植物种群更新的补充限制

补充限制基于生态位理论, 从种子萌发、幼苗存活和生长、繁殖体扩散等生活史阶段的种群统计特征及环境因素(土壤水分、养分、凋落物等)着手, 探讨种群的更新问题。种源限制和微生境限制是补充限制理论研究的核心内容, 但是哪个更为重要并没有统一的结论。种源限制与种子生产、土壤种子库和地下芽库中的繁殖体数量不足有关。其中, 气候的年际波动、土壤种子库寿命和动物的捕食都会影响种子生产在种群更新中的作用; 土壤种子库常被视为种群更新的保险库, 与地上种子雨共同促进种群更新, 但是, 如果土壤里种子具有较高的死亡率和休眠率, 将会降低种子库的作用; 地下芽库及其产生的无性分株对于种群更新的意义更多地体现在干扰后种群更强的恢复能力上。扩散限制是种群更新中的普遍现象, 与种子产量、散布能力、传播媒介、幼苗密度等因素有关。微生境限制主要表现为水分、养分、凋落物等非生物因素以及竞争、捕食等生物因素对种子的活力、萌发性、幼苗的存活力、物质分配等过程的影响, 其重要性随着植物生活史阶段而发生变化。未来需要进行综合的、长期的实验, 并应着重加强种源限制及相关生态过程的进化与生态相结合的机理性研究, 从而更深刻地认识和理解种群更新问题, 建立更为综合、系统的种群更新理论体系。     

动物传播者对植物更新的促进与限制

在动物 植物的关系网络中,传播者对植物更新具有促进与限制两方面作用.本文从种群尺度总结了传播者取食、空间利用等行为对植物更新的影响;从群落尺度分析了多种传播者传播有效性对植物更新的促进与限制.传播者对食物的处理方式决定了种子的命运,且具有明显的种间差异;植物在传播者食谱中的地位亦决定其更新的成败,成为动物偏好的食物可助其摆脱森林破碎化等不利事件的影响.动物的空间行为可导致种子命运发生改变.传播者移动距离能否逃逸同种成树对种子的距离限制,影响种子的更新命运;动物偏好的适宜生境与适于植物萌发生境的空间一致性程度影响传播者传播的成效.有效传播者的非冗余性促使种子传播网络更稳定,利于植物更新;无效传播直接限制植物更新,但为其他植物定殖提供了可用空间.今后应将传播者行为融入植物种群更新研究,而从生态系统服务角度揭示传播者在植被恢复的作用应是未来恢复生态学研究的重点.     

珍稀植物连香树在其中国分布区北缘的种子性状及幼苗更新限制

连香树(Cercidiphyllum japonicum)是第三纪孑遗植物, 存在严重的幼苗更新限制。为验证生活史早期(种子萌发)限制中国分布区北缘连香树种群幼苗更新, 并探讨其主要成因, 本研究在秦岭和太行山脉采集不同种源地的种子, 测定其形态性状、营养元素含量和质量、不同贮存时间的活力及不同温度条件下的萌发性状, 通过方差分析、相关分析等方法对不同种源地的种子性状进行分析。结果表明: 在中国分布区北缘, 其种子长度(P < 0.001)、萌发率(P < 0.001)、平均萌发时间(P < 0.001)、氮(P < 0.05)和磷含量(P < 0.001)在不同种源间存在显著差异; 而在区域尺度上(秦岭与太行山), 仅种子碳含量存在显著差异(P < 0.01)。天水种群的种子萌发率最高(21.77%), 平均萌发时间最长(11.12 d); 栾川的萌发率最低(1.38%), 平均萌发时间最短(3.47 d)。在25℃条件下, 济源种群的种子萌发率显著高于10℃、15℃和20℃条件下(P < 0.05), 而其他种源地的萌发率在不同温度条件下无显著差异。在4个温度条件下, 栾川种群种子的初始萌发时间无显著差异, 而其他4个种源地的初始萌发时间都随温度升高而缩短。相关分析结果表明, 种子萌发率与种子活力密切相关, 而种子活力与种子质量、种子的氮和磷含量显著相关。在中国分布区北缘, 连香树种子的自身属性(质量、氮和磷含量)通过影响种子活力间接影响萌发率; 且种子萌发对温度的响应主要表现在萌发时间上。本研究证实种子萌发是限制连香树种群幼苗更新的关键阶段, 主要原因如下: (1)连香树种子在9月成熟后, 10月的温度仍适宜种子萌发, 但较短生长期的幼苗在冬季低温下不能存活; (2)连香树种子萌发率低(14.4%); (3)第二年春天种子活力骤降。     

濒危植物银缕梅(Parrotia subaequalis)不同径级个体的光合能力差异与更新限制

银缕梅(Parrotia subaequalis)是我国华东地区特有种,属国家Ⅰ级濒危保护植物.分别于5月份与8月份采用Li-6400光合系统测定了江苏宜兴龙池大垅西芥银缕梅种群内大 (胸径＞10 cm)、中 (2 cm＜胸径≤10 cm)、幼 (胸径≤2 cm) 3个径级个体的光合生理参数、光合-光强响应曲线及光环境因子,探讨了不同径级个体的光合能力差异及其对种群更新的影响.结果表明:幼树最大净光合速率(Pnmax)、光饱和点(LSP)、光补偿点(LCP)和暗呼吸速率(RD) 显著低于大树与中树,但比叶面积(SLA)显著高于大树及中树;低LCP、低RD与高SLA是银缕梅幼树对弱光环境做出的适应性反应.中树LSP仅为大树的二分之一,而其LCP与大树无明显差异,且显著高出幼树;中树LSP与LCP之间的光能利用区间极为狭窄;中树全天平均接受的RAR稍高于LCP,但远低于其LSP;同时,中树的SLA显著低于幼树与大树,致使中树全天碳同化总量严重不足.基于净光合速率(Pn)与光环境参数建立的逐步回归方程进一步证实了光合有效辐射(PAR)对Pn的贡献最大,是影响银缕梅光合碳同化能力的关键因子.由于幼树与中树受到大树遮荫,叶片上方的光合有效辐射均显著降低.尽管银缕梅幼苗在林下表现出较强的弱光适应能力,但在后期建成为中树的过程中,叶片形态与光合生理特性均表现出对遮萌环境的不适应,光资源不足构成银缕梅种群更新的主要限制因子,导致种群的中等径级个体急剧减少.人为干预以形成适宜的林窗环境,有利于银缕梅幼树的后期生长,并促进银缕梅种群的顺利更新.     

啮齿动物捕食和搬运蒙古栎种子对种群更新的影响

2010-2011年,在东北林业大学城市林业示范基地研究了啮齿动物对蒙古栎种子的捕食和搬运。结果表明,蒙古栎种子捕食率在年际间无显著差异,但2010年的蒙古栎种子搬运率显著高于2011年,存留率显著低于2011年。啮齿动物在胡桃楸林、樟子松林、水曲柳林和白桦林内对蒙古栎种子的总捕食率和总搬运率分别达到(5.7±13.5)%和(27.1±37.1)%。蒙古栎种子在樟子松林内的捕食率和搬运率均最高,胡桃楸林内蒙古栎种子存留率最高;2010年蒙古栎种子在与蒙古栎林边缘距离0,10,20,30,40,50m组间的捕食率无显著差异,搬运率和存留率有显著差异,但2011年蒙古栎种子在不同距离组间的捕食率呈显著差异,搬运率和存留率无显著差异。与蒙古栎林边缘距离20m处蒙古栎种子的捕食率最高。2010年和2011年,分别有(37.8±49.7)%和(27.7±49.8)%的蒙古栎种子被啮齿动物利用,这表明啮齿动物是林业示范基地内蒙古栎地表种子的主要捕食者。冬季食物匮乏秋季贮藏种子是造成啮齿动物对蒙古栎种子有较大捕食、扩散压力的主要原因。因此,啮齿动物对蒙古栎种子的捕食和搬运影响了蒙古栎林的种群更新。     

濒危植物长柄双花木自然种群结实的花粉和资源限制

长柄双花木（Disanthus cercidifolius Maxim．vsr．longipes H．T．Chang）只零星分布于湖南、江西和浙江等省的少数县,已处于濒危状态。通过对其野外自然种群试验,研究了花粉和资源的有效性对雌性生殖的影响,并进一步探讨了长柄双花木“花多果少”的生殖机制。结果表明：不同来源的花粉对长柄双花木结果率及结籽率均产生显著影响。花粉来源而不是花粉数量对长柄双花木结实存在显著影响。自然条件下,长柄双花木“花一果转化率”约为54馏,“胚珠-种子转化率”约为83．19。施肥处理有利于花芽发育,可使植株花芽败育率明显下降,开花比例升高;同时座果率和结籽率均显著提高,而果实败育率则显著降低。随剪除叶片量的增加,结果率明显降低,但果实败育率在处理间无显著差异;剪除弱小枝条及病虫枝后,个体总开花数虽有所下降,但是结果率和结籽率均显著提高。去除花后,长柄双花木平均单果重、单粒种子重均显著降低。营养水平的时空异质性使自然种群结籽率受到资源和花粉的限制;补充资源可以通过增强花对传粉者的吸引而间接影响传粉。长柄双花木自然种群的坐果率极低,而且存在严重的“大小年”结果现象,不同的因子可能对其低水平坐果率产生相互作用。选择性败育、子房供应和雄性功能假说似乎是长柄双花木“花多果少”的生殖策略的最合理解释。     

动物分散贮食行为对植物种群更新的影响

分散贮食是许多动物取食行为策略的重要组成部分。对以植物种子为主要贮食对象的动物来说,种子内营养物质含量、种子大小以及种子内次生化合物的含量等因素都直接影响动物的贮食行为。动物偏爱贮藏个体较大的种子,大种子多被搬运并分散贮藏在远离种源的地方,而小种子则多被就地取食,以补充动物贮食过程中的能量消耗。贮食动物主要通过空间记忆、特殊路线以及贮藏点周围的直接线索等途径重新获取贮藏点内食物。在重取过程中,一些贮藏点被遗忘,其中的种子成为植物种群更新的潜在种子库。因此,分散贮食动物不仅是种子捕食者,还是种子传播者,它们对植物种子的捕食、搬运和贮藏,影响了植物种子的存活和幼苗的建成,从而在一定程度上影响植物种群的更新、分布。植物种群为了促进种子的传播,在进化过程中逐渐形成了形式多样的适应性策略,降低种子的直接被捕食率,提高种子的被贮藏率。研究动物分散贮食行为对植物种群更新的影响,将有助于理解贮食动物与植物之间的互惠关系,从而认识贮食动物种群在生态系统中的作用,为生物多样性的保护提供科学依据。     

画稿溪国家级自然保护区珍稀植物桫椤种群结构与更新

根据四川画稿溪国家级自然保护区桫椤(Alsophila spinulosa)种群分布特点,设立4个样地1个样带共44个样方(20 m×20 m),从种群密度、生命表和生境关联(草本层郁闭度和林分受干扰强度)等方面研究了保护区桫椤种群的结构特征。结果表明,每个桫椤种群样方的平均密度为0.088株/m2,幼苗密度为0.049株/m2;桫椤种群的死亡率和消失率曲线变化基本一致,发育过程中出现3个死亡高峰值,分别在第1龄级(H0.01 m)、第7龄级(3.90 m≤H4.80 m)和第10龄级(H≥6.60 m)阶段;种群生存率单调下降,累计死亡率单调上升,第7龄级以后,种群生存率小于1%;根据种群生存率函数Si、累计死亡率函数Fti、死亡密度函数fti和危险率函数λti4个生存函数曲线表明,画稿溪桫椤种群具有前期锐减、中期稳定、后期衰退的特点。桫椤各龄级阶段密度与草本层郁闭度均无显著相关(p≥0.18),然而桫椤幼苗(第1龄级)以上的个体密度、第7龄级密度与干扰度之间呈显著负相关(p0.05)。综合分析表明,画稿溪桫椤种群具有一定的自然更新能力,但人为活动的干扰可能是限制桫椤种群的因素之一。     

黄檗的更新特点及食果实鸟类对其种子的传播

在帽儿山实验林场和哈尔滨实验林场 ,黄檗母树下没有幼苗分布 ,不能进行天然更新 ,需要靠食果实鸟类等将果实和种子传播至远离母树的其他林下。捕食黄檗果实的食果实鸟类有 9种。其中 6种是食果肉鸟类 ,吞入果实后 ,消化果肉 ,而种子完整地随粪便排出而得以传播。其余 3种是食种子鸟类 ,没有传播作用。果实在鸟类消化道内的滞留时间达 2 0～ 30min ,具有很长的潜在传播距离。将鸟类消化后的种子与完整果实和人为去果肉种子进行萌发对比实验 ,消化后种子的累计萌发率与其余二者之间均没有显著性差异 ,说明食果实鸟类的消化 (道 )过程对种子萌发没有明显影响 ,同时证明果肉对种子萌发率没有显著影响 ,果肉中不含萌发抑制物质。黄檗提供多种鸟类以食物 ,而鸟类也同时以多种肉质果植物为食物。因此食果实鸟类和肉质果植物 (包括黄檗 )之间形成了松散的互利共生关系     

食种子动物对三种锦鸡儿属植物繁殖更新的影响

锦鸡儿属(Caragana)植物是黄土高原植被的重要组分,具有固氮、耐旱的特性,对黄土丘陵沟壑区的植被恢复具有重要意义。作者于2002–2003年对该属本地种白毛锦鸡儿(C.licentiana)和甘蒙锦鸡儿(C.opulens)及外来种中间锦鸡儿(C.intermedia)种子被动物取食情况进行了比较研究。2003年白毛锦鸡儿种子散落前的虫害率为88.2%,甘蒙锦鸡儿为29.7%±1.7%,中间锦鸡儿为43.2%±4.8%。种子落地后,白毛锦鸡儿和甘蒙锦鸡儿的种子被小啮齿动物取食的比例在各种微生境下均为100%,且取食速度很快,在种子放置后4d内即全部被取食。中间锦鸡儿种子被取食率则因微生境的不同而不同:在灌丛下被取食率为100%,在半阳坡的开阔草地上为85.0%±10.0%,在种植植物未成功的水平阶地上为31.0%±8.7%,且取食速度相对较慢。研究结果表明,食种子动物的取食大大减少了白毛锦鸡儿和甘蒙锦鸡儿的种子量以及种子传播萌发的机会。但食种子动物可能充当了白毛锦鸡儿4.0%硬实种子的传播者,从而有助于白毛锦鸡儿实现种子途径的更新;中间锦鸡儿种子无论在散落前还是散落后受食种子动物的影响均很小。     

Seedling establishment in an epiphytic orchid: an experimental study of seed limitation

Will increased fruit and seed production in a severely pollination-limited orchid stimulate population growth? We tested whether safe sites for germination and seedling establishment are limiting for the twig epiphyte, Tolumnia variegata, by manipulating fruit set and monitoring subsequent seedling establishment for two seasons (1991–1992, 1992–1993). In the Cambalache Forest Reserve of Puerto Rico, we established 36 plots along a transect. Each plot consisted of nine trees. A center tree was designated as the site for attaching Tolumnia and manipulating fruit set. The other eight potential host trees were 1–3 and 3–5 m from the center tree in each of the cardinal directions. A 1-m length of stem 1 m from the ground was monitored for recruits on each of the nine trees of 24 fruit-enhanced plots and 12 controls (23 and 13, respectively for the 1992–1993 season). Fruit enhancement plots were divided among two treatments: one-fruit and five-fruit additions for the 1st year and one to five and more than five fruits for the 2nd year. Availability of suitable host species was not limiting. T. variegata showed little specificity for host tree species, good host trees and shrubs were common, and there was no evidence that the orchid had a preference for small branches, despite possessing the entire suite of characteristics thought to respresent “obligate” twig epiphytes. Fruit enhancement increased seed rain and seedling establishment consistently in only the high-fruit treatment plots. Most recruitment occurred near fruiting plants. Over the 2-year period, mortality was 18% for adults and 85.5% for the 1991–1992 cohort of recruits. Net recruitment was positive for both the treatment (average = 1.74) and control plots (average = 0.67). Seedling establishment at our study site was not microsite-limited. If selection for increased pollinator attraction occurs, then an increase in seed output should result in population growth.     

Consumer pressure, seed versus safe-site limitation, and plant population dynamics

Plants often suffer reductions in fecundity due to insect herbivory. Whether this loss of seeds has population-level consequences is much debated and often unknown. For many plants, particularly those with long-lived seedbanks, it is frequently asserted that herbivores have minimal impacts on plant abundance because safe-site availability rather than absolute seed number determines the magnitude of future plant recruitment and hence population abundance. However, empirical tests of this assertion are generally lacking and the interplay between herbivory, spatio-temporal variability in seed- or safe-site-limited recruitment, and seedbank dynamics is likely to be complex. Here we use a stochastic simulation model to explore how changes in the spatial and temporal frequency of seed-limited recruitment, the strength of density-dependent seedling survival, and longevity of seeds in the soil influence the population response to herbivory. Model output reveals several surprising results. First, given a seedbank, herbivores can have substantial effects on mean population abundance even if recruitment is primarily safe-site-limited in either time or space. Second, increasing seedbank longevity increases the population effects of herbivory, because annual reductions in seed input due to herbivory are accumulated in the seedbank. Third, population impacts of herbivory are robust even in the face of moderately strong density-dependent seedling mortality. These results imply that the conditions under which herbivores influence plant population dynamics may be more widespread than heretofore expected. Experiments are now needed to test these predictions. Received: 3 November 1999 / Accepted: 15 February 2000     

Recruitment limitation: A theoretical perspective

Abstract A theoretical analysis of the concept of recruitment limitation leads to the conclusion that most populations should he regarded as jointly limited by recruitment and interactions between individuals after recruitment. The open nature of local marine systems does not permit avoidance of density-dependent interactions; it simply may make them more difficult to detect. Local populations consisting of settled organisms may not experience density-dependent interactions under some circumstances, but the entire species population consisting of the collection of local populations and their planktonic larvae must have density-dependent dynamics. Any local population of settled individuals can escape density dependence if sufficient density dependence occurs among planktonic larvae or within other local populations. Common conceptions of density dependence are too narrow, leading too often to the conclusion that it is absent from a system. It is equally wrong to expect that density-dependent interactions after settlement determine local population densities independently of recruitment. Special circumstances allowing density dependence to act strongly and quickly are needed before density dependence can neutralize the effects of recruitment. Recruitment limitation and density-dependent interactions therefore should not be regarded as alternatives but as jointly acting to determine the densities of marine benthic populations. Moreover, the interaction between fluctuating recruitment and density dependence is potentially the most interesting feature of recruitment limitation. For example, this interaction may be an important diversity-maintaining mechanism for marine systems.     

Dispersal and establishment filters influence the assembly of restored prairie plant communities

Community assembly filters, which in theory determine the suite of species that arrive at and establish in a community, have tremendous conceptual relevance to restoration. However, the concept has remained largely theoretical, with a paucity of empirical tests. As such, the applicability of assembly filters theory to ecological restoration remains incompletely known. We tested the relative strengths of dispersal and establishment filters by comparing the plant species composition, measured by species' presence/absence, in 29 restored prairies with the seed mixes used to restore each prairie. We found that both establishment and dispersal filters limited prairie similarity to the seed mix. Sown species responded differentially to filters, with a few species limited only by dispersal (seed density), many others limited only by establishment conditions (i.e. organic matter and sand content of soils, land use history, and fire frequency), and others limited by both dispersal and establishment filters. A few species, typically those sown most often, were not restricted by dispersal or establishment filters, likely because they were sown in high enough densities and all sites had suitable environmental conditions. Finally, one group of species established poorly, but we could not attribute this to either dispersal or establishment filters. This information can help land managers select species likely to establish in restorations when sown at sufficient densities. These results illustrate that dispersal and establishment filters limit the establishment of species in restored communities and these filters are species‐dependent. Identifying the most limiting filter(s) for species will inform strategies to increase their establishment success.     

Plant recruitment bottlenecks in temperate forest fragments: seed limitation and insect herbivory

Numerous studies have documented declines in plant diversity in response to habitat loss in fragmented landscapes. However, determining the mechanisms that lead to species loss is challenging using solely a correlative approach. Here we link correlative assessments of plant community composition with seed additions for a focal species to test the hypothesis that distributions of forests plants within a fragmented landscape are limited by seed dispersal. Woody plant species richness of fragments declined as fragments (n=26) became more isolated by agricultural fields. We predicted that if these isolation effects were driven by poor dispersal rather than other effects associated with habitat loss, then plants should vary in their response to isolation in relation to their seed size (i.e., stronger effects for plants with larger seeds). As predicted under this dispersal limitation hypothesis, sensitivity of bird-dispersed shrubs to isolation was related to their seed mass, with species with heavy seeds (e.g., Lindera benzoin) exhibiting stronger declines in presence across isolation gradients than species with light seeds. Seed addition experiments were performed for Lindera benzoin in two high isolation forest fragments (nearest neighbor mean distance=803 m) where Lindera was naturally absent, and two low isolation fragments (nearest neighbor mean distance=218 m) with naturally occurring Lindera populations. Seed addition and control plots (n=50 1 m² plots per fragment) were monitored for 13 censuses over 3 years. Across all four fragments, seed additions resulted in significant increases in Lindera seedling recruitment with no differences in final seedling establishment among fragments. However, insect herbivory was higher on Lindera seedlings in high isolation compared to low isolation fragments and was negatively correlated with seedling survival over some years. Consistent with prior work, our results confirm that seed dispersal plays a significant role in affecting plant diversity in fragmented landscapes. However, results also suggest the need for a better understanding of how additional processes, such as herbivory, may be altered as habitat is lost and what effects such changes have for forest plants.