川西亚高山65年人工云杉林种子雨、种子库和幼苗定居研究

川西亚高山人工针叶林已成为亚高山森林的重要组成部分, 它们是否具有持续的自然更新能力, 是决定川西亚高山针叶林群落演替方向和维持该区针叶林大面积存在的基础。以川西米亚罗亚高山人工云杉(Picea asperata)林(65 a)为研究对象, 对种子雨量年际变化、土壤种子库动态、种子萌发和幼苗定居等更新过程的关键环节进行了连续7年(2002-2008年)的野外观测, 以研究人工云杉林更新潜力及影响其更新的限制因素。结果表明: 该区云杉林种子雨一般从每年的10月初开始下落, 一直到翌年的1月底或2月初结束; 云杉种子散落存在明显的大小年现象, 种子散落周期为4年, 且大小年之间种子产量差异极大; 云杉种子从下落到土壤到种子完全失去活力不到1年时间, 属于Thompson和Grime定义的第II类土壤种子库类型。腐烂死亡和动物取食是土壤种子库损耗的主要因素, 而种子通过萌发真正转化为幼苗的比例非常低, 仅占2002年下落种子总量的3.6%。种子萌发后, 环境筛的作用导致云杉幼苗大量死亡, 尤其是在种子萌发后的一个生长季节内, 其死亡率高达78%。凋落物和苔藓是构成人工云杉林下地表的两种主要地被物类型, 二者占所有调查幼苗数量的93%左右; 两种地被物类型上0-2 cm层幼苗存活率最高, 分别占存活幼苗总数的76.07%和86.72%, 随地被物厚度增加, 幼苗存活率呈明显下降趋势, 而幼苗死亡率呈明显升高的趋势, 表明林下地被物厚度也是影响云杉幼苗定居的重要因素。两种地被物对幼苗生长的影响不同, 除株高之外, 分布在苔藓上的云杉幼苗生长参数(地径、分枝数、干重以及干重年增长率)明显高于分布于凋落物上的幼苗, 表明苔藓地被物更有利于云杉幼苗定居。尽管该区大量云杉种子下落, 但由于种子的高损耗率、幼苗的低输出率以及萌发幼苗的高死亡率, 使得人工云杉林下种子通过萌发转为实生幼苗的数量非常少, 最终真正能补充到云杉种群的个体数量非常有限。     

长白山暗针叶林苔藓植物对三种针叶树种子萌发及幼苗生长的影响

研究了长白山暗针叶林两种主要地面苔藓植物——塔藓和拟垂枝藓对红松、红皮云杉和落叶松种子萌发及幼苗生长的影响.结果表明,在一定浓度下,塔藓浸提液抑制红松发芽,拟垂枝藓浸提液促进红松发芽;苔藓浸提液对红皮云杉和落叶松的发芽率没有显著影响,但不同程度地加快了种子日萌发高峰的出现.用苔藓植物体直接培养红皮云杉和落叶松种子,发现活植物体量大时对发芽率没有显著影响,但延迟了红皮云杉种子萌发高峰的出现,加速了落叶松种子萌发高峰的出现;杀青后的苔藓植物体则明显地抑制种子发芽.通过幼苗培养实验,发现苔藓植物浸提液抑制红皮云杉和红松幼苗生长,但促进了落叶松幼苗的生长;不同处理的苔藓植物体对幼苗生长的影响差异较大,活植物体量大时,两种苔藓均促进了红皮云杉和落叶松幼苗生长,而杀青后植物体则明显地抑制幼苗生长.     

红松天然更新对动物的依赖性—一个排除动物影响的球果发芽实验

红松球果成熟后不能自行散出种子,需要依赖动物将种子从球果中取出再传播出去。为了检验红松种子能否直接在落地球果中发芽、成苗。在球果成熟期采集球果,将球果封闭在铁丝笼中,在排除动物干扰的情况下放置在林地上。2年后回收了10个球果,其中9个可看到少量种子发芽。对5个球果中115粒发育正常种子的逐个检查表明发芽率为11％。53％种子腐烂,另外35％的种子虽未腐烂也已失去发芽能力。发芽后形成的幼苗均为残缺或畸形苗。实验结果支持没有动物帮助,红松不能进行天然更新的结论及有关动物可视为红松林生态系统中的关键种或关键种组的意见。     

星鸦的贮食行为及其对红松种子的传播作用

1994－1998年秋在小兴安岭凉水自然保护区对星鸦（Nucifraga caryocatact）的贮食行为及具对红松（Pinus koraiensis）种子的传播作用进行了研究,星鸦在秋季至翌年早春主要以红松种子为食,偶食浆果。秋季种子成熟时,星鸦在取食的同时储藏大量种子作为冬季和早春的食物,这些种子中未被完全吃掉的遗留部分在第三年春天即可发芽,在地上取食部分种子后,星鸦通常储存30－50粒于舌下囊中,飞行2－5km后,将种子埋藏在土层下2.5-3.5cm深处,每个贮点从为2－4粒,贮点生境主要有8种：林缘（路边）、保护区周围空阔地、人工落叶松林、人工自冷杉林、人工红松林、人工红皮云杉林、天然更新的杨桦林和天然原始红松阔叶林（母树林）。星鸦每天至少搬运种子10次,达400粒。一个贮藏季节一只星鸦可贮藏至少16000粒种子,对红松1年生和2年生幼苗调查表明,幼苗在上述8种生境中均有分布,并且在人工落叶林、天然更新杨桦林和母树林下有幼树分布,因此星雅能有效地扩散红松种子,有利于红松的天然更新。     

动物与红松天然更新关系的研究综述

红松球果成熟后,种子自身不能脱落到达适宜的萌发地,因而无法自身完成天然更新而要依赖动物帮助传播种子。捕食红松种子的动物包括啮齿类、鸟类和一些大型兽类。其中,松鼠、星雅和普通币鸟具有分散贮藏的行为,以前两者为重要。这些动物从球果中取出种子,吃掉一部分后,靠颊囊或舌下囊搬运至贮藏地,将种子埋藏在地被物下准备冬、春食物缺乏时再重新捕食。搬运的距离远达数百米至几公里。分散贮藏的贮点数量很大。每个贮点有一至数粒种子。贮点小生境比较适合红松种子的萌发,因此大量的未被重新发现和捕食的贮点的种子在20个月后可能萌发。虽然动物贮藏的生境适合幼苗的萌发,但是不一定适合幼苗的继续生长。由于动物的捕食和贮藏行为,阔叶红松林内红松种子运转途径在4个层次（树上、地面、地被物下和洞穴）进行,最后只有少数种子萌发为幼苗。幼苗的生长过程同样受到动物的捕食和破坏。由于红松结实的丰欠年度变化,动物种群也随之变动。     

红松阔叶林中红松种子的分布及更新

 天然阔叶红松林的红松种子经过动物活动的处理,在林内呈四个层次的垂直分布．其中在地被物下是以簇状埋藏着,数量相当多,尤其在红松密林中高大树木的周围,这些簇状分布的种子成为红松天然更新的基础。在次生阔叶林中红松种子埋藏量则很小。而埋藏位置与红松幼苗适生生境的不吻合是红松天然更新的主要障碍。     

生境异质性对刺五加种子萌发的影响及其种子库动态

本文通过定位观测研究了中国东北红松阔叶林及其次生林下重要灌木刺五加种子在异质生境中转化为幼苗的差异,人工落叶松林下模拟种子库的动态。结果表明：人工落叶松林下的转化率最高,为１６．８％。以下顺序为蒙古栎林下为４．ｌ％,白桦林下为２．７％,人工红松林下为０．８％,硬阔叶林下为０．５％。人工落叶松林下模拟种子库中刺五加种子的寿命为４ａ。幼苗输出率第２年为１４．５％,第３年为１０．１％,第４年为１．８％。其它输出为：腐烂３３．ｌ％;生理衰老２２．３％;鼠类捕食１４．１％,但其很不稳定,这与鼠类种群数量动态有关。昆虫和土壤动物捕食最少,仅１．２７％。     

长白山阔叶红松林细根周转的研究

系统研究了长白山阔叶红松林细根生物量、生产力、年周转率及其在净生产力分配中的作用,生物量调查结果表明,阔叶红松林活细根的生物量为5049kg.ha~(-1),死细根生物量的平均值为1883kg.ha~(-1),细根的年周转率为0.96,年生产量为4860kg.ha~(-1),约占总净初级生产力的19.40％,年死亡量为2343kg.ha~(-1),相当于阔叶红松林枯枝落叶年凋落量的60％,由此提出了森林凋落物应包括枯枝落叶和根系凋落物的论点。     

不同年龄香果树种子雨和种子库及其更新特征

对宝天曼自然保护区香果树种子雨、土壤种子库进行了观测,并进行不同微生境播种实验,研究其种子萌发及幼苗生长动态。结果表明:(1)不同龄级香果树种子雨持续时间及其高峰期有所不同,种子雨强度存在极显著差异。(2)20~50龄级香果树的种子饱满率、土壤种子库中种子密度均显著低于其他龄级,但其不同类型种子的水平和垂直分布规律一致;大约80%的香果树种子在其萌发前消失,剩余种子中以霉烂种子密度最高(9.81粒·m-2),饱满种子密度最低(1.94粒·m-2)。(3)野外育苗实验表明,不同龄级香果树所产饱满种子的萌发率及幼苗存活率差异不显著,其种子萌发率约为16.93%,但仅有3.86%的幼苗寿命超过5个月;不同微生境对香果树幼苗存活率产生显著影响,林窗是其最适宜微生境。研究认为,人工采集香果树种子,于次年4月份在原生境下播种;清理母树冠下和冠缘的地被物,增加林缘空地地被物覆盖度,以减少香果树种子的损失,改善香果树种子萌发和幼苗生长的光照条件,以利于幼苗根系的生长,促进香果树的自然更新。     

鄂东南低丘地区枫香林下枫香幼苗更新限制因子

深入理解幼苗更新的生态过程及其限制因子是植物种群保育和森林管理的基础。在鄂东南的低丘地区, 采用控制试验研究了限制枫香(Liquidambar formosana)幼苗在其自身林下更新的生态因子。由于枫香幼苗在马尾松(Pinus massoniana)林下偶尔更新良好, 在枫香林下几乎没有更新, 以马尾松林为对照, 设计了4个试验来检验: (1)低温和土壤湿度对枫香种子储藏过程中寿命的影响, (2)两种林型的土壤、凋落物对枫香种子越冬及存活的影响, (3)凋落物对枫香种子萌发是否具有化感作用, (4)林下的光强变化是否影响幼苗的存活。结果表明, 枫香种子在很低的温度下仍可以立即萌发, 这表明枫香种子没有休眠特性是影响枫香幼苗更新的重要因子; 土壤、凋落物和凋落物滤液对种子越冬后的存活没有影响, 不能解释“枫香林下没有枫香苗”的问题; 两种林型的主要凋落物滤液对枫香种子萌发没有抑制作用或促进作用; 一年生枫香幼苗遮阴试验的结果表明, 3%透光率下幼苗的死亡率接近90%。因此枫香林下弱的光环境和种子的低温萌发特性是影响枫香幼苗更新的重要因子。     

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.     

Inhibition of seedling survival under Rhododendron maximum (Ericaceae): could allelopathy be a cause?

In the southern Appalachian mountains a subcanopy species, Rhododendron maximum, inhibits the establishment and survival of canopy tree seedlings. One of the mechanisms by which seedlings could be inhibited is an allelopathic effect of decomposing litter or leachate from the canopy of R. maximum (R.m.) on seed germination, root elongation, or mycorrhizal colonization. The potential for allelopathy by R.m. was tested with two bioassay species (lettuce and cress), with seeds from four native tree species, and with three ectomycorrhizal fungi. Inhibitory influences of throughfall, fresh litter, and decomposed litter (organic layer) from forest with R.m. (+R.m. sites) were compared to similar extractions made from forest without R.m. (-R.m. sites). Throughfall and leachates of the organic layer from both +R.m. and -R.m. sites stimulated germination of the bioassay species above that of the distilled water control, to a similar extent. There was an inhibitory effect of leachates of litter from +R.m. sites on seed germination and root elongation rate of both bioassay species compared with that of litter from -R.m. sites. Native tree seed stratified in forest floor material from both forest types had a slightly higher seed germination rate compared with the control. A 2-yr study of seed germination and seedling mortality of two tree species, Quercus rubra and Prunus serotina, in field plots showed no significant influence of litter or organic layer from either forest type. Incorporating R.m. leaf material into the growth medium in vitro depressed growth of one ectomycorrhizal species but did not affect two other species. Leaf material from other deciduous tree species depressed ectomycorrhizal growth to a similar or greater extent as leaf material from R.m. In conclusion, R.m. litter can have an allelopathic effect on seed germination and root elongation of bioassay species as well as some ectomycorrhizal species. However, this allelopathic affect is not manifest in field sites and is not likely to be an important cause for the inhibition of seedling survival within thickets of R.m.     

浇水频率和凋落物覆盖量对黧蒴锥种子萌发及幼苗存活的影响

黧蒴锥是华南用于人工造林和生态恢复的优良乡土树种.选取浇水频率和凋落物覆盖量两个控制因子,研究在不同的土壤湿度和凋落物覆盖量条件下,黧蒴锥种子萌发及幼苗存活和生长特征.结果表明: 凋落物对黧蒴锥种子萌发和幼苗生长的影响与环境的潮湿程度相关.当浇水频率为每天1次时,凋落物覆盖会抑制黧蒴锥种子萌发和幼苗早期生长,导致种子萌发率降低、幼苗死亡率提高; 在浇水频率为2 d 1次和3 d 1次时,凋落物覆盖可以改善土壤的水分状况,有助于种子萌发和幼苗存活.凋落物覆盖还可以显著地提高幼苗的生物量.在通过播种来实现亚热带灌木林地更新时,可利用凋落物覆盖促进黧蒴锥的种子萌发和幼苗存活.     

Light limitation and litter of an invasive clonal plant,Wedelia trilobata,inhibit its seedling recruitment

Background and Aims

Invasive clonal plants have two reproduction patterns, namely sexual and vegetative propagation. However, seedling recruitment of invasive clonal plants can decline as the invasion process proceeds. For example, although the invasive clonal Wedelia trilobata (Asteraceae) produces numerous seeds, few seedlings emerge under its dense population canopy in the field. In this study it is hypothesized that light limitation and the presence of a thick layer of its own litter may be the primary factors causing the failure of seedling recruitment for this invasive weed in the field.

Methods

A field survey was conducted to determine the allocation of resources to sexual reproduction and seedling recruitment in W. trilobata. Seed germination was also determined in the field. Effects of light and W. trilobata leaf extracts on seed germination and seedling growth were tested in the laboratory.

Key Results

Wedelia trilobata blooms profusely and produces copious viable seeds in the field. However, seedlings of W. trilobata were not detected under mother ramets and few emerged seedlings were found in the bare ground near to populations. In laboratory experiments, low light significantly inhibited seed germination. Leaf extracts also decreased seed germination and inhibited seedling growth, and significant interactions were found between low light and leaf extracts on seed germination. However, seeds were found to germinate in an invaded field after removal of the W. trilobata plant canopy.

Conclusions

The results indicate that lack of light and the presence of its own litter might be two major factors responsible for the low numbers of W. trilobata seedlings found in the field. New populations will establish from seeds once the limiting factors are eliminated, and seeds can be the agents of long-distance dispersal; therefore, prevention of seed production remains an important component in controlling the spread of this invasive clonal plant.     

Ingestion by an endemic frugivore enhances seed germination of endemic plant species but decreases seedling survival of exotics

Aim To test whether ingestion by endemic frugivores differentially affects the seed germination time, germination percentage and seedling survival of endemic, native and exotic fleshy fruited plant species, and to identify the principal processes and attributes driving such effects. Location Round Island, Mauritius. Methods We conducted a germination and seedling survival experiment for 3 months to test whether ingestion (gut passage and deposition in faeces) by the endemic Telfair’s skink (Leiolopisma telfairii) had a differential effect on the germination time, germination percentage and seedling survival of two endemic, four native and two exotic fleshy fruited plant species. To assess the importance of factors involved in the ingestion process, we used a factorial design with gut passage (gut‐passed vs. not gut‐passed), depulping (whole fruit vs. manually depulped seed) and the presence of faecal material (faeces vs. without faeces). In addition, the roles of species‐specific traits, seed size and deposition density (average number of seeds per faeces) were examined. Results Exotic species had a higher germination percentage than indigenous (native and endemic) species when not ingested. Following skink ingestion, there was no longer a difference, as ingestion enhanced germination percentage most in endemic species. The exotic species still germinated faster overall than the indigenous species, despite ingestion accelerating the germination time of endemics. However, ingestion strongly reduced seedling survival of the exotic species, while having no negative effect on the survival of indigenous seedlings. Overall, ingested indigenous seeds were more likely to germinate and the seedlings more likely to survive than ingested exotic seeds and seedlings. Seed size, deposition density and the removal of fruit pulp by either manual depulping or gut passage were important predictors of germination time, germination percentage and seedling survival. Main conclusions These endemic frugivores can enhance the competitiveness of endemic compared with exotic fleshy fruited plants at the critical germination and seedling establishment stage. Consequently, conservation and restoration of mutualistic endemic plant–animal interactions may be vital to mitigating the degradation of habitats invaded by exotic plants, which is of particular relevance for island ecosystems in which large numbers of endemics are threatened by exotic invaders.     

Effect of canopy position on germination and seedling survival of epiphytic bromeliads in a Mexican humid montane forest

BACKGROUND AND AIMS: Seeds of epiphytes must land on branches with suitable substrates and microclimates to germinate and for the resulting seedlings to survive. It is important to understand the fate of seeds and seedlings in order to model populations, but this is often neglected when only established plants are included in analyses. METHODS: The seeds of five bromeliad species were exposed to different canopy positions in a Mexican montane forest, and germination and early seedling survival were recorded. Additionally, the survival of naturally dispersed seedlings was monitored in a census over 2.5 years. Survival analysis, a procedure rarely used in plant ecology, was used to study the influence of branch characteristics and light on germination and seedling survival in natural and experimental populations. KEY RESULTS: Experimental germination percentages ranged from 7.2 % in Tillandsia deppeana to 33.7 % in T. juncea, but the seeds of T. multicaulis largely failed to germinate. Twenty months after exposure between 3.5 and 9.4 % of the seedlings were still alive. There was no evidence that canopy position affected the probability of germination, but time to germination was shorter in less exposed canopy positions indicating that higher humidity accelerates germination. More experimental seedlings survived when canopy openness was high, whereas survival in census-seedlings was influenced by moss cover. While mortality decreased steadily with age in juveniles of the atmospheric Tillandsia, in the more mesomorphic Catopsis sessiliflora mortality increased dramatically in the dry season. CONCLUSIONS: Seedling mortality, rather than the failure to germinate, accounts for the differential distribution of epiphytes within the canopy studied. With few safe sites to germinate and high seedling mortality, changes of local climate may affect epiphyte populations primarily through their seedling stage.     

Effects of litter on seedling establishment: an indoor experiment with short-lived Brassicaceae species

Recruitment by seeds is essential both in vegetation dynamics and in supporting biodiversity in grasslands. The recruitment by seeds is feasible in suitable vegetation gaps from the seed rain and/or by establishment from persistent soil seed banks. Cessation of grassland management results in litter accumulation, which leads to the decline of species diversity by the decreased availability of open patches. Low amounts of litter is often beneficial, while high amounts of litter is detrimental for seed germination and seedling establishment of short-lived species. In a designed indoor experiment, we explored the effect of litter on seedling establishment by germinating six short-lived Brassicaceae species with both increasing seed mass and litter cover. We found that both seed mass and litter had significant effect on germination and establishment of the sown species. Small-seeded species were significantly negatively affected by the 300 and/or 600 g/m² litter layers. No negative litter effect was detected for species with high seed masses (Lepidium spp.). No overall significant positive litter effect was found, although for most of the species cumulative seedling numbers were not the highest at the bare soil pots. Our results suggest that the negative effects of litter are less feasible on the large-seeded short-lived species than on that of small-seeded ones.     

R.M. Cowling,D.M. Richardson & S.M. Pierce (eds.) 1977. Vegetation of Southern Africa.

We determined the role of bird dispersal in seed and seedling dynamics of the tree Kalopanax pictus from 7 years of observing seed rain and seedling emergence in a broad-leaved deciduous forest in central Japan. We also performed an experiment on the influence of seed pulp on germination of seeds of K. pictus. Seeds of this species can lie dormant for several years, and this causes rather constant yearly seedling emergence in spite of irregular seed production. The spatial distribution of the seedlings that emerged each year (maximum distance from nearest conspecific seed-bearing tree of 90 m) was wider than that of gravity-dispersed seeds (max. distance of 37 m), suggesting seed dispersal by birds in winter. Emerged seedling densities at sites over 20 m from the nearest conspecific seed-bearing tree were highest in the spring of 1991, about half a year after the largest seed fall of the observation period. However, emerged seedling densities within 20 m from seed-bearing trees were highest in 1992, 1.5 years after the largest seed fall. These field observations may be explained by the experimental results on the effects of seed pulp on germination. Intact seeds germinate slowly at low germination rates, while seeds without seed pulp germinate quickly at high germination rates. Fallen seeds with seed pulp thus appear to form a seed bank near seed sources (temporal dispersal), while seeds scattered by birds appear to increase the possibility of reaching the present safe sites in distant areas with quick germination (spatial dispersal).     

Effects of storage,mucilage presence,photoperiod, thermoperiod and salinity on germination of Farsetia aegyptia Turra (Brassicaceae) seeds: implications for restoration and seed banks in Arabian Desert

Seed viability and germination are key factors in the success of restoration efforts, especially when stored seeds are used. However, the effect of seed storage on germination of most of the native Arabian species is not well documented. We investigated the effect of storage time and role of the seed mucilage in regulating germination, dormancy, salinity tolerance and consequential survival strategy of F. aegyptia in an unpredictable arid desert setting. Effect of light and temperature during germination was studied under two photoperiods and two thermoperiods using intact and de-mucilaged seeds. Presence of mucilage and thermoperiod did not affect the germination. However, seed collection year and photoperiod had a highly significant effect on the germination. Increasing salinity levels decreased the germination of F. aegyptia but ungerminated seeds were able to germinate when salinity stress was alleviated. Seed storage at room temperature enhances the germination percentage, indicating that F. aegyptia seeds have physiological dormancy and it can be alleviated by after-ripening at dry storage. In addition, F. aegyptia seeds show ability to germinate at lower salinity concentration and remain viable even at higher saline conditions, indicating their adaptability to cope with such harsh environmental conditions.     

盐生植物种子萌发对环境的适应对策

盐生环境是一种严峻的胁迫环境,对植物的生长、发育、繁殖等生活史的各阶段都产生着重要的影响。盐生植物是生长在盐渍土壤上的一类天然植物区系,它们在长期的进化过程中形成了一系列适应盐生生境的特殊生存策略。一般情况下,盐生植物种子对环境的适应能力,是植物对盐生环境适应性的重要体现;而植物发育早期对盐度的适应能力又是决定物种分布和群落组成的关键因素。在对国内外相关文献进行分析归纳的基础上,从盐分对种子萌发的影响机理及植物种子萌发对盐生环境的适应对策两个方面综述了植物种子休眠萌发与盐生环境的关系。