西双版纳不同地点和生境鼠类及蚂蚁的多样性比较

于2004年4月至2005年5月在西双版纳的3个不同地点(勐仑植物园、新山和补蚌)选择荒地、野芭蕉林和森林3种不同植被类型的生境,对2个重要的雨林种子传播类群--鼠类和蚂蚁的多样性进行了研究.采用铗日法捕获9种鼠共97只,采用陷阱法捕获104种蚂蚁共2 600头.结果表明,蚂蚁聚集生活在干燥的荒地生境中,而鼠类则集中生活在食物资源丰富的野芭蕉林生境中.不同地点鼠的种类和数量无显著差异(P＞0.05),但不同地点蚂蚁的种类和数量差异显著(P＜0.05);不同生境鼠和蚂蚁的种类和数量均差异显著(P＜0.05).地点和生境对2个类群分布有不同的影响,这种影响也通过它们对种子散布的不同作用,最终影响地区内植被的更新和循环.     

犬蝠对小果野芭蕉的取食及种子传播

2004年9月到12月,在西双版纳热带植物园沟谷雨林内,通过详细的野外观察和雾网实验以及种子的定时收集方法,对犬蝠(Cynopterussphinx)取食小果野芭蕉(Musaacuminata)的行为及规律进行了研究。发现在夜晚24:00以前,犬蝠取食小果野芭蕉有明显的2个活动高峰,分别发生在20:30和22:30左右,这一结果与雾网采样结果较为一致。在研究期间的雨季(9-10月)和干季(11-12月),犬蝠平均被捕获量为2.2±0.3只/d、1.4±0.3只/d,二者之间没有显著的差异;同时这两个季节收集的种子团数量分别为9.0±1.1个/d和7.2±1.4个/d,也没有显著差异。犬蝠对小果野芭蕉种子的传播受生境影响较为显著,各样地间种子传播的结果大不相同;地点和季节对犬蝠传播种子也不存在交互影响。犬蝠对小果野芭蕉种子的传播距离为50-200m,是小果野芭蕉有效的种子传播者     

橡胶树在西双版纳引种区的种子捕食与散布

在纯橡胶林、胶-竹复合林、沟谷季节雨林、次生林及毗邻次生林的橡胶林共5种不同生境中释放标记的橡胶树种子,观测种子捕食者以及种子被捕食、埋藏和扩散情况。结果表明:纯橡胶林和沟谷季节雨林内的橡胶树种子大多数都原地存留(80%和98%),而胶-竹复合林内几乎100%的种子均被捕食;次生林内释放的橡胶树种子捕食率也几为100%,而毗邻次生林的橡胶林种子捕食率约50%;卡方检验表明,胶-竹复合林的种子捕食率显著高于沟谷季节雨林和纯橡胶林,次生林内显著高于毗邻的橡胶林(P0.001)。绝大多数橡胶树种子被捕食者原地或搬运后取食,仅有少量种子被埋藏。大多数种子(80%)被搬运距离在10 m以内。红外感应相机拍摄和饲喂实验发现,黄胸鼠(Rattus tanezumi)和红刺鼠(Rattus surifer)为主要的种子捕食者。可见,在西双版纳引种区橡胶树种子主要是被小型鼠类取食,但捕食强度较大,而被埋藏种子比例很低,散布距离较短(10 m)。     

不同扰动生境中动物对酸苔菜种子的捕食和散布

有效的种子散布是木本植物形成入侵种需要经历的过程之一,但在预测入侵种时却常常被忽略.紫金牛科东方紫金牛(Ardisia elliptica)原产热带亚洲而在北美成为入侵植物,分布在云南南部的其同属种酸苔菜(A.solariacea)与之具有相似的生物学特征.本文以酸苔菜为研究对象,于2004年12月至次年2月分别在人为干扰轻的野象谷和人为干扰重的植物园进行酸苔菜的种子散布及捕食研究,试图了解生境变化对其种子散布和种子捕食的影响.结果表明,酸苔菜在两地的种子散布者均为白喉冠鹎(Alophoixus pallidus)、黑冠黄鹎(Pycnonotus melanicterus)和灰眼短脚鹎(Iole propinqua),但3种食果实鸟类的组成比例、拜访行为、频率及种子捕食者的影响在两地均不相同.人为干扰轻的野象谷生境中白喉冠鹎、黑冠黄鹎与灰眼短脚鹎的拜访频率分别为25%、32%和26%.取食后的第一次停栖地点有4%在10 m以外;人为干扰重的植物园生境中3种鸟的拜访频率分别为67%、8%、5%,取食后的第一次停栖地点有26%在10 m以外.人工摆放种子试验表明,地面上种子捕食者主要是啮齿类;在两生境中种子捕食率均较低(2-6%),但野象谷生境中种子捕食率仍显著高于植物园生境.野象谷生境中种子还受到象鼻虫幼虫的危害,危害率为17.9±3.5%(n=512);而植物园生境中未发现种子被象鼻虫危害(n=489).干扰对生境中的动物组成及行为造成了明显影响,并可能通过种子散布与捕食的改变而间接影响与其有密切关系植物的种群动态.     

秦岭森林鼠类对华山松种子捕食及其扩散的影响

森林鼠类的种子贮藏行为对植物的扩散和自然更新有着非常重要的影响。通过塑料片标记法,2008和2009年的9月—11月分别在秦岭南坡的佛坪国家级自然保护区内调查了森林鼠类对华山松(Pinus armandii)种子的捕食和扩散,结果显示:森林鼠类对华山松种子有着非常大的捕食压力。在2008年,几乎所有的种子(96.4%)在第3天后就被全部取食,而在2009年,也有将近一半的种子(49.6%)在第3天后被取食。但与此同时,鼠类对华山松种子的扩散也起着非常重要的作用。尤其在2009年,第3天时鼠类最高分散贮藏了17.75%的种子,而且直到第19天后仍然有12.25%被贮藏的种子存活下来。华山松种子在两个年份间的扩散历程有很大差异。在2008年,几乎所有的种子都被鼠类取食,贮藏量非常小;而在2009年,种子被贮藏的比例显著的增加。这个结果可能与种子大小年现象有着十分紧密的联系。2008年是华山松种子的小年,产量非常低。鼠类为了满足其日常的能量需求,只能大量的取食有限的种子,而减少其贮藏量。而2009年是华山松种子的大年,产量非常高。鼠类在满足其日常能量需求的同时,还有大量剩余的种子供其贮藏。     

垂叶榕种子的二次散布:蚂蚁和非蚁传植物互惠关系的新证据

蚂蚁对热带森林中小种子的二次散布会影响种子最终到达地点和种子命运.本文以一种由鸟散布的榕树垂叶榕(Ficus benjamina)为研究对象,观察蚂蚁对不同处理种子二次散布的影响.共记录到菱结大头蚁(Pheidole rhombinoda)、法老小家蚁(Monomorium pharaonis)、横纹齿猛蚁(Odontoponera transversa)和布立毛蚁(Paratrechina bourbonica)等4种蚂蚁参与了种子的搬运,其出现频率分别为71.7%、23.3%、2.5%、2.5%.蚂蚁对种子的搬运距离为179 cm±13 cm(n=159).其中,最主要的散布者菱结大头蚁把种子搬进蚁巢后,取食了内果皮,并把58.6%的净种子抛弃在巢外垃圾堆中.蚂蚁取食内果皮显著提高了种子萌发特性(萌发率由49.3%提高到93.3%).去除内果皮后种子被蚂蚁搬走的比率显著下降(由75.0%降为29.5%).垂叶榕种子外着生的内果皮,明显提高了对蚂蚁的吸引力,而蚂蚁取食内果皮显著提高了种子萌发能力,垂叶榕和二次散布的蚂蚁之间存在着明显的互惠关系.     

昆虫蛀蚀对鼠类介导下的锐齿槲栎种子扩散的影响

森林鼠类的种子贮藏行为对植物的扩散和自然更新有着非常重要的影响。然而,鼠类是否具有鉴别虫蛀种子的能力还存在一定的争议。此外,鼠类的鉴别能力是否受到食物丰富度变化的影响也未见相关报道。采用标签标记法,2011年秋季(9—11月,食物丰富季节)和2012年春季(4—6月,食物匮乏季节)分别在秦岭南坡的佛坪国家级自然保护区内,调查了森林鼠类对完好和虫蛀锐齿槲栎(Quercus aliena)种子的选择差异。结果显示:1)在秋季,尽管2种类型种子的存留动态没有显著差异,但是在后期虫蛀种子的存留时间相对更长;而在春季2种类型种子的存留动态则极为显著,几乎所有的完好种子(99%)在释放后的第3天就被鼠类全部扩散,虫蛀种子的存留时间则相对较长。2)在秋季,鼠类更喜好扩散后取食完好种子;而在春季,鼠类则喜好在原地取食绝大部分的种子,并且优先取食完好种子。3)在秋季,鼠类贮藏了更多的完好种子;而在春季,尽管完好种子在释放后第1天便达到贮藏高峰,然而由于后期的大量被捕食,2种类型种子在贮藏动态上没有显示出显著差异。研究结果表明秦岭地区森林鼠类可以准确区分完好与虫蛀种子,但是食物丰富度会影响鼠类对种子的选择策略。在食物丰富的秋季,鼠类更多地选择贮藏完好种子;而在食物相对匮乏的春季,鼠类更倾向于同时取食2种类型种子。森林鼠类通过对2种类型种子的鉴别和选择,影响不同种子的命运,从而可能对种子的扩散和自然更新产生重要影响。     

亚热带森林啮齿类动物对林区两种植物果实的捕食和扩散研究——以宽阔水自然保护区为例

2007年9月至10月,以贵州省遵义市绥阳县宽阔水国家级自然保护区亚热带森林系统内食果(或食种子)啮齿类动物与肉果植物香港四照花Dendrobenthamia hongkongensis和干果植物西南山茶Camellia pitardii为研究对象,初步研究鼠类捕食对两种植物种子扩散的影响.结果表明啮齿动物对两种植物果实及种子的搬运存在较大差异:鼠类对于肉果植物主要表现为种子的散布者,香港四照花90%的种子被成功散布,但距离较靠近母树;鼠类对于干果植物则主要表现为种子的掠食者,超过90%的西南山茶种子被鼠类采食,仅极少数能够偶然幸存下来进入土壤种子库,鼠类对干果的平均搬运距离大于肉果.     

食果鸟类对红楠种子的传播作用

肉质果植物通常依赖食果鸟类等取食果实后经消化道过程传播种子, 以完成种群的更新。红楠(Machilus thunbergii)是亚热带森林的代表性种类, 其果实具有依赖鸟类等动物取食后传播的特点。2012 年、2013 年在梅花山国家级自然保护区,研究了鸟类对红楠果实的取食及种子的传播作用。结果如下: 取食红楠种实的鸟类共计18 种, 其中整吞果实的鸟类12 种,啄取果肉的鸟类5 种, 啄食种子的鸟类1 种。整吞果实鸟类中4 种鹎科和2 种鸦科鸟类访问频次和取食量较大, 是主要的种子传播者。鸟类主要以呕吐方式传播种子。取食后初停栖点与母树距离及地面种子散布地点的分析结果表明, 鸟类可以远距离扩散种子。地面种实可被啮齿动物或蚁类再次捕食或搬运。研究表明, 红楠可以借助鸟类实现种子传播和种群更新。     

食物丰富度对鼠类鉴别和扩散虫蛀短柄枹栎种子的影响

时空因素和种子特征对森林鼠类扩散植物种子有着非常重要的影响。为了解食物丰富度对鼠类鉴别和扩散虫蛀种子的影响,于2012年春季(4—5月,食物匮乏季节)和2012年秋季(9—11月,食物丰盛季节),采用塑料片标记法,在秦岭南坡的佛坪国家级自然保护区内调查了森林鼠类对完好的和虫蛀的短柄枹栎Quercus glandulifera种子的扩散差异。结果显示:1)森林鼠类可以较为准确地区分完好种子和虫蛀种子。完好种子的扩散速度较快,而虫蛀种子在原地的存留时间相对较长。2)食物丰富度会对鼠类鉴别和扩散虫蛀种子产生一定的影响,但是这种影响并不显著,鼠类在春、秋季均倾向于优先取食或贮藏完好种子。3)完好种子在秋季被鼠类贮藏的比例(20%)显著高于春季(12.5%),说明种子在食物丰盛季节能够获得更大的利益。     

Poachers and Forest Fragmentation Alter Seed Dispersal,Seed Survival,and Seedling Recruitment in the Palm Attalea butyraceae,with Implications for Tropical Tree Diversity1

We examined the interaction between a palm and two bruchid beetles along with several mammal species to explore how poachers and habitat fragmentation may indirectly alter the spatial pattern of seed dispersal, seed predation, and seedling recruitment in central Panama. The large, stony endocarps of Attalea butyraceae decay slowly and bear distinctive scars when opened by rodents or beetles. We determined the final distance between endocarps and reproductive trees (which we call an ecologically effective dispersal distance), the predation status of each endocarp, and the distance between seedlings and reproductive trees. The 68 focal trees were divided among 14 sites and four levels of anthropogenic disturbance. Levels of disturbance included full protection from poachers, light and heavy pressure from poachers, and small island habitat fragments. Ecologically effective seed dispersal distances were greatest for protected sites, intermediate for lightly poached sites, and shortest for heavily poached sites and habitat fragments. Seed predation by rodents increased with distance to the nearest reproductive Attalea and was greatest for fully protected sites, intermediate for lightly poached sites, and least for heavily poached sites and habitat fragments. Seed predation by beetles reversed the patterns described for seed predation by rodents. Total seed predation by beetles and rodents combined was independent of distance, greatest for fully protected sites, and lower for poached sites and habitat fragments. Seedling densities were always greatest close to reproductive trees; however, the increase in seedling densities close to reproductive trees was minimal for fully protected sites, clearly evident for poached sites, and pronounced for habitat fragments. Increased seedling recruitment near conspecific trees may in time reduce tree diversity where humans disrupt mammal communities.     

Seed Removal and Predation as Factors Affecting Seed Availability of Tree Species in Degraded Habitats and Restoration Plantings in Rainforest Areas of Queensland,Australia

The post‐dispersal removal or predation of seeds of native tree species was investigated in Queensland, Australia, at degraded habitats and rainforest restoration sites where direct seeding might be used to facilitate tree regeneration (old fields or open habitats, lantana thicket, rainforest edge, and 5‐ and 10‐year‐old restoration plantings). Seed removal/predation was assessed in relation to tree seed weight and canopy density of the habitats during the wet season period. Results indicated that seed removal/predation imposes limitations on seed availability, particularly for small seeded species. In most situations, larger seeds were less removed/predated, most likely due to the more limited range of large seed consumers. The use of large, hard‐coated seeds may potentially reduce seed loss in open situations (from both seed removal and desiccation), unless large seed consumers frequent the site. Canopy cover exerted an influence on seed removal/predation, though trends varied in relation to site and the time of season. Broadcast sowing of seed under planted tree canopies at the more advanced stages of closure may in some areas result in higher seed removal/predation. Likewise, seeding in areas dominated by woody weeds may result in high seed losses to consumers such as rodents. Results suggested that undertaking direct seeding to coincide with the maximal period of fruit production may in some situations be beneficial to minimize seed loss. Overall, site context, canopy cover, and species selection appear to be important considerations when aiming to reduce loss of seeds to animal seed consumers in restoration work.     

SPATIAL PATTERNS OF SEED DISPERSAL AND POSTDISPERSAL SEED PREDATION OF CRYPTANTHA FLAVA (BORAGINACEAE)

I examined the spatial patterns of seed dispersal and postdispersal seed predation of the semidesert perennial Cryptantha flava (A. Nels.) Payson (Boraginaceae) at two sites in north-eastern Utah. Most flowers mature only one seed (nutlet) which is permanently retained within a pubescent calyx. The calyx and enclosed seed abscise from the plant as a unit. These dispersal units are effectively dispersed by wind as evidenced by the highly directional seed shadows and the long distances some of them travel (up to 31.3 m). Potential seed predators at the sites include five species of rodents, of which Peromyscus maniculatus is the most common, and two species of ants, Pogonomyrmex occidentalis and an undescribed species of Conomyrma. There were no strong spatial patterns of postdispersal seed predation. More seeds were removed from dishes placed at the bases of fruiting adults than from dishes ≥ 1.0 m away in one of three experiments. More seeds were removed from under shrubs or clumps of grass than in the open in one of four experiments. After 3–4 days, there was a consistent tendency for more seed removal from high density (75 seeds per .25 m²) quadrats than from low density (75 seeds per 6.25 m²) quadrats, but the difference was not always significant. There was a similar nonsignificant difference between high- and low-density quadrats exposed for 21 days. The pubescent calyx greatly discourages seed predation by ants, and probably also reduces predation by rodents. In addition, by increasing the surface area of the dispersal unit, the calyx may facilitate dispersal by wind.     

Seed dispersal in Erythronium dens-canis L. (Liliaceae): variation among habitats in a myrmecochorous plant

Erythronium dens-canis is a geophyte which produces a single flower each season. The fruits produce small seeds with relatively large elaiosomes. We performed experiments to investigate primary and secondary seed dispersal mechanisms of this species in different habitats in the western part of the Cantabrian Range in northwest Spain. Sticky traps were used to measure primary dispersal of seeds up to 0.5 m from mother plants. Seed cafeteria experiments were performed in different habitats to examine the role of ants and rodents in secondary seed transport and seed predation. Our results indicate that: (a) primary seed dispersal is positively skewed (99% of seeds fall within 20 cm of the mother plant) and seed dispersal distances vary significantly among plants; (b) secondary dispersal is exclusively by myrmecochory, although the proportion of seeds removed by ants differs significantly among habitats; (c) ant species composition and abundances vary among habitats; and (d) freshly dropped seeds are more likely to be removed than seeds that have begun to dry out. We conclude that secondary dispersal of seeds is greatly influenced by habitat but not by small-scale microhabitat. This revised version was published online in June 2006 with corrections to the Cover Date.     

斑块生境中啮齿类和蚁类动物对南方红豆杉地面种子的搬运

南方红豆杉(Taxus chinensis)在我国南方地区多分布在村落附近的斑块生境中,成熟后的种子直接落在母树下或被鸟类等动物搬运至其他斑块中。林下地面种子会被地面活动的动物搬运,继而影响种子命运及种群更新。于2016年、2017年南方红豆杉果期,在浙江天目山一个红豆杉种群分布地(临安市桐坑村),采用野外种子摆放实验的方法研究了动物对地面种子的搬运情况。结果表明:啮齿类动物取食是南方红豆杉地面种子消失的主要原因,但种子消失率在斑块间及边缘生境中的差异较大。母树林和竹林是啮齿类动物的主要觅食生境,两种生境的动物取食率明显高于山核桃种植园。边缘生境成为啮齿类斑块间移动的通道,而非觅食场所。4种啮齿类动物中,淡腹松鼠(Callosciurus pygerythrus)在母树林斑块以外的生境中出现频率最高,而其他地面搬运者回避利用山核桃种植园,仅在母树林和竹林斑块中搬运南方红豆杉地面种子。可见,斑块生境中动物改变了南方红豆杉地面种子的空间沉积格局,进而影响植物种群的更新。     

Survival and Scatterhoarding of Frugivores-Dispersed Seeds as a Function of Forest Disturbance

The effect of forest disturbance on survival and secondary dispersal of an artificial seed shadow (N= 800) was studied at Brownsberg Natural Park, Suriname, South America. We scattered single seeds of the frugivore‐dispersed tree Virola kwatae (Myristicaceae), simulating loose dispersal by frugivores, in undisturbed and disturbed secondary forest habitats. Seed survival rate aboveground was high (69%) within 2 wk and was negatively correlated with scatterhoarding rate by rodents, the latter being significantly lower in the undisturbed forest (9%) than in the disturbed forest (20%). Postdispersal seed predation by vertebrates was low (3%) and infestation of seeds by invertebrates was almost zero in all instances. Therefore, secondary seed dispersal by rodents in forest is not as critical for recruitment as observed among other bruchid‐infested large‐seeded species. Secondary seed dispersal by rodents may, however, facilitate seedling recruitment whether cached seeds experience greater survival than seeds remaining above ground surface.     

Response of ants to human‐altered habitats with reference to seed dispersal of the myrmecochore Corydalis giraldii Fedde (Papaveraceae)

Anthropogenic habitat disturbance has potential consequences for ant communities. However, there is limited information on the effects of ant responses on associated ecological processes such as seed dispersal. We investigated the effect of disturbance on the abundance, richness, and composition of ant communities and the resulting seed‐dispersal services for a herbaceous myrmecochore, Corydalis giraldii (Papaveraceae), in an undisturbed habitat (forest understory), moderately disturbed habitat (abandoned arable field), and highly disturbed habitat (road verge) on Qinling Mountains, China. In total, we recorded 13 ant species, and five out of these were observed to transport seeds. The community composition of dispersers was significantly different amongst habitats. The richness of the dispersers did not differ among the habitats, but their total abundance varied significantly across habitats and was 21% lower in the road verge than in the abandoned arable fields. The major seed‐dispersing ant species in both the forest understory and the abandoned arable field were large‐bodied (Myrmica sp. and Formica fusca, respectively), whereas the major seed‐dispersing ants found in the road verge were the small‐bodied Lasius alienus. This difference resulted in lower seed removal rates and dispersal distances in the road verge than in the other two habitats. The different dispersal patterns were attributed primarily to differences in dispersing ant abundance and identity, most likely in response to habitats with different degree of anthropogenic disturbance. The possible influence of disturbance on the ecological specialization of ant‐seed dispersal interaction is also discussed.     

Leaf-cutting ant nests support less dense and impoverished seed assemblages in a human-modified Caatinga dry forest

Regenerating forests make up an increasingly large portion of tropical landscapes worldwide and regeneration dynamics may be influenced by leaf-cutting ants (LCA), which proliferate in disturbed areas and collect seeds for fungus culturing. Here, we investigate how LCA influences seed fate in human-modified areas of Caatinga dry forest. We evaluate the seed deposition and predation on Atta opaciceps nests, foraging habitat surrounding nest and control habitat away of nest influence of 15 colonies located along a forest cover gradient during the rainy and dry seasons. For each habitat, four 50-cm² plots were established and all seeds on the soil surface were collected along 1 year. We recorded 13,628 seeds distributed among 47 species and 36.57% of the total seeds did not show any sign of predation. Nest mound habitats supported low-density and species-poor seed assemblages, which were taxonomically distinct from the control habitats. These effects only occurred in the rainy season. The proportion of undamaged seeds were similar across the habitats. While forest cover did not influence seed assemblage in terms of species richness or seed predation, it did interact with habitat type via increments in seed abundance as forest cover increased across the nests. Forest cover also affected seed composition, but only in the rainy season. These results indicate that LCA decrease seed deposition in areas under their influence, particularly on the nest mounds. As LCA profit from human disturbance in the Caatinga, their role as seed ‘sinks’ should be enhanced in disturbed Caatinga patches, particularly during the rainy season, when most of the plant recruitment occurs. Our findings reinforce the importance of LCA as drivers of forest dynamics and resilience in human-modified landscapes.     

种子重量的生态学研究进展

作为植物生活史中的一个关键性特征,种子重量与其它许多植物性状和生态因子有关,种子重量的分异与其它一些植物性状及环境的变化关系在进化生物学上已经成为一个非常有意义的研究内容,且具有一定的实践意义。种子重量被发现与下列的一些植物学和群落学性状有关:植物的生活型、种子的散布能力、种子的散布方式、植物的高度、植物的冠幅、植物的比叶面积、植物的寿命、动物的捕食、植被中植物的数量或多度、土壤中种子的数量或多度、种子的休眠、种子在土壤中的持久性和植物的净初级生产力等,另外生态因子如降雨、温度、坡向、海拔、经度、纬度、光强和干扰等都影响种子的重量。种子的重量被认为是在大量小种子和少量大种子之间的进化折衷,在一定的能量限度内,较大重量的种子一般具有较少的数量,而较小重量的种子一般数量较多,这是种子重量和数量方面具有的一种反向关系。与其它性状相比,很多研究都表明种子重量和植物的生活型的关系密切。没有散布结构或风散布的种子比以动物和水作为散布媒介的种子重量要小。种子重量与捕食的关系现发现有3种格局。种子重量和形状与种子在土壤中的持久性的关系有4种格局。在干旱和阴暗的环境条件下,种子有变大的趋势。大重量种子比小种子赋予幼苗较优势的竞争地位,其原理尚有争论,尚不清楚是否是幼苗阶段的竞争决定了世界上大部分植被类型的物种组成。未来的研究方向主要有以下几个方面:1) 种子重量与植物系统学相结合,探索种子重量的变化规律;2)调查群落三向(纬度、经度和海拔)性的种子重量谱变化规律;3) 群落演替与群落种子重量谱的变化;4) 种子重量与群落中植物个体和种子的数量的关系及机理研究;5) 微生境、微地形如坡向、坡位和林间隙等对种子重量的影响;6) 全球气候变化和种子重量变化的关系。