子(0 ℃以上积温、降水量)对木本植物以及不同传播方式物种的种子雨密度的影响。结果表明:(1)2015—2020年6年间,在29个样岛用240个收集器共收集到877 178粒木本植物的成熟种子,属于26科40属52种。(2)动物传播是木本植物主要的种子传播方式,不同传播方式物种的种子雨时间动态存在较大差异。(3)木本植物的种子雨年密度与岛屿面积和年积温呈显著正相关,与年降水量呈显著负相关。(4)自主传播物种的种子雨月密度与距最近岛屿的距离呈显著正相关,而动物传播物种的种子雨月密度则与距大陆的距离呈显著正相关,风力传播物种的种子雨月密度与月积温呈极显著正相关。综上表明,生境片段化通过岛屿空间特征影响了木本植物种子雨的时间动态。     

Spatial and Temporal Variation of Seed Rain in the Canopy and on the Ground of a Tropical Cloud Forest

Spatial and temporal patterns of seed rain impact plant fitness, genetic and demographic structure of plant populations, and species' interactions. Because plants are sessile, they rely on biotic and abiotic dispersal agents to move their seeds. The relative importance of these dispersal agents may shift throughout the year. In tropical forests, seed dispersal of epiphytes constitutes a major but hitherto unknown portion of seed rain ecology. For the first time, we report on patterns of seed rain for both epiphytic and terrestrial plants across an entire year in a Neotropical montane forest. To examine seed rain, we placed traps in the canopy and on the ground. We analyzed seed dispersal syndrome (bird, mammal, wind) and plant habit (epiphyte, liana, shrub, small tree, large tree) across all seasons of the year (dry, misty, wet). We found that the community of species collected in canopy traps was significantly different from the community in ground traps. Epiphytes were the most common plant habit found in canopy traps, while large trees were most common in ground traps. Species with bird‐dispersed seeds dominated all traps. Species richness was significantly higher during the dry season in ground traps, but did not vary across seasons in canopy traps. Our results highlight the distinct seed rain found in the canopy and on the ground and underscore the importance of frugivores for dispersing both arboreal and terrestrial plants in tropical ecosystems.     

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.