植冠与土壤种子库储存种子的萌发特性及策略

为明确黄土丘陵沟壑区植物种子库如何调控种子萌发来提高个体适合度,选择研究区7种具有种子库的主要物种为研究对象,以刚成熟和室内储存种子为对照,比较植冠宿存(5个宿存期)和土壤埋藏(5a埋藏期)对植物种子萌发特性的影响,探讨植冠种子库与土壤种子库储存下的种子萌发策略。结果表明:7种植物种子经过不同种子库储存后萌发特性表现出明显的种间差异,黄刺玫(Rosa xanthina)和水栒子(Cotoneaster multiflorus)种子萌发力表现为植冠宿存不变型、土壤储存增强型,土壤储存明显提高水栒子种子萌发速率;达乌里胡枝子(Lespedeza davurica)和狼牙刺(Sophora davidii)种子萌发力表现为植冠宿存增强型、土壤储存减弱型,种子萌发历时表现为植冠宿存延长型,土壤种子库储存还可加快达乌里胡枝子萌发速率、缩短萌发历时;茭蒿(Artemisia giralaii)和铁杆蒿(Artemisia gmelinii),种子萌发率随植冠宿存时间先升高后降低,随土壤储存时间先降低后升高,土壤储存可推迟其萌发,铁杆蒿种子萌发速率在植冠与土壤储存后均加快;紫丁香(Syringa oblata)种子萌发率随植冠宿存先升高后降低,土壤储存明显加快其种子启动萌发与速率。在黄土丘陵沟壑区,植物种子经过植冠或土壤种子库储存,或增加、加快、提早萌发充分利用有利条件提高占据性,或减少、减缓、推迟萌发分摊不利条件的风险;而且该区植物植冠与土壤储存后种子萌发特性间的关系,体现各自适应环境与应对干扰的分工与协作策略,主要表现为:单一主导型和相辅相成型。     

塔里木河下游不同退化区地表植被和土壤种子库特征

对塔里木河下游不同退化程度的4个典型断面进行了植被和土壤种子库的取样调查,采用种子萌发试验研究了不同退化区植被和土壤种子库的特征,结果表明:(1)塔里木河下游地表植被表现为严重的逆行演替,具体体现为胡杨林都为过熟林,几乎没有胸径在10cm以下的幼林;植被盖度、密度和多样性指数均维持在一个较低的水平上;随退化程度不断加重,地表植被中草本植物的相对密度、相对盖度和相对频度逐渐降低,而灌木和乔木的相对密度、相对盖度和相对频度逐渐增加;(2)研究区土壤种子库的基本特征是:土壤种子库种类贫乏、密度低、多样性指数和相似性系数不高;(3) 随退化程度的加重,土壤种子库物种数不断减少、密度明显下降、优势种组成趋于单一、表层种子库比例升高、1年生草本植物占优势逐渐向多年生草本植物和灌木植物转变及土壤种子库物种组成与地上植被物种组成上差异显著.     

植冠种子库及其生态意义研究

植冠种子库由成熟的种子宿存在植冠上形成。种子可在植物植冠中存留1～30年或者更长时间。40属木本植物中有1200种植物具有植冠种子库,它们大多分布在火灾易发区和干旱区。植冠种子库是植物繁殖对策的一个方面,具有延缓繁殖体传播、使繁殖体免受捕食、调节萌发时间、将种子降落在适宜位置等生态功能。本文对植冠种子库的概念、分类、具有植冠种子库物种的地理分布、植冠种子库的研究现状及植冠种子库研究中存在的问题进行了论述。应加强对干旱区植冠种子库种子脱落机制和沙漠地区植冠种子库生态意义的研究。     

黄土丘陵沟壑区主要物种植冠种子库动态及其生态策略

植冠种子库是植物适应环境并应对外界干扰的种子生态策略之一,研究了黄土丘陵沟壑区12种主要植物植冠种子库动态,结果表明:杠柳不具有植冠种子库,其他11种植物均具有植冠种子库;除了黄刺玫种子在翌年5月达到脱落高峰,其他植物大部分种子在冬季脱落,其中杠柳、达乌里胡枝子、茭蒿、黄柏刺和水栒子的大部分种子脱落集中偏早,铁杆蒿和土庄绣线菊的大部分种子脱落集中偏晚;植冠宿存对大部分植物种子的萌发特性表现为促进作用;但不同植物种子的萌发时滞对植冠宿存响应差异较大;9种植物种子在植冠上宿存至翌年2月底,其种子活力仍能维持达60%以上;该区植物表现出不同的植冠种子库策略,通过不同的方式来减少干扰的威胁,提高成功萌发与更新的几率,它们或具有较大规模的宿存量、或调控种子萌发特性、或提高种子维持活力的百分比。此外,全面了解该区植物形成植冠种子库的机理及对应的生态策略还有待于全面、深入的研究。     

河水漫溢对塔里木河下游荒漠河岸林地表植被与土壤种子库的影响

采用种子萌发法,研究了河水漫溢对塔里木河下游荒漠河岸林地表植被与土壤种子库的影响.结果表明：塔里木河下游漫溢区地表植被分属8科13属14种,非漫溢区为10科21属26种植物;与非漫溢区相比,漫溢区地表植被中出现了一些浅根系和喜湿的草本植物;漫溢区单位面积物种数、植被盖度、植株密度、物种多样性指数和丰富度指数均比非漫溢区有明显增加.漫溢区土壤种子库中有物种19种,比非漫溢区增加了5种;漫溢区土壤种子库总密度比非漫溢区增加了3.94倍;与非漫溢区相比,漫溢区1年生草本植物种子的比例增加了23.07%,而灌木植物种子比例减少了20.99%;多年生草本的变化则不明显;河水漫溢提高了土壤种子库的生物多样性.漫溢区和非漫溢区土壤种子库与地表植被的共有物种分别为18和9种,土壤种子库与地表植被的相似性系数分别为0.842和 0.667.     

城市绿地种子库的初步研究

对城市绿地土壤种子库的组成、种群数量、空间分布和物种多样性进行了初步研究,结果表明：城市绿地土壤种子库存中共出现植物42种,其中在第一层中共出现植物1 477株,多样性指数为2.05,均匀度是0.58,最大的物种多样性指数是3.56,第二层的物种多样性指数,均匀度和最大物种多样性指数均比第一层要低,分别为1.94、0.56和3.47,植物种类也要少一些。城市绿地土壤种子库中种的丰富度、种子苗量、多样性指数均不如远郊区山地天然次生林。城市绿地土壤种子库中蕴藏着一些潜在种群,有些植物种可以用于城市绿化建设。     

荒漠草原土壤种子库对草地管理方式的响应

以宁夏中部不同封育年限(封育1a、3a、5a和7a)、不同放牧方式(中等强度自由放牧、中等强度4区轮牧)、补播改良(2006年补播沙打旺Astragalus adsurgens和紫花苜蓿Alfalfa stiva)及未封育等管理方式下的荒漠草原为研究对象,采用空间梯度代替时间梯度法,于2012年3月下旬用样线法采集土壤样品,采用温室萌发法,统计土壤种子库物种组成和种子数,研究了不同草地管理方式下的种子库与地上植被相似性、多样性等特征。结果表明:1)宁夏荒漠草原土壤种子库中共出现14种植物,分属6科14属,除牛枝子(Lespedeza potanimii)为半灌木外其余均为草本植物。多年生植物占土壤种子库植物总数的65.3%,一年生植物占土壤种子库植物总数的35.7%;多年生植物种子数量随封育年限增加而表现为下降趋势,封育1a和未封育草地最多;2)不同管理方式下土壤种子库和地上植被的共有物种数为2—6种,封育时间延长导致土壤种子库物种与地上植被相似性下降,土壤种子库和地上植被共有物种数有减少趋势,自由放牧降低了土壤种子库和地上植被的共有物种数,补播恢复的草地土壤种子库中未出现补播物种;3)种子库物种多样性与管理方式无明显的相关性;种子库物种多样性指数4区轮牧草地最高,封育7a草地最低,物种丰富度指数封育1a草地最高,均匀度指数自由放牧草地最高,补播草地最低。若荒漠草原仅依靠土壤种子库恢复自然植被,则不建议封育时间过久,需间以适当的干扰(放牧);补播外来种未对荒漠草原土壤种子库产生贡献,反而对乡土种产生了一定限制。     

沙丘区植物植冠储藏种子的活力和萌发特征

研究了科尔沁沙地10种植物的植冠储藏(成熟后在植冠中保留至翌年5月)种子的活力及萌发特性.结果表明：典型沙生植物(流动沙丘及半固定沙丘植物)沙蓬、乌丹蒿和差巴嘎蒿的植冠储藏种子80%以上具有活力,而非典型沙生植物的植冠储藏种子有活力的不足80%,甚至在30%以下;典型沙生植物的植冠储藏种子表现出“速萌型”的萌发特征.植冠种子库使典型沙生植物种子的脱落延迟到风季结束、雨季来临的时候,而且此时的种子活力较高、萌发迅速.植冠种子库是典型沙生植物适应流沙和季节性干旱的主要方式之一.     

青藏高原东缘封育和退化高寒草甸种子库差异

采用幼苗萌发法研究了青藏高原东部高寒草甸封育和退化地区土壤种子库的差异.结果表明:(1)两个样地总共有10161株幼苗萌发,分属55个物种,23个科.封育样地多年生植物比例高于退化样地,禾草和单子叶植物比例在两个样地间差异不大.(2)退化样地土壤种子库种子密度((6105±1530) m-2)显著高于封育样地((3883±798) m-2),而物种丰富度差异不显著.所研究区域拥有较为丰富的种子库资源,退化地区资源更丰富,说明恢复不存在种子限制问题,土壤种子库可以成为植被恢复的潜在资源.(3)种子密度和物种丰富度在垂直分层上差异显著,且随着深度的增加而显著减小.(4)用Srensen coefficient指数计算出土壤种子库和地上植被之间的相似性在整体上较低,封育样地(45.3%)略高于退化样地(40.4%),两个样地地上植被之间的相似性为50%,而种子库之间相似性高达84.6%,说明种子库起到一个"缓冲器"的作用.(5)Shannon-Wiener多样性指数分析显示,无论在地上植被或种子库中,物种多样性都是封育样地显著高于退化样地.过度放牧不仅导致地上植被的生物多样性丧失,而且使种子库中的物种多样性降低,而封育管理可以维持地上植被和种子库中的物种多样性.     

荒漠草原区4种植物群落土壤种子库特征及其土壤理化性质

土壤种子库作为地上植被更新的潜在种源,在植被自然恢复和演替过程以及生态系统建设中起着重要作用。以宁夏盐池县荒漠草原区4种植物群落为研究对象,通过对封育16年后植物群落土壤理化性质的变化与其土壤种子库的特征之间的关系的探讨,揭示荒漠草原植物群落的土壤种子库分布特征和演替趋势,以及土壤质量的改变如何影响土壤种子库特征。研究结果表明：（1）不同植物群落土壤种子库物种组成及种子密度不同,同一物种在不同植物群落土壤种子库中出现时种子密度也存在差异;禾本科、菊科、藜科植物种在4种植物群落土壤种子库中出现比例均较高,分别占到26.19%、21.43%和19.05%,占总物种数的66.7%;灰绿藜、冰草、碱蓬在4种植物群落土壤种子库中均有出现。（2）4种植物群落土壤种子库中多年生植物和一年生植物种子居多,灌木、半灌木植物种较少。（3）4种植物群落土壤种子库中物种数和种子密度均表现为：芨芨草群落 > 苦豆子群落 > 油蒿群落 > 盐爪爪群落;且随着土层的加深,4种植物群落土壤种子库中物种数和种子密度均呈递减趋势。（4）4种植物群落土壤种子库中Shannon-Wiener指数、Simpsin指数、Pielou指数和Patrick指数均表现为芨芨草群落 > 苦豆子群落 > 油蒿群落 > 盐爪爪群落。此外,芨芨草群落土壤种子库与油蒿群落土壤种子库相似性最高,油蒿群落与盐爪爪群落的最低。（5）土壤种子库物种多样性特征指数与土壤pH、含水量、碱解氮呈正相关,与土壤电导率呈负相关,其中土壤pH和电导率对种子库物种多样性的影响较大。荒漠草原封育以后,土壤种子库中植物种数、种子密度和物种多样性均呈增加趋势,且以多年生植物和一年生植物种子居多;禾本科和豆科植物的物种数和种子密度大于菊科和藜科,这表明围封后牧草品质改善,植物群落正向演替。     

Plant traits enabling survival in Mediterranean badlands in northeastern Spain suffering from soil erosion

Question: This study analysed the effect of severe soil erosion on species composition of plant communities by favouring species showing certain growth forms, root‐sprouting and clonal growth abilities. Location: The study area was located between the middle Ebro Valley and the Pre‐Pyrenees (northeastern Spain). Methods: Root‐sprouting and shoot‐rooting abilities, clonal reproduction and growth form were assessed for the 123 most common plant species from eroded lands in the study area. We obtained 260 vegetation relevés in three different substrata (gypsum outcrops, Miocene clays and Eocene marls) on areas with different degrees of soil erosion. The frequency of every plant trait in each relevé was estimated according to species presence. The effect of soil erosion on the frequency of plant attributes was assessed by correlation analyses. Results: Bipolar, non‐clonal plants and annual species decreased their frequency with increasing soil erosion in the three substrata analyzed, whereas root‐sprouters and woody plants (mostly sub‐shrubs) increased their frequency in most of the substrata analysed. Conclusions: Woody sub‐shrubs, root‐sprouters and clonal species are favoured in eroded lands in NE Spain. Bipolar species and annual plants might not be plastic enough to survive the high stress and frequent disturbances prevailing in such eroded areas.     

黄土丘陵沟壑区土壤侵蚀环境下芽库的季节动态及垂直分布

研究了陕北黄土丘陵沟壑区5种土壤侵蚀环境（阳沟坡、阳峁坡、峁顶、阴峁坡、阴沟坡）下的植被组成、芽库组成、芽库季节动态及垂直分布特征.结果表明： 该区拥有永久性芽库的多年生物种占总物种数的80.3%,具季节性芽库的一年生植物占19.7%.在侵蚀严重的阳沟坡和阳峁坡,植物返青期的永久性芽库存量较大,而在侵蚀强度较小的峁顶、阴沟坡和阴峁坡,植物开花结实期的季节性芽库密度较大.不同侵蚀环境下地下部分芽库库存量占总永久性芽库的比例较稳定,地面永久性芽库存量在阳坡较大,而地上部分季节性芽库存量在阴坡和峁顶较大.由于不同侵蚀环境下植物群落的物种组成不同,加上土壤侵蚀干扰和植物季节更新,影响了芽库在季节及垂直分布上的变化.在黄土丘陵沟壑区植被更新过程中, 芽库具有重要作用.     

The Role of Propagule Banks from Drainage Ditches Dominated by Free‐Floating or Submerged Plants in Vegetation Restoration

Dominance by free‐floating plants results in a loss of plant species in many waters. An important source for re‐establishment of non‐floating aquatic plants can be the propagule bank. This study focuses on whether the propagule bank of free‐floating plant–dominated ditch sediments can serve as potential source for recovery of a diverse plant community. The first objective was to determine differences in propagule germination from sediments of ditches in the Netherlands that differ in vegetation composition through a seedling‐emergence experiment. The second objective was to analyze the effect of sediment disturbance on the number of germinating propagules. The results show that, compared to sediments from ditches with submerged vegetation, sediments from free‐floating plant–dominated ditches produced significantly lower numbers of individuals and species of submerged and emergent plants, while numbers of individuals and species of free‐floating plants were higher. These results suggest that sediments from free‐floating plant–dominated ditches have lower potential to recover a diverse plant community probably resulting from positive feedback mechanisms caused by the vegetation present, maintaining the free‐floating plant–dominated state. Sediment disturbance strongly favors the germination of free‐floating plant propagules, especially from free‐floating plant–dominated ditch sediments. Ditch maintenance activities such as mowing and dredging will therefore likely favor persistence of the free‐floating plant–dominated state. To shift from dominance by free‐floating plants to a more diverse plant community, alternative maintenance methods should be considered that cause less sediment disturbance together with measures that promote colonization such as temporary drawdown or re‐introduction of species.     

Regeneration Potential of Floodplain Forests Under the Influence of Nonnative Tree Species: Soil Seed Bank Analysis in Northern Italy

Soil seed banks are the ecological memory of plant communities and might represent their regeneration potential. This study examines the soil seed bank in hardwood floodplain forests of the biosphere reserve “Valle del Ticino” (Northern Italy) to find out whether the natural forest vegetation can potentially be restored by the soil seed bank. We compared near natural forests of the phytosociological association Polygonato multiflori–Quercetum roboris with stands dominated by the nonnative tree species Robinia pseudoacacia and Prunus serotina in order to investigate whether the composition of the soil seed bank is significantly influenced by the composition of the main canopy tree species and soil properties. Soil seed bank samples were taken from 20 randomly selected plots in stands that were differentiated into four groups related to the dominant forest canopy species. The germinated plants were counted and their species determined. A total of 2,427 plants belonging to 84 species were recorded. The composition of the dominant tree species and soil parameters significantly influence the composition of the seed bank. The similarity with the standing vegetation was very low. Only 13% of the species in the soil seed bank represent the target vegetation. The low percentage of target species and the high percentage of nonnative species imply that the regeneration of near‐natural forest vegetation from the soil seed bank is not feasible. Consequently, disturbances that may activate the soil seed bank should be minimized. Thus, we recommend stopping the mechanical removal of the nonnative tree species in the Ticino Park .     

甘南亚高山草甸群落的物候谱研究──兼论群落种多样性维持的机制

本文研究了亚高山草甸群落中常见的４２种植物的物候期。研究结果表明,此４２种植物在返青时间顺序、进入生殖生长的个体数量比例和进入生殖生长阶段的起始时间、历期上各不相同。根据返青顺序可将４２种植物分为三类。以单子叶禾莎草的返青时间为最早,多年生双子叶植物的返育时间较晚。根据进入生殖生长的个体占种群总个体数量的比例可分为：（１）生殖生长的个体比例在２０％以下,主要靠营养繁殖的种类;（２）具有双重繁殖特性的种类;（３）生殖个体占５０％以上的种类和（４）完全依靠有性繁殖的种类共四类。根据进入生殖生长的起始时间可分为三类。以上结果在一定程度上说明了植物种间由于在生长发育时间上的错位,减缓了种间对资源（如土壤矿物质营养）的竞争,使得群落中有多个种共存。     

Comparison of floristic changes on vegetation affected by different levels of soil erosion in Miocene clays and Eocene marls from Northeast Spain

Increased soil erosion on Eocene marls from N Aragón (NE Spain) tends to reduce vegetation cover and plant species number, but little is known about its effect in the neighbouring Miocene clays. In this study, the vegetation of strongly eroded areas on Miocene clays was analysed in terms of erosion intensity and compared with Eocene marls. Relevés were carried out on uniform patches of vegetation affected by different levels of erosion. The degeneration of vegetation cover explained 34% of the variation in species number as opposed to 48% in marls, and a clear pattern of species replacement through the destruction of the vegetation cover was not observed. Approximately 25% of the species decreased significantly and 4% increased, as opposed to 47% and 0% in marls, respectively. Erosion on marls may be more severe (more disturbance) and less stressing for vegetation (more water availability) than on clays. The few species that colonized intermediate degeneration stages and highly eroded sites were more common in non-eroded areas in drier bioclimatic belts. Thus, the degeneration of vegetation by soil erosion favoured the establishment of xeric species. The ecological range of erosion-resistant species was not wider than non-resistant species. Overall, increased soil erosion selected for different plant species in marls than in clays.     

Experimental evidence for a delayed response of the above-ground vegetation and the seed bank to the invasion of an annual exotic plant in deciduous forests

Invasions by alien plants significantly affect native biodiversity and ecosystem functioning. We conducted a 5-year field experiment to investigate potential effects of the annual invasive plant Impatiens glandulifera on both the native above-ground vegetation and the soil seed bank in a deciduous forest in Switzerland. Eight years after the establishment of I. glandulifera, we set up plots in patches invaded by the alien plant, in plots from which the invasive plant had been manually removed and in plots which were not yet colonized by the invasive plant. We examined plant species richness, diversity and plant species composition in the above-ground vegetation and soil seed bank in all plots one year and five years after the initiation of the experiment. The 36 plots (3 plot types × 6 replicates × 2 sites) were equally distributed over two forest sites. Neither the native above-ground vegetation nor the soil seed bank was influenced by the presence of I. glandulifera one year after the start of the field experiment. After five years, however, plant species richness of both the above-ground vegetation and the soil seed bank was reduced by 25% and 30%, respectively, in plots invaded by the alien plant compared to plots from which I. glandulifera had been removed or uninvaded plots. Furthermore, plots invaded by the alien plant had a lower total seedling density (reduction by 60%) and an altered plant species composition in the soil seed bank compared to control plots. Our field experiment indicates that negative effects of the annual invasive plant on the native above-ground vegetation and soil seed bank of deciduous forests become visible with a delay of several years.     

Seed production of co-occurring species: Regenerative strategies,plant economic spectrum or architectural constraints?

Although plant clonality is an important reproductive strategy complementing seed reproduction, their interrelationship is seldom studied. We evaluated how plant clonality, together with plant economics spectrum and architectural constraints, affect the generative reproduction characteristics of co-existing grassland species. For this purpose, we collected data on seed number, seed mass and seed yield (seed mass x seed number) per shoot of 56 herbaceous species from temperate grasslands in the White Carpathian Mts. (SE Czech Republic).Clonality had a weaker effect on seed traits than leaf economics and plant architecture. Metamer size showed the strongest relationship to seed number per shoot, while leaf dry matter content and canopy height were the most important factors affecting average seed mass in the studied species. Seed yield was affected by both leaf nitrogen and metamer size but only for plants with long lived shoots (di- and polycyclic). The negative correlation between seed mass and seed number per shoot for our dataset was confirmed.Our study did not support the idea that clonality is the most important factor affecting seed production in grassland species and pointed out complex relationships of coordinated traits in coexisting species.