种子产量对鼠类扩散栓皮栎坚果的影响

为了解林木种子产量对鼠类种子扩散行为的影响,于2008 年和2009 年,在国有济源市愚公林场调查了栓皮栎的种子雨;在每年的种子雨结束后,选择次生林生境,研究了鼠类对人工释放栓皮栎坚果的扩散,旨在探讨种子产量与种子扩散间的关系。结果表明:1)取食栓皮栎坚果的鼠类主要有大林姬鼠、社鼠和岩松鼠,2008 年的鼠类捕获率(2% )低于2009 年(10%),其差异并未达到显著性水平;2)栓皮栎的种子雨构成和产量存在年际差异,2008 年以完好种子为主且产量高于2009 年,而2009 年以败育种子为主;3)2008 年人工释放栓皮栎坚果的中位存留时间显著高于2009 年;4)2008 年鼠类对栓皮栎坚果的平均扩散距离显著低于2009 年,且两年的搬运距离几乎都集中在9 m 以内;5)2008 年鼠类对栓皮栎坚果的埋藏比例显著高于2009 年。结果提示,在种子高产年份,可能有较多的种子逃脱动物的取食,从而增加种子萌发和幼苗建成的机会,最终促进植物的更新。     

包衣、埋藏的栓皮栎和枹栎种子在鼠类捕食下的存留

鼠类对壳斗科植物种子的捕食作用十分强烈,从而影响此类植物的更新.栓皮栎和枹栎是中国西南地区常绿阔叶林主要树种,不仅有重要的经济价值,而且还可用于植树造林.对栓皮栎和枹栎种子分别进行包衣地表、无包衣地表、包衣埋藏和无包衣埋藏等4种处理,并跟踪各种处理种子被鼠类捕食的情况.结果表明:各处理之间种子的中位存留时间显著不同,包衣埋藏后的种子存留时间显著长于未包衣且放置于地表的种子,埋藏显著地降低了鼠类对这2种种子的捕食;埋藏及包衣处理能够很好地保护栓皮栎和枹栎种子免于鼠类的捕食,有利于种子存留并萌发建成幼苗.该方法可用于人工播种造林以提高造林成活率.     

昆虫寄生对辽东栎种子命运的影响

报道了昆虫寄生对辽东栎 Quercus liaotungensis 种子命运的影响。结果表明：（1）被昆虫寄生的种子发芽率明显低于正常成熟的种子,被昆虫寄生的种子或者不能发芽,或者能发芽但随后死亡;（2）从虫卵到老龄幼虫的发育在橡实内完成,当橡实落地后幼虫破壳而出,整个种子库的幼虫破壳时间延续2～50天,并非高度同步化;种子库的虫寄生率高达45.4%,以柞栎象Curculio dentipes为主,还有少量其它种类,如另一种象虫Curculio sp.、栗白小卷蛾 Cydia kurokoi 和螟蛾科（Pyralidae）的一未定名种;（3）昆虫寄生影响橡实成熟,测量种子大小分布表明,被昆虫寄生的种子的大小分布介于未成熟和成熟种子之间;（4）象虫寄生取食引起16.05% 的种子能量损失,显著低于对照组。结果证明昆虫寄生对辽东栎种子的命运有重要影响,是影响辽东栎种群更新的关键因素之一。     

壳斗科三种植物种子大小对昆虫寄生及种子存活率的影响

种子内的寄生昆虫可以严重影响种子的发育、损害种子活力。种子足余策略理论认为大种子有利于抵御和适应昆虫寄生取食,但动物最优觅食理论推测,大种子更易遭受昆虫寄生。为对这两种对立观点进行验证,本实验以青冈、苦槠和麻栎各2个种群的种子为材料,对昆虫寄生与完好种子间的体积和萌发率进行比较,并对寄生种子萌发率与种子体积的关系进行了分析。结果显示:(1)在6个种群的种子中,只有松阳麻栎和青冈种群的寄生种子体积大于完好种子,其余4个种群的寄生种子体积小于完好种子,但这种差异不显著;(2)所有寄生种子的整体萌发率(18%)显著低于完好种子(45.66%)(P<0.001),在不同种群内,寄生种子的萌发率也分别显著低于完好种子。(3)比较同种植物体积差异显著的寄生种子的萌发率发现,大种子总比小种子具有更高的萌发率,但差异不显著;在不同植物的寄生种子间比较时,体积最大的麻栎种子萌发率显著高于体积较小的青冈和苦槠种子。研究结果表明,象虫在种子上产卵时对大种子没有选择偏好,在昆虫寄生取食严重损害种子活力的压力下,大种子比小种子具有更强的耐受力。     

Effects of seed size on the survival of insect- infested acorns in three oak species

种子内的寄生昆虫可以严重影响种子的发育、损害种子活力.种子足余策略理论认为大种子有利于抵御和适应昆虫寄生取食,但动物最优觅食理论推测,大种子更易遭受昆虫寄生.为对这两种对立观点进行验证,本实验以青冈、苦槠和麻栎各2个种群的种子为材料,对昆虫寄生与完好种子间的体积和萌发率进行比较,并对寄生种子萌发率与种子体积的关系进行了分析.结果显示:(1)在6个种群的种子中,只有松阳麻栎和青冈种群的寄生种子体积大于完好种子,其余4个种群的寄生种子体积小于完好种子,但这种差异不显著;(2)所有寄生种子的整体萌发率(18%)显著低于完好种子(45.66%)(P＜0.001),在不同种群内,寄生种子的萌发率也分别显著低于完好种子.(3)比较同种植物体积差异显著的寄生种子的萌发率发现,大种子总比小种子具有更高的萌发率,但差异不显著;在不同植物的寄生种子间比较时,体积最大的麻栎种子萌发率显著高于体积较小的青冈和苦槠种子.研究结果表明,象虫在种子上产卵时对大种子没有选择偏好,在昆虫寄生取食严重损害种子活力的压力下,大种子比小种子具有更强的耐受力.     

昆虫与栎树的相互关系及其对栎林更新的影响

本文概述了昆虫和栎树的相互关系以及栎林种群更新研究的进展,从叶片和栎实两个方面分别分析昆虫对栎树的危害、栎树对昆虫的防御以及栎林种群更新中的问题等。（1）危害栎树的昆虫种类丰富,它们以取食叶片和栎实为主,昆虫的寄生几乎使栎实完全失去活力。（2）栎树的防御效应能影响昆虫啃食叶片的程度,被昆早寄生的栎实提前下落以及栎实产量周期性的大小年变化,能降低昆虫寄生和取食栎实所带来的损失。（3）昆虫啃食叶片降低栎树的能量利用和营养物质贮存,昆虫对栎实的寄生和取食给栎树的种子库及苗库带来压力,直接影响到栎林的种群更新,此外,昆虫对栎实的寄生也会影响脊椎动物搬运栎实时的选择取向,间接控制栎实扩散,进而影响栎树的种群更新。     

土壤埋藏深度和埋藏时间对辽东栎种子萌发的影响

以采自宁夏六盘山区的辽东栎(Quercus wutaishanica)种子为材料,研究了不同土壤埋藏深度(0、30和50 cm)和埋藏时间(2、4、6、8、10和12周)对种子萌发的影响,以探索辽东栎种子的安全贮藏条件和长期贮藏潜力,为辽东栎的种苗繁育和退化辽东栎灌丛及次生林的恢复实践提供参考.结果表明,尽管土壤埋藏处理前期(2周)种子萌发受到抑制,但随着埋藏时间的延长,萌发率提高,萌发速率加快,且萌发指数和活力指数均呈波动性增大趋势,埋藏12周后,30和50 cm埋深处理种子的萌发率分别为90.1％和96.8％.种子霉烂率表现为随埋藏深度的增加而提高的趋势,且各埋深处理种子的霉烂率随埋藏时间的延长而提高.种子埋藏2周后,开始出现自动萌发现象;埋藏12周后,30和50 cm埋深处理种子的自动萌发率分别为24.5％和43.0％.     

生境类型对啮齿动物扩散和贮藏栓皮栎坚果的影响

在河南省济源市王屋山地区,研究了次生林、灌丛、退耕地三个生境中啮齿动物对栓皮栎坚果的取食、
扩散与贮藏情况。结果表明:1)在研究地区,取食栓皮栎坚果的啮齿动物主要有大林姬鼠、社鼠和黑线姬鼠;
2)在次生林中,啮齿动物对坚果的搬运速率慢于在灌丛和退耕地生境中的搬运速率;3)次生林生境中,贮藏
坚果占被扩散坚果的比例高于灌丛和退耕地,而灌丛生境中被取食坚果占被扩散种子的比例高于退耕地和次生
林;4)栓皮栎坚果的中位存留时间在三个生境间有显著性差异。研究结果提示,在人工造林时,可通过对栓皮
栎坚果进行埋藏处理,以提高种子萌发率。     

温度和土壤含水量对青冈栎种子萌发的影响

运用直播法,研究应用人工气候箱控制温度和土壤含水量对野生青冈栎种子萌发和幼苗生长的影响。结果表明:在12h光照条件下,青冈栎种子在变温为10℃/15℃时,萌发率很低,仅为(12±12.29)%;变温为30℃/35℃时,萌发率达最大,为(81±13.7)%。土壤含水量为30%～40%时,萌发率为(79±15.9)%;土壤含水量为90%～100%时,萌发率为(56.67±19.36)%。青冈栎种子萌发呈现出迅速萌发和推迟萌发的特点;温度对青冈栎幼苗的苗高、叶长和叶宽影响显著,对幼苗的地径、叶片数量影响不显著。恒温下,不同梯度土壤含水量处理对幼苗的苗高、地径、叶长、宽影响不显著。     

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。     

Importance of microhabitat and acorn burial on Quercus ilex early recruitment: non-additive effects on multiple demographic processes

This study investigates the effect of microhabitat and seed burial on the main demographic processes operating during the early recruitment of Quercus ilex, such as postdispersal seed predation, seed germination, and seedling emergence, survival and growth. The effect of burial was positive over all the processes analysed in this study, since predation rate was lower (63.6% vs. 88%), whereas germination (53.1% vs. 21.8%) and emergence (32.0% vs. 5.5%) were higher for buried acorns. The quality of some microhabitats remained similar throughout the stages and processes studied. Thus, afforestation provided especially suitable microhabitats for oak establishment, since seed predation was lower, while germination, emergence and seedling survival were higher, than in any other microhabitat. By contrast, the quality of some microhabitats, such as open sites and Holm oaks, differed between recruitment stages. Acorns in open sites escaped predation and germinated easily, but most seedlings died due to summer drought. Similarly, although acorns under Holm oaks can germinate and survive drought, they cannot survive to postdispersal predators. This uncoupling results in a post-dispersal change in the spatial distribution of Q. ilex recruits. Furthermore, there were significant interactions between burial and microhabitat for some demographic processes. The recruitment was in afforestations high irrespective of burial, suggesting that burial is not as beneficial in high-quality habitats as it is in lower-quality ones. An accurate understanding of plant recruitment requires the determination not only of the direct effects of limiting factors but also the potential interactions occurring between them.     

槲栎超寄生橡子内象甲幼虫的适应对策

栎属植物的橡子常常受到象甲的侵害, 对橡子存活产生影响, 但有关橡子对象甲幼虫适合度影响的研究尚未见报道。本研究旨在通过对槲栎Quercus aliena种子雨进程以及象甲幼虫逃逸过程的调查, 研究二者之间的进化适应关系。结果表明： 当年槲栎的种子产量为51.92±29.26粒/m², 虫蛀率达到42.4%。虫蛀的橡子中65%以上被象甲超寄生, 且显著大于完好的橡子。在种子下落过程中, 早期从橡子中逃逸的象甲幼虫要比后期逃逸的大。对某一超寄生的橡子而言, 首先逃逸出的象甲幼虫要比随后逃逸的幼虫大。另外, 从早期脱落的橡子中逃逸的幼虫明显小于后期脱落的橡子。象甲幼虫的干重随着橡子内象甲幼虫数量的增加而显著下降。象甲幼虫寄生数量与橡子的长径和短径呈现明显的正相关关系。尽管超寄生降低了象甲虫的适合度, 但雌性成体象甲产卵过程中偏好选择大橡子在一定程度上可以减少这种不利影响。因此, 小橡子可能更容易逃脱象甲的超寄生而获得较大的存活机会。     

槲栎种子雨进程中昆虫的捕食特征

为了了解昆虫捕食与槲栎Quercus aliena种子大小和产量的关系以及在栎林更新中的作用,于2008和2009年秋季,分别在暖温带伏牛山系天池山国家森林公园内研究了两个年度槲栎的种子雨过程及昆虫捕食特征。结果表明：（1）种子雨过程从8月中下旬到9月底约经历40 d,存在高峰期且时段较明显,高峰期下落种子量分别占全部种子雨量的78.13%（2008年）和75.91%（2009年）;（2）槲栎的种子雨强度年间存在较大差异,2008年明显小于2009年（两年分别为31.75±16.65粒/m²和51.92±29.26粒/m²）,但2009年的橡子明显比2008年小（两年分别为1.94±0.61 cm³和2.46±0.57 cm³）;（3）种子雨构成比例在两个年份间存在差异,完好种子的量分别为59.05%（2008年）和36.12%（2009年）,虫蛀率在大量结实的2009年显著提高;（4）虫蛀橡子（2.29±0.42 cm³）显著大于完整橡子（1.59±0.32 cm³）,且虫蛀种子中所含虫卵数与种子大小显著正相关,昆虫有选择大种子产卵寄生的偏好。本研究的结果说明,昆虫对槲栎大种子有寄生选择偏好;大部分槲栎种子遭遇象甲虫蛀而降低生命活力,这可能是影响槲栎林更新的主要因素之一。     

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.     

Acorn Pericarp Removal as a Cache Management Strategy of the Siberian Chipmunk,Tamias sibiricus

Numerous recent studies have revealed a variety of behavioral adaptations of rodents for maximizing returns from cached seeds. Herein we report on a novel behavior by the Siberian chipmunk (Tamias sibiricus) in northeastern China, by which they consistently remove the pericarp (shell) of Quercus mongolica acorns before dispersing and caching these nuts. We investigated the effects of pericarp removal on acorn germination, tannin concentrations, cache pilferage, and insect damage, to determine if and how pericarp removal facilitates cache management by Siberian chipmunks and whether or not such behavior influences seed fates. Chipmunks cached acorns only after the pericarps were removed. Chipmunks preferred pericarp‐removed acorns over intact acorns when removing them from seed stations for both consumption and caching. Pericarp removal did not affect germination or tannin concentration of cached Q. mongolica acorns, suggesting that the behavior is not an adaptation for long‐term storage and tannin decomposition. Acetone treatments of the pericarp and artificial pericarp removal failed to alter pilferage rates by Siberian chipmunks and wood mice (Apodemus peninsulae). Since damage of acorns by weevils often leads to cache losses, we also tested the effects of weevil infestation on cache decision following pericarp removal. Siberian chipmunks removed pericarps and then scatter hoarded significantly more sound than weevil‐infested acorns, strongly suggesting that pericarp removal is used to discriminate between the infested and non‐infested acorns. Thus, we argue that the primary function of this behavior is to ensure successful storage of sound acorns, at least for short‐term storage. Future studies should consider the potential impact of pericarp removal on weevil populations and long‐term patterns of seed survival and establishment from the Siberian chipmunk’s caches.     

都江堰地区灌丛内袍栎橡子的捕食和搬运

从2000年9月至2002年12月,以四川都江堰常绿阔叶林灌丛内的优势种--枹栎(Quercus serrata)为研究对象,研究了昆虫、鼠类和真菌对枹栎橡子的捕食和搬运.结果表明(1)取食橡子的鼠类的群落结构和数量在年间差异较小,这可能与三年间袍栎橡子的产量相对稳定有关;(2)袍栎橡子的虫蛀率在三年间存在显著性差异,范围在14.17%-32.49%之间;(3)鼠类的就地取食和搬运是影响种子库内橡子命运的主要因素;(4)橡子虫蛀率的变化影响鼠类的觅食活动.鼠类对饱满橡子和虫蛀橡子的选择性取食和搬运可能影响许多坚果植物(栎树)的自然更新过程[动物学报50(4)535-540,2004].     

都江堰地区灌丛内迺栎橡子的捕食和搬运(英文)

从 2 0 0 0年 9月至 2 0 0 2年 1 2月 ,以四川都江堰常绿阔叶林灌丛内的优势种———栎 (Quercusserrata)为研究对象 ,研究了昆虫、鼠类和真菌对栎橡子的捕食和搬运。结果表明 :(1 )取食橡子的鼠类的群落结构和数量在年间差异较小 ,这可能与三年间栎橡子的产量相对稳定有关 ;(2 )栎橡子的虫蛀率在三年间存在显著性差异 ,范围在 1 4 1 7% - 32 4 9%之间 ;(3)鼠类的就地取食和搬运是影响种子库内橡子命运的主要因素 ;(4 )橡子虫蛀率的变化影响鼠类的觅食活动。鼠类对饱满橡子和虫蛀橡子的选择性取食和搬运可能影响许多坚果植物 (栎树 )的自然更新过程     

秦岭南坡短柄枹栎和锐齿槲栎的种子产量和种子大小及其与昆虫寄生的关系

【目的】为了解昆虫寄生特征与短柄枹栎 Quercus glandulifera 和锐齿槲栎 Quercus aliena var. acuteserrata 的种子产量及种子大小的关系。【方法】本研究于2011和2012年在秦岭南坡佛坪国家自然保护区,对短柄枹栎和锐齿槲栎种群的种子产量和昆虫寄生特征进行了野外调查研究。【结果】2年中短柄枹栎、锐齿槲栎种子雨密度和种子产量均差异明显,2011年短柄枹栎种子雨密度和种子产量分别为145.76±24.56粒/m²和74.97±11.56粒/m²,分别显著高于2012年的64.09±9.61粒/m²和34.30±3.51粒/m²;2011年锐齿槲栎种子雨密度和种子产量分别为238.88±43.97粒/m²和117.34±18.76粒/m²,分别显著高于2012年的112.00±19.20粒/m²和55.96±19.20粒/m²。2011年短柄枹栎和锐齿槲栎的种子产量中完好种子所占比例分别为49.73%和50.73%,分别显著高于2012年的38.69%和44.28%;而2011年虫蛀短柄枹栎和锐齿槲栎种子所占比例分别为28.05%和19.31%,分别低于2012年的39.77%和26.63%,但并无显著差异。2011和2012年,短柄枹栎和锐齿槲栎虫蛀种子个体均显著大于完好种子,且2011年二者虫蛀种子所含幼虫数与种子大小的相关性显著,2012年二者相关性不显著。【结论】短柄枹栎和锐齿槲栎种子的虫蛀率与种子产量有一定的关系,即在种子产量较低年份,虫蛀率有升高趋势,反之有降低趋势。同时,昆虫对这两种栎树种子中个体大的种子有寄生选择偏好,同时虫蛀种子所含幼虫数与种子大小也有一定的关系,即种子越大,所含昆虫幼虫数就越多。     

Acorn crop size and pre-dispersal predation determine inter-specific differences in the recruitment of co-occurring oaks

The contribution of pre-dispersal seed predation to inter-specific differences in recruitment remains elusive. In species with no resistance mechanisms, differences in pre-dispersal predation may arise from differences in seed abundance (plant satiation) or in the ability of seeds to survive insect infestation (seed satiation). This study aimed to analyse the impact of pre-dispersal acorn predation by weevils in two co-occurring Mediterranean oaks (Quercus ilex and Quercus humilis) and to compare its relevance with other processes involved in recruitment. We monitored the patterns of acorn production and acorn infestation by weevils and we conducted experimental tests of acorn germination after weevil infestation, post-dispersal predation and seedling establishment in mixed forests. Monitoring and experimental data were integrated in a simulation model to test for the effects of pre-dispersal predation in recruitment. In both oaks pre-dispersal acorn infestation decreased with increasing acorn crop size (plant satiation). This benefited Q. ilex which exhibited stronger masting behaviour than Q. humilis, with almost a single and outstanding reproductive event in 6 years. Acorn infestation was more than twice as high in Q. humilis (47.0%) as in Q. ilex (20.0%) irrespective of the number of seeds produced by each species. Although germination of infested acorns (seed satiation) was higher in Q. humilis (60%) than in Q. ilex (21%), this could barely mitigate the higher infestation rate in the former species, to reduce seed loss. Conversely to pre-dispersal predation, no inter-specific differences were observed either in post-dispersal predation or seedling establishment. Our results indicate that pre-dispersal predation may contribute to differences in seed supply, and ultimately in recruitment, between co-existing oaks. Moreover, they suggest that seed satiation can barely offset differences in seed infestation rates. This serves as a warning against overemphasising seed satiation as a mechanism to overcome seed predation by insects. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.