子午岭马兰林区辽东栎幼苗定居限制的初步研究

在子午岭马兰林区选择油松林、辽东栎林、灌丛、草地等四种群落,通过播种实验及3 a的跟踪调查,研究四种不同生境下辽东栎幼苗在定居过程中的生长规律.结果显示:在四种生境中,人为增加辽东栎种子密度,均能明显提高出苗量,幼苗生长三年后大量死亡,死亡高峰出现在第三年.郁闭生境下(油松林和辽东栎林),辽东栎出苗率及存活率高于开阔生...     

灌丛高度对啮齿动物贮藏和扩散辽东栎坚果的影响

本文选择高、矮两类灌丛生境, 于2002 年至2003 年在其中释放标记的辽东栎坚果, 并连续记录释放坚果的命运, 以了解生境差异对啮齿动物搬运、取食和贮藏辽东栎坚果的影响以及啮齿动物对微生境的选择等。研究结果表明, 大林姬鼠等啮齿动物对辽东栎坚果的短期取食非常强烈, 而分散贮藏的量则相对较少; 啮齿动物对辽东栎坚果的搬运距离在矮灌丛生境中显著大于高灌丛生境; 啮齿动物倾向于选择灌丛边缘和灌丛下方取食或贮藏辽东栎坚果, 这类微生境也有利于埋藏坚果的萌发。     

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。     

北京东灵山区两种生境条件下辽东栎幼苗补充与建立的比较

 北京东灵山区的人工油松（Pinus tabulaeformis）林中常常混生有萌生的辽东栎（Quercus wutaishanica）种群。局部地带辽东栎与油松形成混交林,在山脊及山坡上部一些特殊生境辽东栎甚至取代油松林成为优势种。为了探讨北京东灵山区辽东栎林的天然更新机制及辽东栎幼苗在人工油松林中的天然更新,在油松人工林的林下和林缘两种生境条件下将辽东栎种子播种于2 cm深的土中,对辽东栎幼苗的补充和建立进行了对照试验。结果显示,在辽东栎结实丰年,两种生境条件下播种后辽东栎种子丢失差异并不显著,而辽东栎幼苗补充和建立方面的差异极显著。由于森林动物特别是啮齿类动物的活动对辽东栎幼苗建立的影响具有二重性：消耗种子与促进萌发,本试验特别关注辽东栎幼苗子叶丢失现象及其丢失后果。由于辽东栎种子萌根较早,在幼苗出土前其粗大的主根长达10～20 cm,并已初步形成根系,表明已有相当比例的营养物质从子叶转移到根部。因此当幼苗出土后如果仅仅只是子叶的丢失对辽东栎幼苗的成活和生长均无明显的影响。通过研究发现辽东栎幼苗期子叶丢失有3种形式：1）仅丢失子叶,幼苗其它部位未受伤害,多发生在林缘生境中,占发芽坚果的29.69%;2）整个幼苗连根被拖出地面,多发生在林下生境,占62.43%;3）在子叶与根颈的连接处主干被咬断,林缘为50.88%,显著高于林下的8.41%。但只有后两种形式才导致幼苗死亡。导致这些现象的原因是由于林缘土壤腐殖质含量低,比较干燥,土质较坚实,以及幼苗出土前坚果主根生长得较长等缘故,当幼苗遭受动物捕食时不会轻易地被拖出地面,拖走的往往仅仅是子叶。至生长季节结束,林缘样方辽东栎幼苗的成活率接近20%,茎干平均高度7.94 cm,芽数11.24·株-1,叶片平均干重为81.14 mg·株-1,且几乎全部由丢失子叶的幼苗所构成;而林下样方辽东栎幼苗的成活率不足2%,茎干平均高度4.74 cm,芽数7.52·株-1,叶片平均干重仅为42.27 mg·株-1,以未丢失子叶的幼苗为主。试验结果表明,林缘及类似林缘的环境条件更有利于辽东栎的实生更新。     

黄土高原马栏林区优势种幼苗与其种群径级结构的演替研究

对黄土高原马栏林区几种典型群落类型中优势种辽东栎(Quercus liaotungensis)、油松(Pinus tabulaefor-mis)、白桦(Betula platyphylla)和山杨(Populus davidiana)的幼苗和种群径级结构进行了研究。结果表明:(1)辽东栎幼苗在马栏林区7种典型群落类型中均有分布,且数量充足,实生苗更新良好,辽东栎在不同的群落类型中表现出不同的种群径级结构。在油松 辽东栎混交林和辽东栎林中趋于稳定型;在油松林、油松 白桦混交林、白桦林、山杨林和白桦 山杨混交林中则为增长型;(2)油松幼苗在油松 白桦混交林、油松林和油松 辽东栎混交林中分布较多,但均少于辽东栎幼苗。在这些群落类型中油松种群的径级结构属于稳定型;(3)白桦和山杨幼苗数量较少,无正常更新能力,种群径级结构趋于衰退,以白桦和山杨为主的群落类型将逐渐被以油松和辽东栎为主的群落类型所取代;(4)马栏林区优势种幼苗分布的广度和丰度受到光照、水和灌草层等非生物和生物因素及其相互作用的影响。表明在黄土高原马栏林区以天然恢复为主的辽东栎种群,将在该地植被的自然恢复过程中产生重要作用;油松种群不仅过去而且将来也同样会在该地区的植被恢复中发挥重要作用。     

鼠类对山杏和辽东栎种子的贮藏

在北京市东灵山地区,选择灌丛生境,人工释放山杏和辽东栎种子,对两种种子在释放处被鼠类就地取食、存留动态、种子被搬运和贮藏的距离、位置、状态、微生境等进行了分析比较。采用活捕饲喂和咬痕比较的方法,对野外取食山杏和辽东栎种子的鼠类进行了鉴定。结果表明：1)取食山杏和辽东栎种子的鼠类主要是大林姬鼠和岩松鼠;2)山杏种子被鼠类搬运的平均距离大于辽东栎种子,两种种子大多数被搬运至21m之内,在9m之内最为集中;3)山杏种子的中位存留时间较辽东栎种子长;4)在释放处,鼠类对辽东栎种子的就地取食强度(54．83％)大于对山杏种子的取食(0．17％);5)在搬运之后,大部分山杏种子被埋藏起来,仅有3粒被取食;而辽东栎种子大部分被取食;6)鼠类趋向于把种子搬运到灌丛下方或灌丛边缘进行贮藏或取食。山杏种子在贮藏点的数量为1—3粒,而辽东栎种子在贮藏点的数量均为1粒。     

黄龙山油松林和油松+辽东栎混交林物种组成及优势种群动态

根据黄龙山油松(Pinus tabulaeformis)林和油松+辽东栎(Quercus liaotungensis)混交林共6个群丛中27个典型样地调查资料,分析了松栎林群落物种组成、多样性指数和优势种群动态.结果表明:(1)松栎混交林中灌木层和草本层植物种数均大于油松林,且灌木和草本种类因群落生境不同而有差异.(2)物种多样性分析显示,油松林和松栎混交林中由于生境不同导致灌木层物种多样性差异最大.(3)优势种群分析表明,油松林中阴坡幼苗数量较多,种群最大的为Ⅳ和Ⅴ龄级个体;松栎混交林中油松和辽东栎幼苗充足,到Ⅱ、Ⅲ龄级二者均大量减少,此后油松个体增加而辽东栎减少,说明松栎林发育过程中各层次植物物种相互制约,优势种群之间也存在竞争,且受生境条件影响较大.(4)主成分分析表明,气温、土壤、光照和总郁闭度是主要影响因子.因此,在未来森林抚育中应适当减少优势种群密度,促进异龄化;采取有效措施合理搭配灌木和草本,强化对森林发育有利的因素,提高林分质量.     

黄土高原油松和辽东栎林下植物β多样性差异及影响因素

β多样性是生态学研究的热点论题,相同的β多样性格局可能由不同的生态过程所决定。该文通过构建零假说模型和典范变异分解的方法,比较了黄土高原油松人工林(Form.Pinus tabulaeformis)和辽东栎天然林(Form.Quercus wutaishanica)林下植物群落β多样性,确定了环境过滤和扩散限制在β多样性形成过程中的相对重要性。结果表明:(1)油松人工林和辽东栎天然林的林下群落均具有相似的β多样性大小,均呈现明显的种内聚集过程。(2)环境和空间共同解释了两种林型林下物种组成的较大变化(草本层[E+S]为33%~45%;灌木层[E+S]为21%~35%),且主要以环境解释量为主(包括纯环境变量[E|S]和空间化的环境变量[E∩S])。(3)油松人工林和辽东栎天然林林下群落β多样性形成的不同过程主要体现在生境异质性差异以及特定的环境因子间,例如,海拔梯度、枯落物厚度以及土壤养分(速效氮和速效钾)是引起两种林型林下物种差异的显著环境因子。综上结果认为,黄土高原的油松人工林和辽东栎天然林的群落构建均由确定性的环境过滤为主导,但两种林型的环境过滤过程并不一样,主要表现为生境异质性的差异和不同的环境限制因子。对于黄土高原区域生态环境建设,首先应该保证不同群落的生境异质性,同时不能忽视扩散限制及其与生境异质性二者交互作用对β多样性的影响。     

植物种群更新限制——从种子生产到幼树建成

更新限制是指种子由于各种原因,不能够萌发并生长成幼树。它作为解释生物多样性的理论,一直受到国内外群落生态学家关注。从种源限制、传播限制和建成限制3个角度,对更新限制机制研究进展进行了综述。从种源限制而言,时空因素是影响植物种群更新限制的重要因素,因为植物结实量存在明显时空变化,造成植物更新个体出现明显的时空规律。从传播限制而言,传播数量、距离和食果动物行为均限制植物种群更新。数量上,缺乏有限传播者势必减少传播数量,但如果种子拥有较高质量,则能逃脱数量限制;距离上,植物更新个体显示出明显的Janzen-Connell格局,但传播距离趋向稳定,形成植物种群的进化稳定对策;食果动物行为上,不同传播者对更新贡献存在差异,捕食者直接降低更新,融入两类动物行为的模型更能反映食果动物对更新的限制。从建成限制而言,环境因子制约植物生长。小尺度下,微生境的好坏对于植物幼苗建成至关重要;大尺度下,植物提供较好的广告效应则能摆脱生境限制。将传播者行为、捕食者行为与幼苗的空间分布格局、种子传播机理模型等结合,建立植物更新限制机理模型应是更新限制未来的研究热点。选择稀有种和古老种为主题的长期更新限制研究,为种群恢复提供指导,也是未来重要研究方向。     

繁殖体与微生境在退化草地恢复中的作用

退化草地的成功恢复主要依赖于种子和母株无性繁殖幼苗的有效建植 ,即草地群落中可利用繁殖体是退化草地得以恢复的内在条件。此外 ,群落中那些提供种子发芽、幼苗生长发育的适宜微生境 (safe sites/ suitable microsites) ,构成了退化草地恢复的外在条件。由于严重退化草地群落缺乏可利用繁殖体和供繁殖体生长发育的适宜微生境 ,使得退化草地恢复受到很大限制 ,因而 ,同时满足繁殖体与微生境是退化草地恢复的先决条件。人为提供繁殖体和适宜微生境可以在很大程度上提高退化草地的恢复速度 ,即在缺乏繁殖体草地群落供给繁殖体 ,或者在缺乏微生境的草地群落中创造适宜微生境。不同植物种群建植需要的环境存在着显著差异 ,因此在人工恢复草地群落过程中 ,对这些植物的繁殖体和繁殖体着床环境给予特殊处理是必需的 ,使之同时满足多种植物种群建植需求。对退化草地植物繁殖体、微生境的重要性及其涵义进行讨论     

Experimental Assessment of Factors Limiting Seedling Recruitment of an Amazonian Understory Herb

Understory herbs are an important and species-rich component of tropical forests, but little is known about factors limiting recruitment of these herbs. In a Peruvian flood plain forest, we studied seedling recruitment of the widespread clonal herb Heliconia metallica . We analyzed natural recruitment and experimentally added seeds, excluded predators and removed the understory vegetation at occupied and unoccupied sites to test whether seed limitation, predation, disturbance, and the presence of mature conspecific plants influence seedling recruitment. The number of naturally recruited seedlings was higher at flooded sites and close to flowering H. metallica ramets. Predation caused considerable seed loss shortly after sowing. Seed survival strongly increased with the openness of a site, but was not affected by the presence of mature conspecifics. Seed addition increased the number of seedlings at both occupied and unoccupied sites. Two years after sowing, the number of established seedlings and their size increased with light availability at a site. In experimentally disturbed plots, light availability, seedling establishment, and survival were higher. The effect of disturbance on seedling survival was stronger at occupied sites, probably due to strong light competition from mature H. metallica plants. Herbivore damage on leaves of seedlings was lower at occupied sites, but seedlings grew faster at unoccupied sites. We conclude that seedling recruitment of H. metallica depends on natural disturbance and is limited by both seed availability and the presence of mature conspecific plants.     

Feedbacks between canopy composition and seedling regeneration in mixed conifer broad‐leaved forests

To address the role of canopy‐seedling feedbacks in the structure and dynamics of mixed conifer broad‐leaved forests in the eastern US, we monitored seedling regeneration patterns and environmental conditions in the understorey of stands dominated by either hemlock (Tsuga canadensis) or red oak (Quercus rubra) for three years. Hemlock seedlings were favoured over other species’ seedlings in hemlock stands (a true positive feedback), due to a combination of high seed inputs, high seedling emergence and relatively high seedling survival during the growing season, which allowed hemlock to remain dominant under its own canopy. Red oak stands favoured a suite of mid‐successional broad‐leaved species over hemlock. A more even age structure of broad‐leaved species in red oak stands revealed that high seedling survival in such stands were driving this feedback. Canopy‐mediated variations in both understorey light availability (1.5% for hemlock vs 3.5% for red oak) and soil pH (3.9 for hemlock vs 4.4 for red oak) were found to be the primary correlates of stand‐level differences in seedling regeneration dynamics. In mixed temperate forests in the eastern US, canopy‐seedling feedbacks could act to slow successional trajectories and contribute to the maintenance of a stable landscape structure over many generations.     

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.     

Seed dispersal and seedling recruitment of trees at different successional stages in a temperate forest in northeastern China

Aims Spatial distribution of adult trees in a forest community is determined by patterns of both seed dispersal and seedling recruitment. The objectives of our study were to understand the processes of seed dispersal and seedling recruitment of dominant tree species in a temperate forest of northeastern China and to identify the factors constraining seed dispersal and seedling establishment at different stages of forest succession.Methods During three summer and autumn sessions between 2006 and 2008, altogether 113080 seeds from 22 different tree species were collected in three large field plots representing different forest types in the Changbai Mountain region of northeastern China. The spatial distribution of seed abundance was analyzed using a Syrjala test. Regeneration success of nine major tree species was assessed using variables defining 'limitations' in 'seeds' and 'seedling establishment'.Important findings We found that seed production fluctuated between years and varied greatly with forest types. Four tree species, Acer spp., Fraxinus mandshurica, Tilia amurensis and Betula spp., had the greatest seed production and the widest range of seed dispersal, whereas Quercus mongolica showed the most sustained seed production pattern. The spatial patterns of seed abundance differed significantly among forest types and years. The tree species investigated in this study differed in the degree of seed limitation, as well as in limitation of seedling establishment. There were both negative and positive correlations between seed density and seedling density, depending on site and parental tree density. Seeds of 16 tree species were found in the Populus davidiana–Betula platyphylla forest (PBF) plot, 11 in the conifer and broad-leaved mixed forest (CBF) plot but only 8 in the broad-leaved-Korean pine mixed forest (BKF) plot. The number of seed-contributing species was not only greater in the secondary forests (CBF and PBF plots) than in the primary forest (BKF plot) but was also more variable during the 3 years of assessment. Results from the correlations between seed density and seedling occurrence and that between parental tree density or seed weight and dispersal limitation confirm our intuitive expectations, i.e. heavy seeds had greater dispersal limitation but higher establishment success than light seeds.     

Factors limiting rosette recruitment in scarlet gilia, Ipomopsis aggregata: seed and disturbance limitation

We performed a factorial combination of seed additions, surface soil disturbance, and protection from potential seed predation in experimental plots to elucidate the relative importance of each factor in the recruitment of scarlet gilia seedlings. We detected a significant interaction between seed addition and surface soil disturbance on seedling recruitment over 2 years of establishment. Plots that received both supplemental seed addition and disturbance established 10.5 times as many rosettes as control plots. We did not detect main or interactive effects of caging on seedling establishment. We explored the importance of density-dependent recruitment by investigating the number of rosettes per experimental plot through time. In addition, we used planned contrasts to compare the absolute and percent changes in control and treatment plot rosette densities between the study years. We found significant downward concave curvature to the seedling recruitment curve, suggesting negative density-dependent recruitment across the experimental range of rosette densities. We found a significant difference between the control plots and the seed addition-disturbance plots in terms of absolute changes in plot density from 1997 to 1998. Greater net mortality occurred in plots that received both seed addition and disturbance. However, we found no differences among treatments in the percent change in rosette density across the study years. Our study demonstrates that seedling recruitment in natural plant populations may be limited by the interaction of seed and microsite availability, and that seedling recruitment in scarlet gilia may be negatively density dependent. Our results suggest that the fecundity effects of particular plant-animal interactions (e.g., pollination, herbivory) may affect scarlet gilia population dynamics, particularly under conditions of high soil disturbance. Received: 11 June 1999 / Accepted: 24 November 1999     

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.     

Establishment of tree seedlings in the understory of restoration plantations: natural regeneration and enrichment plantings

Little is known about the potential of restoration plantations to provide appropriate understory conditions to support the establishment of seeds arriving from neighboring native forests. In this article, we investigated how seedling establishment is affected in the understory of restoration sites of different ages and assessed some of the potential environmental factors controlling this ecological process. We first compared the density and richness of native tree seedlings among 10‐, 22‐, and 55‐year‐old restoration plantations within the Atlantic Forest region of southeastern Brazil. Then, we undertook a seed addition experiment in each study site, during the wet season, and compared seedling emergence, survival, and biomass on local versus old‐growth forest soil (transferred from a reference ecosystem), in order to test whether local substrate could hamper seedling establishment. As expected, the oldest restoration site had higher density and richness of spontaneously regenerating seedlings. However, seedling establishment was less successful both in the oldest restoration planting and using substrate transferred from a reference ecosystem, where emergence and survival were lower, but surviving seedlings grew better. We attribute these results to lower light availability for seedlings in the understory of the oldest site and speculate that higher incidence of pathogens on old‐growth forest soil may have increased seedling mortality. We conclude that the understory of young restoration plantations provides suitable microsite conditions at the early establishment phases for the spontaneous regeneration or enrichment planting of native trees.     

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.