Effects of local biotic neighbors and habitat heterogeneity on seedling survival in a spruce‐fir valley forest,northeastern China

Seedlings are vulnerable to many biotic and abiotic agents, and studying seedling dynamics helps understand mechanisms of species coexistence. In this study, the relative importance of biotic neighbors and habitat heterogeneity to seedling survival was examined by generalized linear mixed models for 33 species in a spruce‐fir valley forest in northeastern China. The results showed that the relative importance of these factors varied with species and functional groups. Conspecific negative density dependence (CNDD) was important to the survival of Abies nephrolepis and Picea koraiensis seedling, whereas phylogenetic negative density dependence (PNDD) was critical to Pinus koraiensis and Betula platyphylla, as well as functional groups of tree, deciduous, and shade‐intolerant seedlings. For shrubs and Acer ukurunduense, habitat heterogeneity was significant. Despite of the significance of CNDD, PNDD, and habitat heterogeneity on seedling survival, large proportions of the total variance were not accounted for by the studied variables, suggesting the needs to examine the influences of other factors such as pests, diseases, herbivores, forest structure, species functional traits, and microclimatic conditions on seedling survival in the future.     

Seedling survival simultaneously determined by conspecific,heterospecific, and phylogenetically related neighbors and habitat heterogeneity in a subtropical forest in Taiwan

Density dependence and habitat heterogeneity have been recognized as important driving mechanisms that shape the patterns of seedling survival and promote species coexistence in species‐rich forests. In this study, we evaluated the relative importance of density dependence by conspecific, heterospecific, and phylogenetically related neighbors and habitat heterogeneity on seedling survival in the Lienhuachih (LHC) Forest, a subtropical, evergreen forest in central Taiwan. Age‐specific effects of different variables were also studied. We monitored the fates of 1,642 newly recruited seedlings of woody plants within a 25‐ha Forest Dynamics Plot for 2 years. The effects of conspecific, heterospecific, and phylogenetically related neighbors and habitat heterogeneity on seedling survival were analyzed by generalized linear mixed models. Our results indicated that conspecific negative density dependence (CNDD) had a strong impact on seedling survival, and the effects of CNDD increased with seedling age. Heterospecific positive density dependence (HPDD) and phylogenetic positive density dependence (PPDD) had a significant influence on the survival of seedlings, and stronger HPDD and PPDD effects were detected for older seedlings. Furthermore, seedling survival differed among habitats significantly. Seedling survival was significantly higher in the plateau, high‐slope, and low‐slope habitats than in the valley. Overall, our results suggested that the effects of CNDD, HPDD, PPDD, and habitat heterogeneity influenced seedling survival simultaneously in the LHC subtropical forest, but their relative importance varied with seedling age. Such findings from our subtropical forest were slightly different from tropical forests, and these contrasting patterns may be attributed to differences in abiotic environments. These findings highlight the importance to incorporate phylogenetic relatedness, seedling age, and habitat heterogeneity when investigating the impacts of density dependence on seedling survival that may contribute to species coexistence in seedling communities.     

Drivers of seedling survival in a temperate forest and their relative importance at three stages of succession

Negative density dependence (NDD) and niche partitioning have been perceived as important mechanisms for the maintenance of species diversity. However, little is known about their relative contributions to seedling survival. We examined the effects of biotic and abiotic neighborhoods and the variations of biotic neighborhoods among species using survival data for 7503 seedlings belonging to 22 woody species over a period of 2 years in three different forest types, a half‐mature forest (HF), a mature forest (MF), and an old‐growth forest (OGF), each of these representing a specific successional stage in a temperate forest ecosystem in northeastern China. We found a convincing evidence for the existence of NDD in temperate forest ecosystems. The biotic and abiotic variables affecting seedlings survival change with successional stage, seedling size, and age. The strength of NDD for the smaller (<20 cm in height) and younger seedlings (1–2 years) as well as all seedlings combined varies significantly among species. We found no evidence that a community compensatory trend (CCT) existed in our study area. The results of this study demonstrate that the relative importance of NDD and habitat niche partitioning in driving seedling survival varies with seedling size and age and that the biotic and abiotic factors affecting seedlings survival change with successional stage.     

Phylogenetic density dependence and environmental filtering predict seedling mortality in a tropical forest

Negative density dependence (NDD) and environmental filtering (EF) shape community assembly, but their relative importance is poorly understood. Recent studies have shown that seedling's mortality risk is positively related to the phylogenetic relatedness of neighbours. However, natural enemies, whose depredations often cause NDD, respond to functional traits of hosts rather than phylogenetic relatedness per se. To understand the roles of NDD and EF in community assembly, we assessed the effects on seedling mortality of functional similarity, phylogenetic relatedness and stem density of neighbouring seedlings and adults in a species-rich tropical forest. Mortality risks increased for common species when their functional traits departed substantially from the neighbourhood mean, and for all species when surrounded by close relatives. This indicates that NDD affects community assembly more broadly than does EF, and leads to the tentative conclusion that natural enemies respond to phylogenetically correlated traits. Our results affirm the prominence of NDD in structuring species-rich communities.     

宝天曼国家级自然保护区落叶阔叶林密度制约效应对树木存活的影响

密度制约对物种共存起着重要作用。随着密度制约效应研究的深入, 亲缘关系较近的物种间表现出的密度制约效应逐渐被人们认识。本研究基于2009和2014年对宝天曼1 ha落叶阔叶林样地的2次调查数据, 利用广义线性混合模型分析了重要值排名前11位的物种不同邻域尺度的密度制约效应对不同径级(小径级(1 cm ≤ DBH < 5 cm)、中径级(5 cm ≤ DBH < 10 cm)、大径级(DBH ≥ 10 cm))目标个体存活的影响。研究表明: (1) 5年间样地中DBH ≥ 1 cm的所有个体的年均死亡率和增员率分别为5.85%和0.27%; (2)有5个物种的个体存活率与同种邻体个体数及同种邻体胸高断面积显著负相关; (3)小径级个体的存活在5 m的邻域范围内受同种密度制约和谱系密度制约的影响都很显著; 中径级个体在3个尺度上受到的密度制约和谱系密度制约的影响都不大; 大径级个体在7.5 m、10 m邻域范围内受谱系密度制约影响显著。结果表明, 同种密度制约和谱系密度制约效应对宝天曼落叶阔叶林不同生长阶段的树木个体影响不同。     

Species abundance is jointly determined by functional traits and negative density dependence in a subtropical forest in southern China

植物物种多度受功能性状和负密度依赖共同影响——以中国南亚热带黑石顶森林样地为例 影响植物群落中植物物种多度的因素较多,确定关键影响因素及阐明其具体机制一直是群落生态学的研究重点之一。目前确定性的影响因素主要有两大类：植物功能性状和负密度依赖。功能性状通过影响植物的竞争能力、资源的获取效率、对环境的适应能力等方面进而影响植物的多度;负密度依赖表现在同种或异种植物个体在空间上聚集,特异性的寄主(病原菌、昆虫等)传播容易导致植物个体(主要为幼苗和小树)死亡,种群发生自疏,进而影响植物的多度。到目前为止,单一探究植物功能性状或负密度依赖对植物多度影响机制的研究已有大量报道,而将两者结合来探讨对植物多度的影响的工作相对较少。为了更深入地探索影响植物多度的机制,本研究中我们综合分析了功能性状与负密度依赖对植物多度的影响。研究地点为广东省黑石顶亚热带森林50 ha大样地;功能性状方面选取了叶面积(LA)、比叶面 积(SLA)、叶片干物质含量(LDMC)、叶片氮含量(LNC)和最大电子传输速率(ETR_max);植物物种所受负密度依赖强度通过构建小树在样地两次普查期间的存活状况受邻体密度影响的层次贝叶斯模型计算;通过结构方程模型构建功能性状-负密度依赖-物种多度的内在联系框架。结构方程模型结果显示,物种多度受功能性状和负密度依赖共同作用,其中功能性状对物种多度的影响包括直接作用和通过负密度依赖的间接作用。具体来说,SLA对多度的影响包括直接和间接两种;LDMC和LNC仅间接影响物种多度;LA和ETR_max对多度只有直接影响;负密度依赖与物种多度之间存在直接的负相关关系,说明多度较高的物种受到的负密度依赖效应更强。结构方程模型对物种多度差异的解释度达到20%。以上结果表明,黑石顶植物多度分布是功能性状与负密度依赖共同作用的结果;比叶面积在所研究的因素中贡献最大。该项工作有助于提高我们对亚热带森林植物常见种和稀有种分布格局的理解。     

Effects of biotic neighbors and habitat heterogeneity on tree seedling survival in a secondary mixed conifer and broad-leaved forest in Changbai Mountain

Aims Our study aimed to understand the effects and the relative importance of biotic neighborhood and habitat heterogeneity for tree seedling survival in a secondary mixed conifer and broad-leaved forest in Changbai Mountain, north-eastern China.
Methods The generalized mixed linear model was used to examine the relative effects of biotic neighborhood and habitat heterogeneity on seedling survival over two years.
Important findings Our results showed that both biotic neighborhood and habitat heterogeneity had significant effects on the seedling survival at community level. The local environment suitable for the adult growth was also suitable for seedling survival. The soil moisture and soil available nitrogen exhibited significant positive effects on seedling survival. On the other hand, seedling density had significant negative effects on seedling survival due to the individual competition. Particularly, we found significant negative density-dependent effects on seedling survival which was caused by conspecific adult and seedling neighbors. As expected, with the increasing of seedling survival age, the habitat heterogeneity became more important on seedling survival. These results suggest that both local biotic neighborhood and habitat heterogeneity drive seedling survival in this temperate forest, and their relative importance varies with different seedling age classes and species traits.     

长白山次生针阔混交林乔木幼苗存活的影响因素分析

基于长白山次生针阔混交林样地, 以520个1 m × 1 m幼苗样方中胸径小于1 cm的乔木幼苗为研究对象, 选取2013年和2014年的幼苗调查数据, 运用广义线性混合模型(GLMM)分析了生物邻体和生境异质性对幼苗存活的影响, 探讨了次生针阔混交林幼苗存活影响因素及物种共存机制。结果表明: (1)适宜大树生长的局域生境同样也适宜幼苗的存活, 幼苗存活率与土壤含水量和有效氮等土壤养分显著正相关。(2)幼苗个体之间存在明显的竞争, 较多的幼苗邻体显著降低幼苗的存活率。同种大树邻体和同种幼苗邻体与幼苗存活显著负相关, 表明存在负密度制约效应。(3)随着幼苗年龄的增加, 生境异质性对幼苗存活的影响逐渐增大。该研究证实了密度制约效应和生境异质性对幼苗存活有着重要影响, 其相对重要性随着幼苗年龄级、功能群以及物种种类而变化。     

亚热带中山常绿阔叶林木本植物幼苗数量动态及其与生境的相关性

以百山祖自然保护区5 hm2永久样地150个幼苗监测站木本植物幼苗为研究对象,2008—2011年定期调查样方中幼苗的种类、数量、萌发、死亡等,探究亚热带中山常绿阔叶林幼苗种类组成、数量动态及其与生境的相关性。结果表明:1)百山祖样地在2008年至2011年出现的幼苗属于26科,40属,共53个物种,不同物种萌发时段有异;2)2009年样地幼苗存活比率为7.7%,2010年为-20.8%,2011年则是-0.3%,幼苗存活比率不高,种类和数量呈减少趋势;3)存活幼苗中有明显的优势物种,分别为光亮山矾(Symplocos lucida)、尖连蕊茶(Camellia cuspidata)、浙闽新木姜子(Neolitsea aurata var.undulatula)、尖叶菝葜(Smilax arisanensis)和短尾柯(Lithocarpus brevicaudatus),5个物种之和占幼苗总数比例50%;4)种子的萌发与生境有极显著的相关性,且与生境因子中水分关联最大;5)存活幼苗数与样站坡位、水分、落叶层厚度呈现显著相关性,水和光照是影响幼苗存活的主要因素。     

Resprouting enhances seedling persistence in a subtropical coastal dune forest

In theory, resprouting enables species with low reproductive output (i.e., few seedlings) to persist. The advantage conferred by seedling sprouts on tree species persistence was evaluated in a subtropical coastal dune forest in South Africa. Species with a higher frequency of seedling sprouts demonstrated greater persistence as evidenced by a larger proportion of seedlings >1 year old and a larger seedling bank than species with few seedling sprouts. Resprouted seedlings had a larger basal diameter than true seedlings. Although resprouting resulted in the maintenance of multiple stems in some seedlings, the proportion of multi‐stemmed seedlings was low. Multi‐stemming was not a favoured form of seedling growth except in one species that occupied relatively open sites. Despite the apparent difference among species in resprouting ability, we found that seedling resprouting was not phylogenetically constrained. These results demonstrate that seedling sprouts form an important component of seedling banks in coastal dune forest.     

八大公山常绿落叶阔叶混交林影响幼苗存活的主要因子分析

幼苗是植物生活史中最脆弱的阶段,对幼苗存活影响因子的分析有助于我们更清楚的了解森林群落的天然更新机制。利用广义线性混合模型（GLMM）对八大公山常绿落叶阔叶混交林中影响幼苗存活的主要生物与非生物因子进行了研究。结果表明：（1）在群落水平上,幼苗存活与生物因子中的同种幼苗密度呈显著负相关,与非生物因子中的冠层开阔度呈显著正相关;（2）从年龄上看,4年生以下龄级的幼苗存活更容易受到同种幼苗密度的影响,与同种幼苗密度呈显著负相关;4年生及其以上的幼苗存活则主要受非生物因子影响;（3）从生活型上看,相对于常绿物种,落叶物种的幼苗存活率更容易受到同种幼苗密度的影响,也与冠层开阔度呈正相关;（4）在物种水平上,生物因子与非生物因子对不同物种幼苗存活率的影响也不相同。其中,宜昌润楠（Machilus ichangensis Rehd.et Wils.）的存活率与冠层开阔度呈正相关;薄叶山矾（Symplocos anomala Brand）幼苗的存活率与同种幼苗密度、异种大树胸高断面积、林冠开阔度、坡向均呈显著负相关,而与异种幼苗密度和海拔呈显著正相关。本研究表明影响幼苗存活的因子是多样的,而且不是随机发生的。在不同水平上影响幼苗存活的因子不同。     

Conspecific density dependence in seedlings varies with species shade tolerance in a wet tropical forest

Density-dependent seedling mortality could increase with a species relative abundance, thereby promoting species coexistence. Differences among species in light-dependent mortality also could enhance coexistence via resource partitioning. These compatible ideas rarely have been considered simultaneously. We developed models of mortality as functions of irradiance and local conspecific density (LCD) for seedlings of 53 tropical woody species. Species varied in mortality responses to these factors, but mortality consistently increased with shading and LCD. Across species, density-dependent mortality on a per-neighbour basis was inversely related to species community abundance, but higher LCD in more common species resulted in a weak relationship between species abundance and density-dependent mortality scaled to species maximum LCD. Species mortality responses to shading and maximum LCD were strongly and positively correlated. Our results suggest that species differences in density-dependent mortality are more strongly related to physiologically based life-history traits than biotic feedbacks related to community abundance.     

Evolutionary history and distance dependence control survival of dipterocarp seedlings

One important hypothesis to explain tree-species coexistence in tropical forests suggests that increased attack by natural enemies near conspecific trees gives locally rare species a competitive advantage. Host ranges of natural enemies generally encompass several closely related plant taxa suggesting that seedlings should also do poorly around adults of closely related species. We investigated the effects of adult Parashorea malaanonan on seedling survival in a Bornean rain forest. Survival of P. malaanonan seedlings was highest at intermediate distances from parent trees while heterospecific seedlings were unaffected by distance. Leaf herbivores did not drive this relationship. Survival of seedlings was lowest for P. malaanonan , and increased with phylogenetic dissimilarity from this species, suggesting that survival of close relatives of common species is reduced. This study suggests that distance dependence contributes to species coexistence and highlights the need for further investigation into the role of shared plant enemies in community dynamics.
Ecology Letters (2010) 13: 51–59     

古田山亚热带常绿阔叶林树皮厚度的变异特征

树皮是木本植物茎干最外层结构,具有保护茎干、养分储存与运输等重要作用。因此,树皮厚度是一项非常重要的功能性状,其变异不仅影响树皮的各种生态功能,还能影响群落构建与物种共存。然而,以往对树皮厚度的研究集中于火灾易发生态系统,对火灾不易发生的亚热带常绿阔叶林的研究仍较缺乏。测量了古田山国家级自然保护区亚热带常绿阔叶林内树种的树皮厚度,并检验了总树皮厚度、内树皮厚度与外树皮厚度在各分类群间以及功能群间的差异。结果发现：1）39个树种807个个体的总树皮厚度、内树皮厚度与外树皮厚度均值分别为1.90 mm、1.38 mm和0.54 mm。漆树科（Anacardiaceae）、杨梅科（Myricaceae）以及亚热带常绿阔叶林代表类群壳斗科（Fagaceae）、山茶科（Theaceae）的树皮厚度较大。短柄枹（Quercus serrata）、木荷（Schima superba）、小叶青冈（Cyclobalanopsis myrsinifolia）等树种的树皮厚度较大。2）种间、科间的各树皮厚度差异均显著。不同功能类群间,乔木类群的各树皮厚度均较灌木类群大,常绿类群的各树皮厚度均较落叶类群大（内树皮厚度除外）。本次研究结果表明,相对于火灾易发生态系统中的树皮厚度,古田山亚热带常绿阔叶林群落内的树皮厚度相对较薄,表明这些森林树种对当地湿润气候的适应性。同时,树皮厚度在各种分类水平与功能群水平间的显著变异,反映了群落内不同生态策略的共存。     

森林种子雨研究进展与展望

种子雨阶段是植物更新的关键环节,它连接着繁殖生产与植物后续生活史阶段,对群落结构有着重要的影响.虽然早在19世纪中叶达尔文就认识到种子扩散的重要性,然而对种子雨的广泛研究于20世纪80年代初才开始.本文聚焦于森林木本植物种子雨研究,首先介绍了种子雨监测方法,包括收集器的布置,种子雨的收集、分离和鉴定.然后综述了种子雨的4个主要研究方向:种子产量的时间和空间变化格局(包括季节变化、年际变化和空间变化)、增补限制及其在物种多样性维持中的作用、验证负密度效应假说、种子雨与其他生活史阶段(土壤种子库、幼苗、幼树及母树)的比较.未来还需要加强对种子雨的长期监测,开展增补限制的跨纬度比较研究,探讨植物早期更新阶段负密度效应沿纬度梯度的变化规律,加强数学模型以及分子标记和稳定同位素技术等新手段的运用.     

Tri-trophic interactions affect density dependence of seed fate in a tropical forest palm

Natural enemies, especially host-specific enemies, are hypothesised to facilitate the coexistence of plant species by disproportionately inflicting more damage at increasing host abundance. However, few studies have assessed such Janzen-Connell mechanisms on a scale relevant for coexistence and no study has evaluated potential top-down influences on the specialized pests. We quantified seed predation by specialist invertebrates and generalist vertebrates, as well as larval predation on these invertebrates, for the Neotropical palm Attalea butyracea across ten 4-ha plots spanning 20-fold variation in palm density. As palm density increased, seed attack by bruchid beetles increased, whereas seed predation by rodents held constant. But because rodent predation on bruchid larvae increased disproportionately with increasing palm density, bruchid emergence rates and total seed predation by rodents and bruchids combined were both density-independent. Our results demonstrate that top-down effects can limit the potential of host-specific insects to induce negative-density dependence in plant populations.     

生态位分化与森林群落物种多样性维持研究展望

一直以来,生态学家和进化生物学家对森林群落物种多样格局及其形成机制持有不同的观点。虽然Robert Ricklefs将进化和生态过程整合的观点已经被群落生态学家广泛接受,但是区域物种进化历史以及局域群落微进化过程是否能够影响群落生态学过程以及这些过程如何影响群落结构和动态还有待商榷。经典的生态位理论同时强调了种间和种内生态位分化对群落多样性维持的影响。但是生态学家普遍认为种间差异足以代表群落内个体间的相互作用关系,并且由于进化过程导致的种内分化往往涉及较长的时间尺度,因此,虽然种内差异是自然选择的重要材料,物种对环境的适应性进化过程所导致的种内分化对群落构建的影响往往被生态学家所忽视。为此,通过回顾种间和个体生态位分化的研究历史,对两类研究分别进行简要阐述,强调在今后的群落生态学研究中需要考虑个体分化对局域群落构建的影响。