六盘山区辽东栎的实生苗更新及其影响因子

研究了六盘山区辽东栎(Quercus liaotungensis)灌丛的种子产量、土壤种子库组成、湿沙和风干贮藏对种子寿命的影响以及动物取食子叶对种子萌发和幼苗建立的影响。结果表明, 辽东栎灌丛种子的完好率为27.51%, 被动物取食或搬运种子的比例(41.51%)显著高于其他类型种子(p < 0.01); 辽东栎次生林土壤种子库中萌发和虫蛀种子分别占35.16%和38.29%, 完好种子仅占13.65%, 捕食动物主要通过贮藏或搬运而影响土壤种子库中的种子密度。湿沙贮藏60天的辽东栎种子自动萌发率高达96.67%, 短期贮藏可加快种子的萌发进程, 提高萌发率和萌发指数, 但随着贮藏时间的延长, 种子萌发进程延迟, 萌发率、萌发指数和活力指数均不同程度地降低。排除动物取食处理的幼苗在林窗和林下生境的存活率分别为80%和83%, 而不排除动物取食幼苗在2种生境中分别仅有25%和31%能够存活, 表明子叶在幼苗建立中具有重要作用。林窗中幼苗子叶的动物取食率(85.00%)高于林下(71.00%), 子叶留存的幼苗在林窗中的存活率(6.00%)低于林下(15.50%), 而子叶被取食幼苗的存活率在两种环境中基本相等(分别为18.50%和18.00%)。     

辽东栎林内不同小生境下幼树植冠构型分析

以黄土高原黄龙山林区辽东栎林内3个小生境（林下、林隙、林缘）下辽东栎天然更新幼树为研究对象,采用典型抽样法对辽东栎幼树侧枝、叶片和树冠的空间分布状况以及生物量分配状况进行调查分析,探讨微生境与幼树植冠构型特征的关系,明确辽东栎幼树对不同小生境的适应策略,为栎林经营和林分结构优化提供理论依据。结果显示：（1）3种生境下辽东栎幼树构型发生了可塑性变化,林下幼树树冠层次比较单一,林隙与林缘的幼树树冠层次更加丰富。（2）由林下至林缘,幼树的树高、枝下高呈逐渐减小的趋势,而地径变化趋势与之相反;幼树的冠幅、树冠面积、树冠率呈先增加后减小的趋势,并且林下与林隙、林缘的差异显著;幼树的总体分枝率、逐步分枝率、枝径比呈先增加后减小的趋势。（3）3种生境下,幼树的一级枝的枝长、直径与倾角随着树高的增加而呈减小的趋势,但3种生境的差异不显著;林下一级枝主要分布在冠层中上部,而林隙与林缘一级枝主要分布在冠层中下、中上部。（4）由林下至林缘幼树叶长、叶宽、单叶面积和比叶面积逐渐降低,而单株叶数、叶总面积、叶面积指数呈先增大后减小趋势;与其他2种生境相比林下叶片分布趋于冠层上部。（5）幼树地上部分生物量中林下主干生物量占83%,枝和叶生物量只占17%;而林隙与林缘虽然各部位生物量有所差异但比例基本一致,其中主干占66%左右,枝和叶生物量占34%左右。研究表明,林隙生境下幼树的构型优于林缘和林下生境,在今后栎林的经营中,可以通过适当间伐来增加林隙数量,为森林更新和结构的优化提供有利条件。     

锐齿槲栎林的天然更新——坚果、幼苗库和径级结构

对四川茂县地区的锐齿槲栎(Quercus aliena var.acuteserrata)天然次生林进行研究,发现(1)坚果完好比例约在14%～26%之间,完好坚果的发芽率和发芽势都比较高,但在天然状况下能够留存到隔年5月的坚果几乎没有.因此,实生幼苗的天然补充就受到了严重的限制.(2)林缘的幼苗密度低于所有其它生境的(p＜0.05),林中的幼苗密度最高,和林窗的比较有显著差异(p＜0.05),而和灌丛的比较则没有差异.在灌丛幼苗密度和林中以及林窗的比较都没有差异.动物对坚果的取食速度则是林中的最慢(10.0%),林窗和林缘的相同(分别为61.1%和66.5%),都比较快(p＜0.01).(3)林中的幼苗在小于100 cm的各高度内都有大量分布,说明在林中树冠下幼苗是可以长期存在的,在幼苗和种子阶段的锐齿槲栎可能是耐荫的.(4)在林中缺失幼树,在林缘则有包括幼树在内的完整的更新大小系列,表明林缘也可以成为更新的关键地点.(5)在栎林的每一个大小级别上,萌生个体和实生个体都按各种比例伴生,说明导致萌生的干扰一直在发生,并可能在森林的更新中扮演重要角色;也说明该地区的锐齿槲栎林由萌生和实生起源的个体混合组成.     

子午岭地区辽东栎和油松林建群种的更新生态位宽度分析

通过实地调查并采用Levins生态位宽度指数对子午岭地区主要树种辽东栎(Quercus liaotungensis)和油松(Pinus tabulaeformis)的幼苗、幼树和成树在不同坡向的辽东栎林、油松 辽东栎混交林和人工油松林3种群落中的生态位宽度进行了分析研究。结果显示:(1)辽东栎和油松的更新生态位宽度在不同群落中各不相同,成树的生态位宽度与群落类型一致;(2)辽东栎幼苗和幼树在阳坡和阴坡人工油松林中的生态位宽度分别是0.951、0.95和0.98、0.94,且生态位宽度显著大于其它2个群落,表明辽东栎幼苗和幼树能很好地适应人工油松林的环境条件;(3)在阳坡,油松幼苗的生态位宽度在辽东栎林最大为0.79,且极显著大于其它2个群落,而阳坡油松幼树的生态位宽度在人工油松林中最大为1.00;在阴坡,油松幼苗的生态位宽度在人工油松林最大为0.63,而阴坡油松幼树的生态位宽度却在油松 辽东栎混交林最大为0.83,且极显著大于其它2个群落;表明油松幼苗适应阳坡的辽东栎林环境和阴坡的人工油松林环境,油松幼树适应阳坡的人工油松林环境和油松 辽东栎林环境。结果表明,人工种植油松林有利于该地区的植被恢复,人工油松林和油松 辽东栎混交林会在该地区存在较长时间,而阳坡的油松 辽东栎混交林存在时间会更长。     

东灵山地区动物对辽东栎坚果的捕食和传播Ⅰ.排除啮齿目动物对坚果丢失的影响

动物对辽东栎(Quercus liaotungensis Koidz.)坚果的捕食被认为是影响辽东栎坚果命运的主要因素,因此直接影响幼苗的建立和自然更新。在东灵山一个落叶阔叶林中调查了辽东栎坚果被脊椎动物转运和就地消耗状况。在排除部分小型啮齿目动物前后,辽东栎坚果的丢失动态曲线不同。到实验结束时,基本上所有放置的辽东栎坚果都消失了。排除动物前的曲线上有一个拐点,而排除动物后的丢失曲线上有多个拐点。两曲线相似的地方是：最初一两天辽东栎坚果的丢失非常迅速。排除小型啮齿目动物对辽东栎坚果的丢失有影响,然而坚果尺寸和微生境条件对坚果的丢失没有影响。任意2个因子的交互作用以及3个因子的交互作用对辽东栎坚果的丢失也都没有影响。排除啮齿目动物和坚果尺寸对辽东栎坚果的就地消耗有影响,微生境以及其它因子的交互作用对坚果的就地消耗没有影响。在9月份3d捕鼠的时间里,共捕到啮齿目动物3种：大林姬鼠(Apodemus speciosus)、社鼠(Niviveniter confucianus )和花鼠(Tantias sibiricus)。研究结果表明,尽管啮齿目动物的排除能够降低辽东栎坚果的丢失速度,但它们(或者和其它的脊椎动物一起)有足够的能力将辽东栎坚果捕食或搬运到其它地方分散埋藏或将它们搬运到它们的洞穴中用作漫长冬季的主要食物来源。     

秦岭北坡不同生境栓皮栎种子雨和土壤种子库动态

为了阐明栓皮栎种子雨和种子库动态,以秦岭北坡林下、林窗、林缘3种生境栓皮栎天然林为对象,采用离地和地面收集种子的方法,连续定位观测了栓皮栎种子雨、种子库的数量、质量动态,以及幼苗发育过程.结果表明:栓皮栎种子雨从8月中旬开始,9月中旬到10月上旬达到高峰期,11月上旬结束;3种生境种子雨降落历程、发生时间和组成不同;林下种子雨强度最大,为(39.55±5.56)粒·m-2,林窗种子雨降落时间最早,持续时间最长,种子活力最高,而林缘成熟种子占其种子雨的比例最大,达58.7％.从种子雨降落结束到翌年8月,土壤种子库总储量均以林下最大,林缘最小;各生境的土壤种子库中成熟、未成熟、被啃食种子数量和种子活力均随时间变化呈递减趋势,而霉烂种子数量则相反.各生境的土壤种子库中的种子均主要集中在枯落物层,其次为0～2 cm土层,2～5 cm土层中只有极少量种子存在.3种生境中栓皮栎幼苗的密度差异显著,林窗幼苗最多,林缘次之,林下最少.说明林窗更适合栓皮栎种子的萌发和幼苗生长.在栓皮栎林的经营中,通过适当间伐、增加林窗数量,可以促进栓皮栎林的天然更新.     

种皮和播种深度对辽东栎种子萌发和幼苗早期生长的影响

在玻璃温室内遮荫条件下,研究种皮和播种深度(0、2、5、10和15 cm)对辽东栎种子萌发和幼苗早期生长的影响.结果表明： 种皮对辽东栎种子萌发具有明显的抑制作用,去除种皮后,种子萌发率显著升高,不同播种深度处理种子萌发率平均升高19.4%,且萌发指数和活力指数显著增大,萌发进程加快.去皮种子和非去皮种子的萌发率均在2 cm播种深度下最大,分别为78.9%和62.2%;萌发指数和活力指数均在2 cm播种深度下最大,而萌发速率系数在5 cm播种深度下最大.去皮处理幼苗的单株叶面积和干质量与非去皮处理相比均显著增大,但比根长显著减小.去皮和非去皮处理幼苗株高均在5 cm播种深度下最大,分别为13.8和14.2 cm;基径随播种深度增加逐渐增大,而主根长、侧根数和最大侧根长均减小;幼苗干质量受播种深度影响较小.     

子午岭三种生境下辽东栎幼苗定居限制

辽东栎(Quercus wutaishanica)是子午岭地区的乡土乔木树种,也是该地区气候顶极群落的建群种,其幼苗的补充更新影响着森林群落的结构及物种组成。在3种典型生境(辽东栎林、人工油松林、灌草丛)中,设置3因素(种子、干扰、遮荫)两水平的野外播种实验(随机区组设计),记录辽东栎幼苗出苗量,并监测幼苗同生群在3种生境中3a间的生长状况,以确定种子及微生境在辽东栎幼苗补充过程中的限制作用。播种实验样方大小30cm×30cm,共计216个样方。结果显示,在辽东栎林及油松林内,增加种子和干扰强度(去除枯落物),能引起出苗量和幼苗补充量的显著增加,且2种处理方式间存在交互作用,表明在郁闭林冠下,辽东栎幼苗的补充受到了种子和微生境的双重限制,枯落物是导致微生境限制的主要因素之一。在灌草丛生境,各种处理方式均不能增加幼苗的补充量,表明辽东栎无法在开阔生境(强光照、干旱)中完成实生幼苗的补充更新。3种生境中的幼苗同生群生存分析表明,辽东栎幼苗在森林群落中存活率显著高于灌草丛群落。根据幼苗生长指标判断,在3种生境中,人工油松林是辽东栎幼苗定居的最佳场所。与实验预期相反,灌木对辽东栎幼苗的补充无显著影响。     

辽东山区长白落叶松人工林天然更新障碍分析

通过对辽东山区不同间伐强度下长白落叶松林内种子库,种子萌发,幼苗出现、成活与生长的观测,分析了长白落叶松天然更新的主要影响因素.结果表明:40年生落叶松人工林的种子雨中,有30%的种子有活力,可以满足天然更新的需要;土壤种子库的种子主要分布在枯枝落叶层,4月下旬林内种子开始萌发,6月幼苗数量达到最大,幼苗累计出现率与间伐强度(光照)之间无显著相关,只受到地被物的影响;随间伐强度的增加,幼苗平均存活率提高,但林下幼苗生长缓慢,苗高均不超过6 cm;林内当年幼苗9月全部消失,没有超过1年生的幼苗,但皆伐迹地的更新幼苗存活较多、生长良好.初步确定地被物和光照是辽东山区长白落叶松天然更新的主要障碍因子.     

东灵山地区辽东栎幼苗的建立和空间分布

研究树木实生苗的建立对森林生态系统的保育和恢复具有重要意义,在北京东灵山地区调查了辽东栎(Quercus liaotungensis)实生和萌生幼苗在几种典型的植被类型中的空间分布以及辽东栎一年生实生幼苗在1个林窗梯度上的建立。选择几种典型的植被类型,研究辽东栎实生幼苗和萌生幼苗在森林中的空间分布。结果显示,在不同植被类型中辽东栎实生和萌生幼苗的密度存在空间差异,实生苗密度随幼苗年龄增大逐渐降低,种群的更新主要依靠萌生苗完成。在1个落叶阔叶林中选择1个林窗,在不同梯度(即林窗中间、林窗和树冠连接处,以及树冠下)上,播种辽东栎坚果,第二年秋季调查实生幼苗的2个生长指标和出苗率。结果显示,林窗对一年生实生幼苗的生长高度有影响。幼苗高度在林窗中比在树冠下大;林窗对辽东栎幼苗最长叶片长度和幼苗的出苗率没有影响。说明辽东栎实生幼苗的良好生长需要阳光比较充足的生境条件。本项研究结果建议,对辽东栎种群的更新,应该适当择伐一些较大个体,使森林形成一些林窗或林中空地,以利于辽东栎实生幼苗的建立,这样才能使辽东栎种群通过实生苗进行更好的自然更新。     

Alternative strategies of seed predator escape by early-germinating oaks in Asia and North America

Early germination of white oaks is widely viewed as an evolutionary strategy to escape rodent predation; yet, the mechanism by which this is accomplished is poorly understood. We report that chestnut oak Quercus montana (CO) and white oak Q. alba (WO) (from North America), and oriental cork oak Q. variabilis (OO) and Mongolian oak Q. mongolica (MO) (from Asia) can escape predation and successfully establish from only taproots. During germination in autumn, cotyledonary petioles of acorns of CO and WO elongate and push the plumule out of the cotyledons, whereas OO and MO extend only the hypocotyls and retain the plumule within the cotyledons. Experiments showed that the pruned taproots (>6 cm) of CO and WO acorns containing the plumule successfully germinated and survived, and the pruned taproots (≥12 cm) of OO and MO acorns without the plumule successfully regenerated along with the detached acorns, thus producing two seedlings. We argue that these two distinct regeneration morphologies reflect alternative strategies for escaping seed predation.     

人工埋藏植物种子对鼠类采食下辽东栎坚果存活及萌发率的影响

辽东栎林（Q. liaotungensis)是北京东灵山地区典型地带植被。过去研究表明,鼠类对辽东栎更新有很大影响。本研究主要调查3个问题：（1）采食辽东栎坚果的鼠类种类;（2）将坚果埋藏于土中能否减少鼠类对坚果的采食,并提高出苗率？（3）地表植被是否影响辽东栎的出苗率？1996年9月在北京东灵山地区山地选取5个类型的样地37块,每样地在地表放置10粒坚果,检查地表种子消失率。使用木板夹来调查采食坚果的鼠类种类,诱饵分别为坚果和花生米。于1997年10月,选取5块样地,间隔25 m,每样地在1m²范围内埋50粒坚果,深度5 cm,次年5月调查出苗情况。研究结果表明,大林姬鼠、社鼠和黑线姬鼠为取食坚果的主要种类,取食比例与其丰富度有关。370粒坚果放在地表后,1d、10d、20d的消失率分别达15.1%、78.1%、和 100%。250粒坚果埋入土层后,出苗率达11.2%,苗平均高度为7.8cm,出苗率与地表植被盖度有一定负相关。本研究认为,人工埋藏植物种子可以减少鼠类捕食坚果,并能增加其出苗率。     

Facilitation of tree seedling establishment in a sand dune succession

Abstract. A nucleated pattern of establishment by pine seedlings of Pinus strobus and P. resinosa around pre-established oak trees of Quercus rubra in a sand dune succession in Ontario, Canada was examined using field observations, seed planting and habitat manipulation. Densities of young pines beneath oak canopies were approximately six times greater than in treeless areas, and densities on the north sides of the trees were significantly greater than on the south sides. However, oaks younger than 35 yr showed no preferential establishment beneath them, while the pine population structures beneath older oaks indicated single periods of successful recruitment. Pine seed planted beneath and beyond oak canopies of three sizes germinated primarily beneath the canopies of medium and large-sized oaks, but subsequent survivorship of seedlings over two growing seasons was poor. Several micro-environmental conditions were changed by oak canopies, but only shade showed a pattern closely corresponding to that of seedling establishment. A habitat manipulation experiment confirmed the primary role of shade in facilitating pine seedling establishment beneath oaks. Failure of pine to continue recruiting successfully beneath facilitating trees is tentatively attributed to intraspecific competition among pine individuals.     

High Acorn Predation Prevents the Regeneration of Quercus liaotungensis in the Dongling Mountain Region of North China

The mature oak (Quercus liaotungensis Koidz) forests in the Dongling mountains of northern China have become degraded in recent years because regeneration has been limited. To determine whether or not seedling establishment of the oak is seed limited, microsite limited, or predator limited and to determine whether seedling establishment is affected by ground cover, we conducted field experiments during a mast year and investigated the fate of seeds and the soil seed bank dynamics of the oak. A large acorn crop (128.8 acorns/m²) was observed in the study period, and the peak density of acorns on the forest floor reached 46.5 acorns/m², suggesting that tree recruitment was not seed limited. Acorns in the soil seed bank were mainly lost through decay (principally after fungal attack), consumption in situ, and removal by animals. Predation (including consumption in situ and removal) accounted for 86.4% of acorn loss and was therefore likely to have been the most important factor influencing seed dynamics. More than 70% of acorns were found to have germinated, but no established seedling was observed on the forest floor. Using cages to exclude predators, it was estimated that 87% of acorns germinated and 49% became established as seedlings, indicating that the acorns on the forest floor could emerge and grow in the absence of predators. We conclude that the regeneration of the tree population is limited by predators rather than by the availability of microsites. The presence of ground cover increased the germination rate and increased the chance of seed survival in the early stage of the experiment, but at the end of the investigation, no established seedling was found in the quadrats both with and without ground cover, possibly because of high density of animal predators. On the basis of these results, we suggest that selective tree felling will increase the coverage of the herbaceous layer, which can further decrease the population density of the rodents, and thereby improve the regeneration of oak trees.     

鼠类对辽东栎橡子的搬运

在北京东灵山地区,在灌丛、森林2 种生境类型中共设计了5 个处理,以检验种子质量、种子可视度、生境类型如何影响鼠类对辽东栎（Quercus liaotungensis）橡子的搬运。每个处理包括200 枚种子。每日检查种子状态,实验持续16 d。鼠类在几天之内将大多数种子搬运走。完好橡子比虫蛀橡子消失得快。橡子在森林生境中明显比在灌丛生境中消失的快。鼠类对放置在灌丛下方和灌丛之间的橡子的搬运速率差别不大。放置在落叶下方的橡子消失速率明显慢于地表的橡子。本研究利用半存活时间测量种子的搬运率。各处理的半存活时间范围是0. 6 ～ 8. 6 d。各处理的橡子搬运率均较高,这说明鼠类有能力在种子下落到冬季来临的2 个月时间内,搬运绝大多数的橡子。实验说明,鼠类是辽东栎橡子重要的捕食者和扩散者。     

Influence of insect-infested cotyledons on early seedling growth of Mongolian oak, <Emphasis Type="Italic">Quercus mongolica</Emphasis>

Insect-infested (II) acorns germinated 3 d earlier than the healthy (H) ones. However, germination ratio of II-acorns was strongly decreased compared with H-acorns and there were great differences in activities of amylase. We found an apparently lower net photosynthetic rate and total chlorophyll contents of the first true leaf of II-acorns than of the H-ones. Maximal photochemical efficiency of photosystem 2 (PS2, F_v/F_m) decreased in seedlings germinated from II-acorns than from the H-ones. Infestation of insects, especially for weevil (Curculio spp.) had significantly negative effects on length of taproots, height of plants, dry mass (DM) of roots and the first fully expanded true leaf. Leaf area and total N content of the first true leaf declined due to limitation of resource reserves in cotyledons. Damage of cotyledons caused by weevil accounted much for poor development of seedlings germinated from II-acorns. A mutual relationship between seedling establishment and seed-infesting insects may exist due to high predation on H-acorns by small rodents. An erratum to this article is available at .     

The Importance of Cotyledons for Early-Stage Oak Seedlings Under Different Nutrient Levels: A Multi-species Study

Oaks produce acorns containing a large amount of reserves in their cotyledons, which are important for the early development of seedlings. However, it remains unclear at what level oak seedlings will rely on cotyledonary reserves under different soil nutrition levels. We carried out an indoor experiment to quantify seedling growth, mobilization of cotyledonary reserves, and soil-derived N in seedlings of eleven red and white oak species in response to contrasting soil nutrition levels. Our results showed that dry masses of shoots, roots, whole seedlings, and the remnant cotyledons did not vary significantly in response to soil nutrition levels. Moreover, soil-derived N contents in oak seedlings were not significantly different under contrasting soil nutrition levels, reflecting a low morphological and physiological plasticity of oak seedlings to soil fertilization at early growth stage. Lower soil-derived N incorporated into seedlings suggests that oaks rely mainly on cotyledonary reserves for seedling development regardless of soil fertility. A positive correlation between mean fresh masses of acorns and mean dry masses of seedlings at the interspecific level further indicates that reserves in acorns rather than soil nutrition play an important role in supporting oak seedling growth at the early stage.     

Comparative habitat susceptibility to invasion by Chinese ash (<Emphasis Type="Italic">Fraxinus chinensis</Emphasis>: Oleaceae) in a tropical Andean landscape

Chinese ash (Fraxinus chinensis) is an exotic tree species that has been used in non-commercial monospecific plantations for revegetation programs in the Central Andes of Colombia. At the Otún river watershed, these plantations occur in patches intermixed with old pastures, oak forests, and successional forests. In this heterogeneous landscape, the ash has been able to invade some of its surrounding habitats. This study evaluates the invasion patterns of ash to each of these habitats and experimentally quantifies seed and seedling survivorship and seedling growth as three processes that could determine ash establishment. Of the four habitats examined, old pastures were the most vulnerable to invasion, followed by oak plantations, and successional forest. Ash plantations exhibited recruitment levels intermediate between pastures and oak plantations. Abandoned pastures showed the highest seed germination, seedling survivorship, and seedling growth. In the ash plantations, recruitment seemed negatively affected by the low number of germinated seeds, high mortality of seedlings, and low growth. Invasion in oak plantations was constrained by high seed mortality and burial of seedlings by leaf litter, although the seedlings that did survive grew fast and produced a high number of leaves. Within the successional forest, Chinese ash seeds germinated but establishment was constrained by a reduced seedling survivorship and low growth. This research offers new evidence of how different processes affecting the establishment of an invader may differ among landscape elements, and it also yields important information for the management and control of ash in these Andean landscapes.