Effects of herbivore-bearing adult trees of the oak Quercus crispula on the survival of their seedlings

The acorn-producing oak Quercus crispula Blume is shade-tolerant and its seedlings were found to be underdispersed on the forest floor, forming seedling aggregations near fertile canopy trees. However, 84.4% of the seedlings were dead within 2 years, with the highest mortality near the adult canopy trees of Q. crispula. This was mostly due to the high levels of herbivory by lepidopteran larvae. The most serious damage was caused by spring consumers, which fed on the leaves of canopy trees in June and, when the food quality of the leaves declined, fell down to the forest floor to consume the juveniles, including the seedlings. Contrary to the expectation of the herbivore–host arms race theory, most of the herbivores were polyphagous. The most dominant spring consumer Telorta edentata (Leech) was also polyphagous, and its distribution on the forest floor showed a high level of association with the distribution of dead oak seedlings.     

鼠类对山杏种子存活和萌发的影响

山杏（Prunus armeniaca)是广布于北京山区阳坡的矮林或灌丛,易生存在土壤贫脊、干旱、或严重退化的山坡、裸地,对于保护生态环境,减少水土流失及促进山区经济发展均具有重要作用。有关鼠类在山杏更新中作用的研究较少,本研究主要是确定影响山杏更新的鼠类种类、鼠类对山杏种子的取食压力、人工埋藏种子及植被因素对种子存活和萌发的影响。于1997年10月3－4日,采用夹捕法同时调查取食花生和山杏种子的鼠类种类及种群密度。于1997年9月24－25日,通过在地表放置山桔种子,调查在鼠类取食下,地表种子消失的速度。于1997年10月20日,作种子埋藏实验,调查种子存活及萌发情况。于1997年10月26日,进行扣网实验,研究网内、外山杏种子存活及萌发情况。结果表明：大林姬鼠（Apodemus speciosus)、社鼠（Rattus confucianus),黑线姬鼠（Apodemus agrarius)为取食山杏种子的主要鼠种种类;与取食花生相比,社鼠更偏爱山杏种子,大林姬鼠对山杏种子喜食性略低,黑线姬鼠对二者喜食程度相近;山杏种子放置地表后1、10、20、30d的消失率分别为13.06%,64.46%,90.70%,96.69%;扣网实验说明,山杏在地表的萌发率极低,生长也慢;而埋入土层内可明显提高萌发率和生长率（若除去鼠类的取食,萌发率还要高）;人工埋藏实验表明,将山杏种子埋入土层5cm后,能够有效地减少鼠类对种子的取食;通过分析山杏种子萌发与植被的关系,发现山杏易在开阔、阳光充分的草丛中存活和生长,而不易在阴闭的灌丛下存活和生长。     

不同密度下辽东栎幼苗子叶丢失及其对幼苗存活和生长的影响

在宁夏六盘山区龙潭林区华北落叶松(Larix principis-rupprechtii )人工林的不同样带, 按6个密度梯度(3.24、2.56、1.96、1.44、1.00和0.64株·m^-2)移栽萌发生根的辽东栎(Quercus wutaishanica)幼苗, 研究了啮齿动物对幼苗子叶取食的密度效应及其对幼苗存活与生长的影响。结果表明, 辽东栎幼苗的子叶留存率随着幼苗密度的增大而减小。2个高密度处理(3.24和2.56株·m^-2)的幼苗子叶留存率分别在移栽6周和5周后稳定在最低水平, 分别为8.64%和7.81%; 而2个较低密度处理(1.44和0.64株·m^-2)的幼苗在最后一次观测时子叶留存率仍很高, 分别为44.44%和31.25%, 且二者均显著高于其他密度处理(p < 0.05)。受啮齿动物取食的影响, 在从高到低的不同密度处理下, 幼苗的主根留存率分别为64.16%、80.46%、87.23%、80.25%、69.84%和69.84%; 在1.44株·m^-2处理下, 顶芽留存率最高(25.23%), 显著大于3.24株·m^-2处理(4.19%) (p < 0.05), 其他密度处理的顶芽留存率均不足20%。子叶被动物取食的幼苗和子叶完整幼苗的最终留存率基本一致, 仅在个别密度处理的某一观测时期差异显著; 顶芽被动物取食的幼苗留存率略低于顶芽完整幼苗的留存率, 但个别密度处理完全相反; 不同类型幼苗的留存率随时间推移而上下波动, 可能与幼苗顶芽被动物取食后萌生形成新芽有关。幼苗的株高、基径、叶片数和单株叶面积均随密度的减小而略有增大, 但除株高外, 其他3个生长参数均在1.44株·m^-2处理时最大, 与子叶是否被动物取食有一定的对应关系, 即高密度下更多的幼苗丢失了供应其生长所需营养物质的子叶, 从而影响了幼苗的早期生长。研究结果不仅丰富了“负密度效应”理论和种群更新的“增补限制”理论, 也可为六盘山区退化辽东栎灌丛和辽东栎次生林的恢复及生态系统管理提供 参考。     

埋藏和环境因子对辽东栎(Quercus liaotun-gensis Koidz)种子更新的影响

辽东栎萌生丛零散地分布在北京山区,虽然其种子量很大,但自然状况下的实生苗非常稀少。鼠类搬运并吃掉大量散落在地表的辽东栎种子,但鼠类是否影响辽东栎的自然更新尚不清楚。本研究主要在于了解：（1）将种子埋入土层内后对辽东栎种子出苗率的影响;（2）环境因子（如坡向、地表植被等）对辽东栎种子出苗率的影响。1996年9月24日,选取5块样地（Site）,在每块样地各设置1条样线（Transect),每条样线上设置若干样点（Plot）,样点间隔10m,用于种子埋藏实验,在每个样点设置6条平行线（Parallel transect),间隔10cm,各埋入10粒种子,种子间隔2cm;第1、2、3、4、5、6条线的埋藏深度分别为0、2、4、6、8、10cm。分别于1997年6月15日和10月4日调查出苗情况。研究结果表明,和秋季辽东栎种子出苗率均与埋藏深度呈正相关,埋藏通过改善湿度和减少鼠类等动物的取食而增加了出苗率,其贡献分别占18％和12％。但鼠类对埋入土层内种子的取食率依然很高（87％）。辽东栎种子在阴坡和阳坡的最终出苗率都很高,但阳坡地要早于阴坡。夏季辽乐栎种子出苗率均与地表植被覆盖度负相关,地表植被可能是通过与辽东栎种子竞争水分和养分而降低辽东栎种子出苗要早于阴坡。夏季辽东栎种子出苗率均与地表植被覆盖度和高度相关不显著,从夏季到秋季,从东栎种子出苗率有所增加,主要是由于埋藏在深处或阴坡处的种子出苗较晚的缘故。     

The relative importance of carbohydrate and nitrogen for the resprouting ability of Quercus crispula seedlings

BACKGROUND AND AIMS: Plants need some kind of stored resources to resprout after shoot destruction. The aim of this study was to determine the relative importance of carbohydrate and nitrogen (N) storage levels for their ability to resprout. METHODS: A shoot clipping experiment was conducted on Quercus crispula seedlings, which were grown in a factorial experimental design, with two light levels (40% and 3% of full light) and three nutrient concentrations (low, medium and high). KEY RESULTS: At the time of shoot clipping (the end of spring leaf expansion), seedlings exposed to 40% light had an average total non-structural carbohydrate (TNC) concentration of 17.0% in their roots compared with 4.9% in the roots of seedlings exposed to 3% light, and the average amount of TNC (TNC pools) in the roots was 203.8 mg and 20.0 mg at 40% light and 3% light, respectively. In contrast, root N concentration averaged 2.3% in the 3% light treatment compared with 1.2% in the 40% light treatment, and it increased with successive rises in nutrient concentrations at both light levels. Regardless of the nutrient status, at the 40% light level >80% of the seedlings resprouted after shoot clipping. Few seedlings, however, resprouted at the 3% light level, particularly in the medium- and high-nutrient treatments. Furthermore, both root TNC concentrations and TNC pools decreased after resprouting, but the amount of root N remained constant. CONCLUSIONS: These results suggest that carbohydrate storage has a stronger influence on resprouting in Quercus crispula than N storage. However, the size of the resprouting shoot was positively correlated with the amount of both N and TNC in roots. The level of N storage is, therefore, also important for the growth of resprouting shoots.     

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.     

Gnawing damage by rodents to the seedlings ofFagus crenata andQuercus mongolica var.grosseserrata in a temperateSasa grassland-deciduous forest series in southwestern Japan

The effects of dwarf bamboo,Sasa, cover on the initial morrality of hardwood seedlings were investigated by transplanting 1-year-old beech (Fagus crenata) and current-year oak (Quercus mongolica var.grosseserrata) seedling to three different stands; old-growth beech and secondary oak forests withSasa undergrowth, and aSasa grassland in a grassland-forest series near the top of Mt Jippo, southwestern Japan. The most frequent cause of seedling morrality was gnawing of the stems by rodents. In the beech forest, the gnawing was more likely to occur underSasa cover, suggesting that it provides a good habitat for rodents on the beech forest floor. TheSasa under growth may thus play an imporrant role in regeneration of beech forest. In the oak floor, mortality of both species was low and only a little gnawing occurred during a year. However, no natural oak seedling were found in the forest even after a mast year. This may be because most of the acorns disappeated before establishment. The early-stage demography of hardwood seedling as oak may thus play an imporrant role in regeneration of oak forest. In theSasa grassland where the seed supply is small, almost all of the seedlings died fromo gnawing regardless of the presence ofSasa cover. These factors prevent the recruitment of a sizable seedling bank. Rodents may thus play an imporrant role in maintenance of theSasa grassland.     

覆网保护和埋藏对坚果树种种子直播的影响

作者以都江堰林区6种代表性坚果树种为对象,研究了覆网保护和埋藏等措施对直播种子存活和幼苗建成的影响,同时调查了鼠类在春秋两季的数量变化。结果表明:秋季鼠类对种子的捕食压力大,使得种子在无保护的情况下很少存活到出苗;而秋季覆网保护(第二年春季解除覆网)和完全保护(覆网一直保留到第二年10月调查时为止)均增加了直播种子的存活和出苗率;将种子埋藏在土壤1–4cm也显著增加直播种子的存活率和出苗率,这与鼠类密度(12%)在秋季显著高于春季(2%)有直接关系。此外,坚果树种的种子存活和出苗与其种子特征有一定联系。因此,作者认为如果根据鼠类数量的季节波动规律而采取适当育林措施(如保护和埋藏等)能够增加森林种子直播的效率。     

Competitive suppression of Quercus douglasii (Fagaceae) seedling emergence and growth

Reduced recruitment of blue oak (Quercus douglasii) seedlings in California grasslands and woodlands may result from shifts in seasonal soil water availability coincident with replacement of the native perennial herbaceous community by Mediterranean annuals. We used a combination of container and field experiments to examine the interrelationships between soil water potential, herbaceous neighborhood composition, and blue oak seedling shoot emergence and growth. Neighborhoods of exotic annuals depleted soil moisture more rapidly than neighborhoods of a perennial grass or "no-neighbor" controls. Although effects of neighborhood composition on oak seedling root elongation were not statistically significant, seedling shoot emergence was significantly inhibited in the annual neighborhoods where soil water was rapidly depleted. Seedling water status directly reflected soil water potential, which also determined the extent and duration of oak seedling growth during the first year. End-of-season seedling height significantly influenced survival and growth in subsequent years. While growth and survival of blue oak seedlings may be initially constrained by competition with herbaceous species, subsequent competition with adult blue oak trees may further contribute to reduced sapling recruitment.     

黑龙江省不同地点蒙古栎林生态特点研究

通过对黑龙江省６个地点的天然蒙古栎林的结构和更新特点的分析,蒙古栎林可划分为不同特点的蒙古栎群落,即纯蒙古栎群落、蒙古栎桦树群落、蒙古栎槭树群落、蒙古栎红松群落和蒙古栎松混交林群落,其演替趋势如下：红蒙古栎群落、蒙古栎桦树群落至蒙古栎槭树落、蒙古栎红松群落,再至蒙古栎红松混交林群落,蒙古栎群落类型的多样性主要反应群落不同的演替阶段,造成蒙古栎群落多样的原因是人类活动和自然因素作用的结果,随着群落的演替,蒙古栎的优势地位逐渐被消弱,乔木种类丰富度增多,草本种类丰富度增多;蒙栎的相对密度下降,林内环境由于干燥逐渐变中性至较湿润,蒙古栎幼苗和幼树在总幼苗和幼树中所占的比例下降,耐荫物中色木槭等的幼苗和幼树所占比例上升;在演替过程中,蒙古栎分布格局－聚集分布的聚集程度逐渐降低,并向随机分布的方向发展。     

季节变化对锐齿栎种子扩散的影响

季节变化是影响鼠类扩散植物种子的一个主要环境因素.2008年9-11月(秋季)和2009年3-4月(春季)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锐齿栎种子的扩散,并与种子大小年假说的预测进行了比较分析.结果表明:春季锐齿栎种子的扩散速度比秋季快,平均扩散距离(取食距离和贮藏距离)也比秋季远,该结果与种子...     

Taxonomic scale-dependence of habitat niche partitioning and biotic neighbourhood on survival of tropical tree seedlings

In order to differentiate between mechanisms of species coexistence, we examined the relative importance of local biotic neighbourhood, abiotic habitat factors and species differences as factors influencing the survival of 2330 spatially mapped tropical tree seedlings of 15 species of Myristicaceae in two separate analyses in which individuals were identified first to species and then to genus. Using likelihood methods, we selected the most parsimonious candidate models as predictors of 3 year seedling survival in both sets of analyses. We found evidence for differential effects of abiotic niche and neighbourhood processes on individual survival between analyses at the genus and species levels. Niche partitioning (defined as an interaction of taxonomic identity and abiotic neighbourhood) was significant in analyses at the genus level, but did not differentiate among species in models of individual seedling survival. By contrast, conspecific and congeneric seedling and adult density were retained in the minimum adequate models of seedling survival at species and genus levels, respectively. We conclude that abiotic niche effects express differences in seedling survival among genera but not among species, and that, within genera, community and/or local variation in adult and seedling abundance drives variation in seedling survival. These data suggest that different mechanisms of coexistence among tropical tree taxa may function at different taxonomic or phylogenetic scales. This perspective helps to reconcile perceived differences of importance in the various non-mutually exclusive mechanisms of species coexistence in hyper-diverse tropical forests.     

Mycorrhizas on nursery and field seedlings of Quercus garryana

Oak woodland regeneration and restoration requires that seedlings develop mycorrhizas, yet the need for this mutualistic association is often overlooked. In this study, we asked whether Quercus garryana seedlings in nursery beds acquire mycorrhizas without artificial inoculation or access to a mycorrhizal network of other ectomycorrhizal hosts. We also assessed the relationship between mycorrhizal infection and seedling growth in a nursery. Further, we compared the mycorrhizal assemblage of oak nursery seedlings to that of conifer seedlings in the nursery and to that of oak seedlings in nearby oak woodlands. Seedlings were excavated and the roots washed and examined microscopically. Mycorrhizas were identified by DNA sequences of the internal transcribed spacer region and by morphotype. On oak nursery seedlings, predominant mycorrhizas were species of Laccaria and Tuber with single occurrences of Entoloma and Peziza. In adjacent beds, seedlings of Pseudotsuga menziesii were mycorrhizal with Hysterangium and a different species of Laccaria; seedlings of Pinus monticola were mycorrhizal with Geneabea, Tarzetta, and Thelephora. Height of Q. garryana seedlings correlated with root biomass and mycorrhizal abundance. Total mycorrhizal abundance and abundance of Laccaria mycorrhizas significantly predicted seedling height in the nursery. Native oak seedlings from nearby Q. garryana woodlands were mycorrhizal with 13 fungal symbionts, none of which occurred on the nursery seedlings. These results demonstrate the value of mycorrhizas to the growth of oak seedlings. Although seedlings in nursery beds developed mycorrhizas without intentional inoculation, their mycorrhizas differed from and were less species rich than those on native seedlings.     

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。     

Predating strategy of rodents on acorns of Quercusaliena var. acuteserrata under different predating risks and fate of acorns

The effects of rodents on forest regeneration have been highlighted in many ecological studies. In 2002 and 2003, the acorns of Quercus aliena var. acuteserrata were subjected to 12 different treatments. The daily dynamics and the amount of acorns that were finally left intact, predated in situ, or removed off were examined and documented. The ratios of acorns that were infested by bugs before and after predation by rodents were carefully documented. It was found that: (1) the ratios of acorns infested by bugs before and after predation by rodents showed significant difference (P > 0.05), suggesting that rodents would not prey on acorns during the predating process if acorns had been already infested by bugs. (2) When compared with the controls, the fate of acorns could be roughly classified into four types: acorns that were simply buried or placed on black paper showed no significant differences with the controls in their response to rodents, suggesting that rodents have no sensitivity to the little change of odor resulting from burying and may be more adapted to black background. Acorns attached with strings, dyed with black ink, cut into halves, or placed on white/green/red paper were predated in situ with much lower predation rates and lasted the same duration. Compared with the controls, the acorns were removed away by rodents with the same first day rate and the removal lasted longer. The final removal rate of the acorns by rodents was increased, whereas that of predation in situ was decreased, and none were left in the spot. This may suggest that rodents in this case increased their predation risk expectation and adopted a strategy of “less predation proportion in situ”. Acorns that were burnt were not predated in situ as well as removed in the first day, but the rates were raised to the maximum in the forth and fifth day and then declined to zero. Consequently, the final ratios of predation in situ and removal considerably decreased with many acorns, nearly 50%, being left intact. It can be hypothesized that the rodents responded to the change in odor of acorns by using the strategies of “less predation proportion in situ” and “less predating activities”, thereby resulting in larger number of acorns remained intact. Acorns that were enwrapped were barely predated in situ, removed in the first day with a normal predation rate, which declined to nearly zero in the second day and then increased till it declined again from the fourth day to the final day. The final rate of predation in situ was lower than 5% without acorns remained intact, and the rate of removal of acorns showed a considerable relative increase. This suggests that when there was a change in both odor and status of acorns, rodents also adopted the “spying” and “less predation proportion in situ” strategy, Because the predation risk is critical, the sense of sight would play an important role in rodent's living. (3) Definite amounts of acorns were predated in situ, discarded, removed away, predated ex situ, lost and hoarded separately.     

Litter dynamics in post-fire successional forests of Quercus ilex

Forest floor regeneration after fire in a holm oak forest was studied in a chronosequence in Catalonia (NE Spain). The sampled area represented situations of high fire disturbance because of the frequency of wildfires and the high impact of erosive processes. Forest floor bulk samples from the L, F and H layers were taken in 5 burned plots, aging from 0 to 35 years. They were analyzed for dry standing weight, and organic carbon and N content.Plant cover reached almost 100% two years after the fire. During the first 20 years, shrubs and herbs had the highest surface covering, and thereafter, holm oaks became dominant.Holm oak litterfall stabilized by the fourth year after burning.The major shrub species which were present in the chronosequence had two different patterns with respect to nitrogen: 1) a low litter C/N ratio (i.e. nitrogen fixing species); 2) a high litter C/N ratio.The litter decay coefficients were relatively high, allowing for a rapid structuration and formation of the L and F layers after the fire. Indeed, 95% of the maximum steady standing weight accumulates in 8–9 years. The pattern of cover in the different strata influenced the lateral distribution and variable accumulation of the forest floor layers and determined different turnover rates.     

Abiotic constraints on the establishment of Quercus seedlings in grassland

High evaporative demand and periodic drought characterize the growing season in midwestern grasslands relative to deciduous forests of the eastern US, and predicted climatic changes suggest that these climatic extremes may be exacerbated. Despite this less than optimal environment for tree seedling establishment, deciduous trees have expanded into adjacent tallgrass prairie within the last century leading to a dramatic land cover change. In order to determine the role of light and temperature on seedling establishment, we assessed carbon and water relations and aboveground growth of first‐year Quercus macrocarpa seedlings exposed to one of three conditions: (1) intact tallgrass prairie communities (control), (2) aboveground herbaceous biomass removed (grass removal), and (3) shade plus biomass removal to reduce light (PFD) to levels typical of the grassland‐forest ecotone (shade). In the 2000 growing season, precipitation was 35% below the long‐term average, which had a significant negative effect on oak seedling carbon gain at midseason (photosynthesis declined to 10% of maximum rates). However, net photosynthesis and stomatal conductance in the shade treatment was ca. 2.5 and 1.5 times greater, respectively, than in control treatment seedlings during this drought. During this period, leaf and air temperatures in control seedlings were similar whereas leaf temperatures in the shade treatment remained below air temperature. A late‐season recovery period, coincident with decreased air temperatures, resulted in increased net photosynthesis for all seedlings. Increased photosynthetic rates and water relations in shaded seedlings compared to seedlings in full sun suggest that, at least in dry years, high light and temperature may negatively impact oak seedling performance. However, high survival rates for all seedlings indicate that Q. macrocarpa seedlings are capable of tolerating both present‐day and future climatic extremes. Unless historic fire regimes are restored, forest expansion and land cover change are likely to continue.     

Recruitment and survival of Argyrodendron actinophyllum seedlings in an Australian rainforest

The capacity of seedlings to survive for extended periods beneath intact forest increases the likelihood of regeneration of many species of canopy trees in rainforests. I studied the demographics of Argyrodendron actinophyllum (F.M.Bail.) H.L.Edlin seedlings in a subtropical rainforest in northern New South Wales. A mast seeding of A. actinophyllum was observed and subsequent survival of seedlings monitored over a four year period. Densities of seedlings that emerged correlated with seedfall, while seedfall depended on the size and distance to the surrounding trees. Mortality of seedlings showed density-dependence at higher seedling densities (above about 100 seedlings m^?2), apparently in response to browsing pressure that varied with the density of seedlings. Seedlings that were protected from vertebrates by exclosure cages had lower mortality rates than unprotected seedlings and showed no density response. Glasshouse experiments showed seedling growth was reduced by defoliation, light intensity and initial seed weight, and that seedlings could not persist at light intensities below about 1% ambient, which occur in darker patches on the forest floor. Possible mechanisms whereby the observed spatial and temporal patterns of seedling recruitment could reduce the likelihood of the species becoming more common relative to other tree species in the forest are discussed.     

Effects of Herbivory and Light Conditions on Induced Defense in Quercus crispula

Quercus crispula saplings, and the relation between defense and tolerance involving photosynthesis and growth. We conducted field experiments with herbivorous insects under two light conditions. As an induced defense, the leaf mass per area increased with herbivory regardless of the amount of light available, and the concentration of condensed tannin was significantly higher when light was plentiful. On the other hand, as induced tolerance, the photosynthetic rate decreased with herbivory under conditions of ample light. In conclusion, we found that both the availability of light and herbivory affected defense and tolerance in Q. crispula. In addition, we suggest that the interaction between defense and tolerance was noticeable when much light was available because of the photosynthetic reduction caused by herbivory. Received 21 February 2001/ Accepted in revised form 13 August 2001     

Effect of bryophytes on survival of conifer seedlings in subalpine forests of central Japan

Bryophytes form the major seedbed for coniferous trees in the subalpine forests of central Japan. Field experiments were conducted on the floor of a closed stand dominated byTsuga on Mt Fuji to examine the seedling survival ofAbies veitchii andTsuga diversifolia in seven substrate types in relation to the morphological characteristics of the seeds and seedlings. NeitherAbies norTsuga seedlings could survive on dwarf-bamboo litter beyond 2 years after the seed rain.Abies seedlings survived not only in all types of bryophyte communities but also in larch litter. In contrast, the survival ofTsuga seedlings was restricted to communities of smaller-statured bryophytes found on logs. The seeds and 1st year seedlings ofA. veitchii were larger than those ofT. diversifolia. Abies seedlings produced hypocotyls taller than any bryophyte community and radicles which were long enough to penetrate into the moist humus layer, whereasTsuga seedling radicles presumably penetrate into humus only in bryophyte communities forming a thin mat on logs. The effect of bryophytes on survival of tree seedlings therefore depends on the relationships between the morphology of seeds and seedlings and the structure of bryophyte communities.