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。     

长白山阔叶红松林中影响乔木幼苗存活的关键因素

应用广义线性混合模型,对长白山阔叶红松林中影响幼苗存活的生物因素和非生物因素进行分析。结果表明:(1)在群落水平上,幼苗存活率和生物因素中同种幼苗邻居显著负相关,说明在阔叶红松林群落中存在负密度制约效应。(2)生物因子和非生物因子对不同年龄阶段的幼苗存活率影响不同。对于1年生的幼苗,幼苗存活率与异种大树邻居呈显著负相关,与同种大树邻居呈显著正相关;对于2—3年生的幼苗,其存活率和同种幼苗邻居、同种大树邻居均呈显著负相关,和非生物因子相关不显著;对于4年生以上的幼苗,其存活率和土壤主成分分析的PC1(低的有机质、速效钾、速效氮等比较贫瘠的土壤)显著负相关。(3)种子的传播方式不同,幼苗存活率的影响因子也不同。对于风传播的物种,存活率与同种幼苗邻居密度显著正相关。对于重力传播的物种,幼苗存活率与土壤PC3(高的全氮和速效氮,含氮较高的土壤)、异种大树邻居、草本密度呈负相关,与林冠开阔度和草本盖度成正相关。(4)对于不同的物种,影响幼苗存活的因素也不同。紫椴的幼苗存活率与土壤PC3、异种大树邻居、草本密度呈显著负相关,与土壤主成分PC2(高的有机质和全氮等养分比较好的土壤)呈显著正相关。     

长白山阔叶红松林乔木幼苗组成及多度格局的影响因素

以长白山原始阔叶红松林样地为平台,以样方中胸径小于1 cm的乔木幼苗为研究对象,基于2018年两次的幼苗调查数据,利用广义线性混合模型分析影响群落乔木幼苗多度的生物邻体和生境因素。结果表明:该样地所有幼苗样方共调查到10064株乔木幼苗,累计17个物种,分属9科9属,水曲柳幼苗多度极高,在乔木树种幼苗中占优势地位;在群落水平,幼苗多度与异种大树效应、草本密度和林冠开阔度呈显著正相关,与草本盖度和土壤含水量呈显著负相关;在物种水平,水曲柳和紫椴幼苗多度的影响因素与群落水平筛选后的结果一致,红松幼苗多度与异种大树效应呈正相关,与同种大树效应及林冠开阔度呈负相关。该研究证实了生物邻体和生境因素共同影响幼苗多度格局,并且生物邻体和生境因素的相对重要性随幼苗物种种类不同而变化。     

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。     

天童常绿阔叶林定居幼苗存活和生长的关联

群落幼苗的更新过程是维持群落物种共存的关键环节,其中幼苗成功定居后的发展阶段是群落幼苗更新过程的重要组成部分,对群落可繁殖个体补充格局的形成起着十分重要的作用。然而,相比于新生幼苗,群落内定居幼苗短期死亡率相对较低,需要较长时间的监测才能完整地了解其存活格局,所以相关的研究还相对较少。本研究假设群落内已经成功定居的幼苗的生长状况能够较好地预测其存活情况,通过分析影响其生长的外在因素,尝试间接地估计定居幼苗可能的存活格局,从而了解定居幼苗的存活和生长过程。为验证该假设,本文选取浙江天童20 ha森林动态监测样地内的定居幼苗作为研究对象,利用广义线性混合模型分析其相对生长速率,以及生物邻体因子(同种/异种成树邻体指数,同种/异种幼苗邻体密度,同种/异种成树邻体的凋落叶产量)和微生境因子(冠层开度、草本覆盖率、海拔、坡度、坡向、土壤p H、全氮和全磷含量)对定居幼苗短期存活概率的影响,并利用线性混合模型分析生物邻体因子和微生境因子对定居幼苗相对生长速率的影响。结果显示,群落内定居幼苗的相对生长速率对其本身的短期存活概率有显著正影响,且其影响程度最高;冠层开度对群落幼苗相对生长速率有显著正影响,且是唯一有显著影响的因子。上述结果说明,在群落幼苗成功定居后,其生长状况能够较好地预测其存活情况,加之林下较好的光照条件能够促进幼苗的生长,这可能间接导致在林下光照较好的生境条件下定居幼苗长期存活的可能性相对较高。     

吉林蛟河针阔混交林乔木幼苗组成及其密度格局影响因素

以吉林蛟河次生针阔混交林42 hm ²固定监测样地中209个幼苗监测站内的乔木幼苗为研究对象, 基于2016和2017年幼苗调查数据, 探究幼苗物种组成、数量动态等特征, 并运用广义线性混合效应模型分析了幼苗密度与生物邻体及其生境因素的相关性。结果表明: (1)该样地内所有幼苗监测站共调查到幼苗4 245株, 分属10科12属18种, 新生幼苗的数量在物种和调查年份间均有明显差异, 其中水曲柳(Fraxinus mandschurica)和紫椴(Tilia amurensis)幼苗表现出大量出生和死亡的动态特征。(2)群落水平上, 幼苗密度与局域同种成体胸高断面积之和、土壤全磷和有效钾等养分含量显著正相关, 适宜大树生长的生境同样适宜幼苗的生长。(3)水曲柳幼苗密度的影响因素与群落水平一致, 红松(Pinus koraiensis)存在明显的生境偏好, 在湿度较小、土壤全磷、有效氮和有效磷含量较低的生境中密度更高。相对于多年生幼苗, 同种成年邻体对当年生幼苗密度影响更加显著。该研究证实了扩散限制和生境过滤共同影响幼苗密度格局, 生物邻体和生境异质性的相对重要性随幼苗物种种类和年龄级变化。     

Drivers of composition and density pattern of tree seedlings in a secondary mixed conifer and broad-leaved forest,Jiaohe, Jilin,China

以吉林蛟河次生针阔混交林42 hm ²固定监测样地中209个幼苗监测站内的乔木幼苗为研究对象, 基于2016和2017年幼苗调查数据, 探究幼苗物种组成、数量动态等特征, 并运用广义线性混合效应模型分析了幼苗密度与生物邻体及其生境因素的相关性。结果表明: (1)该样地内所有幼苗监测站共调查到幼苗4 245株, 分属10科12属18种, 新生幼苗的数量在物种和调查年份间均有明显差异, 其中水曲柳(Fraxinus mandschurica)和紫椴(Tilia amurensis)幼苗表现出大量出生和死亡的动态特征。(2)群落水平上, 幼苗密度与局域同种成体胸高断面积之和、土壤全磷和有效钾等养分含量显著正相关, 适宜大树生长的生境同样适宜幼苗的生长。(3)水曲柳幼苗密度的影响因素与群落水平一致, 红松(Pinus koraiensis)存在明显的生境偏好, 在湿度较小、土壤全磷、有效氮和有效磷含量较低的生境中密度更高。相对于多年生幼苗, 同种成年邻体对当年生幼苗密度影响更加显著。该研究证实了扩散限制和生境过滤共同影响幼苗密度格局, 生物邻体和生境异质性的相对重要性随幼苗物种种类和年龄级变化。     

鼎湖山南亚热带常绿阔叶林不同树种存活对邻体组成的响应差异

理解群落结构和动态的主导机制是生态学研究的基本目标之一。群落内树种的存活受到其邻近树木的显著影响。为探究不同树种的存活对邻体组成的响应差异, 本研究基于鼎湖山南亚热带阔叶林20 ha森林动态监测样地中常见的90个树种的存活监测数据和功能性状数据, 建立了一系列关于邻体效应的树种存活模型。结果表明: 约58%的树种存活对邻体组成有敏感的响应, 共存树种间的功能性状差异影响着50%的树种存活动态。不同树种对邻体组成的响应差异与其耐阴性相关, 耐阴能力较弱的树种更倾向于表现出对邻体的敏感性。低比叶面积、高叶干物质含量、木材密度和最大胸径意味着较强的耐阴能力, 与光资源利用策略有关的生态位分化可能是邻域尺度上物种共存的原因。本研究为量化邻体间的相互作用和解释局域群落的物种共存提供了新的视角。     

植物群落物种共存机制:负密度制约假说

负密度制约假说主要描述由于资源竞争、有害生物侵害(比如病原微生物、食草动物捕食)等, 同种个体之间发生的相互损害行为; 它主要强调同种个体之间的相互作用, 解释自然群落物种共存的机理; 负密度制约机制主要在小尺度上降低群落内同种个体生长率, 同时提高个体死亡率, 从而为其他物种的生存提供空间和资源, 促进物种共存。目前负密度制约假说的检验研究主要侧重密度制约、距离制约、群落补偿效应等三个方面。最近, 研究者又探讨了近缘物种之间由于对相似资源的竞争所产生的负效应, 扩展了负密度制约假说, 进而提出异群保护假说和谱系多样性制约假说。负密度制约假说引起生态学家长久的探讨和关注, 世界范围内大尺度森林动态样地的建立, 又为探索不同尺度上密度制约效应的研究提供了条件。然而目前的研究仍然存在不足, 比如负密度制约假说的检验受到其他因素的干扰、区域研究不平衡等。因此, 生态学家们仍然怀疑负密度制约效应调节群落物种共存的重要性, 但是目前的研究还没有发现否定负密度制约假说的充分证据。     

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。     

Density dependence and habitat preference shape seedling survival in a subtropical forest in central China

Aims Seedlings are vulnerable to many kinds of fatal abiotic and biotic agents, and examining the causes of seedling dynamics can help understand mechanisms of species coexistence. To disentangle the relative importance of neighborhood densities, habitat factors and phylogenetic relatedness on focal seedling survival, we monitored the survival of 5306 seedlings of 104 species>15 months. We address the following questions: (i) How do neighborhood densities, habitat variables and phylogenetic relatedness affect seedling survival? What is the relative importance of conspecific densities, habitat variables and phylogenetic relatedness to seedling survival? (ii) Does the importance of the neighborhood densities, habitat variables and phylogenetic relatedness vary among growth forms, leaf habits or dispersal modes? Specially, does the conspecific negative density dependence inhibit tree and deciduous seedlings more compared with shrub and evergreen species? Does density dependence affect the wind and animal-dispersed species equally?Methods We established 135 census stations to monitor seedling dynamics in a 25-ha subtropical forest plot in central China. Conspecific and heterospecific seedling density in the 1-m 2 seedling plot and adult basal area within a 20-m radius provided neighborhood density variables. Mean elevation, convexity and aspect of every 5- × 5-m grid with seedling plots were used to quantify habitat characteristics. We calculated the relative average phylodiversity between focal seedling and heterospecific neighbors to quantify the species relatedness in the neighborhood. Eight candidate generalized linear mixed models with binominal error distribution were used to compare the relative importance of these variables to seedling survival. Akaike's information criteria were used to identify the most parsimonious models.Important findings At the community level, both the neighborhood densities and phylogenetic relatedness were important to seedling survival. We found negative effects of increasing conspecific seedlings, which suggested the existence of species-specific density-dependent mortality. Phylodiversity of heterospecific neighbors was negatively related to survival of focal seedlings, indicating similar habitat preference shared among phylogenetically closely related species may drive seedling survival. The relative importance of neighborhood densities, habitat variables and phylogenetic relatedness varied among ecological guilds. Conspecific densities had significant negative effect for deciduous and wind-dispersed species, and marginally significant for tree seedlings>10cm tall and animal-dispersed species. Habitat variables had limited effects on seedling survival, and only elevation was related to the survival of evergreen species in the best-fit model. We conclude that both negative density-dependent mortality and habitat preference reflected by the phylogenetic relatedness shape the species coexistence at seedling stage in this forest.     

The contribution of understory light availability and biotic neighborhood to seedling survival in secondary versus old-growth temperate forest

Seedling survival plays an important role in the maintenance of species diversity and forest dynamics. Although substantial gains have been made in understanding the factors driving patterns of seedling survival in forests, few studies have considered the simultaneous contribution of understory light availability and the local biotic neighborhood to seedling survival in temperate forests at different successional stages. Here, we used generalized linear mixed models to assess the relative importance of understory light availability and biotic neighborhood variables on seedling survival in secondary and old-growth temperate forest in north eastern China at two levels (community and guild). At the community level, biotic neighborhood effects on seedling survival were more important than understory light availability in both forests. In both the old-growth and secondary forests, conspecific basal area had a negative effect on seedling survival, consistent with negative conspecific density dependence. At guild levels, the relative importance of light and biotic neighborhood on seedling survival showed considerable variation among guilds in both forests. Available understory light tended to have positive effects on seedling survival for shrub and light-demanding species in the old-growth forest, but negative effects on survival of shade-tolerant seedlings in the secondary forest. For tree species and shade-tolerant species, the best fit models included neighborhood variables, but that was not the case for shrubs, light-demanding, or mid shade-tolerant species. Overall, our results demonstrate that the relative importance of understory light availability and biotic factors on seedling survival vary with species life-history strategy and forest successional stage.     

Effects of local biotic neighbors and habitat heterogeneity on tree and shrub seedling survival in an old-growth temperate forest

Seedling dynamics play a crucial role in determining species distributions and coexistence. Exploring causes of variation in seedling dynamics can therefore provide key insights into the factors affecting these phenomena. We examined the relative importance of biotic neighborhood processes and habitat heterogeneity using survival data for 5,827 seedlings in 39 tree and shrub species over 2?years from an old-growth temperate forest in northeastern China. We found significant negative density-dependence effects on survival of tree seedlings, and limited effects of habitat heterogeneity (edaphic and topographic variables) on survival of shrub seedlings. The importance of negative density dependence on young tree seedling survival was replaced by habitat in tree seedlings ??4?years old. As expected, negative density dependence was more apparent in gravity-dispersed species compared to wind-dispersed and animal-dispersed species. Moreover, we found that a community compensatory trend existed for trees. Therefore, although negative density dependence was not as pervasive as in other forest communities, it is an important mechanism for the maintenance of community diversity in this temperate forest. We conclude that both negative density dependence and habitat heterogeneity drive seedling survival, but their relative importance varies with seedling age classes and species traits.     

Conspecific density dependence and community structure: Insights from 11 years of monitoring in an old‐growth temperate forest in Northeast China

Forest community structure may be influenced by seedling density dependence, however, the effect is loosely coupled with population dynamics and diversity in the short term. In the long term the strength of conspecific density dependence may fluctuate over time because of seedling abundance, yet few long‐term studies exist. Based on 11 years of seedling census data and tree census data from a 25‐ha temperate forest plot in Northeast China, we used generalized linear mixed models to test the relative effects of local neighborhood density and abiotic factors on seedling density and seedling survival. Spatial point pattern analysis was used to determine if spatial patterns of saplings and juveniles, in relation to conspecific adults, were in accordance with patterns uncovered by conspecific negative density dependence at the seedling stage. Our long‐term results showed that seedling density was mainly positively affected by conspecific density, suggesting dispersal limitation of seedling development. The probability of seedling survival significantly decreased over 1 year with increasing conspecific density, indicating conspecific negative density dependence in seedling establishment. Although there was variation in conspecific negative density dependence at the seedling stage among species and across years, a dispersed pattern of conspecific saplings relative to conspecific adults at the local scale (<10 m) was observed in four of the 11 species examined. Overall, sapling spatial patterns were consistent with the impacts of conspecific density on seedling dynamics, which suggests that conspecific negative density dependence is persistent over the long term. From the long‐term perspective, conspecific density dependence is an important driver of species coexistence in temperate forests.     

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.     

Density‐dependent Survival in Seedlings Differs among Woody Life‐forms in Tropical Wet Forests of a Caribbean Island

Negative density dependence contributes to seedling dynamics in forested ecosystems, but the relative importance of this factor for different woody plant life‐forms is not well‐understood. We used 1 yr of seedling survivorship data for woody seedlings in 17 different plots of lower to mid‐montane rain forests on the island of Dominica to examine how seedling height, abiotic factors, and biotic factors such as negative density dependence are related to seedling survival of five different life‐forms (canopy, midstory, and understory trees; shrubs; and lianas). Across 64 species, taller seedlings in seedling plots with higher canopy openness, greater seedling density, lower relative abundance of conspecific seedlings, and lower relative abundance of conspecific adults generally had a greater probability of surviving. Height was the strongest predictor of seedling survival for all life‐forms except lianas. Greater seedling density was positively related to survival for canopy and midstory trees but negatively related to survival for the other life‐forms. For trees, the relative abundance of conspecific seedling and adult neighbors had weak and strong negative effects on survival respectively. Neither shrub nor liana seedling survival was affected by the relative abundance of conspecific neighbors. Thus, negative density dependence is confirmed as an important structuring mechanism for tree seedling communities but does not seem to be important for lianas and shrubs in Dominican rain forests. These results represent the first direct assessment of controls on seedling survival of all woody life‐forms – an important step in understanding the dynamics and structure of the entire woody plant community.     

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.     

Local-scale drivers of tree survival in a temperate forest

Tree survival plays a central role in forest ecosystems. Although many factors such as tree size, abiotic and biotic neighborhoods have been proposed as being important in explaining patterns of tree survival, their contributions are still subject to debate. We used generalized linear mixed models to examine the relative importance of tree size, local abiotic conditions and the density and identity of neighbors on tree survival in an old-growth temperate forest in northeastern China at three levels (community, guild and species). Tree size and both abiotic and biotic neighborhood variables influenced tree survival under current forest conditions, but their relative importance varied dramatically within and among the community, guild and species levels. Of the variables tested, tree size was typically the most important predictor of tree survival, followed by biotic and then abiotic variables. The effect of tree size on survival varied from strongly positive for small trees (1-20 cm dbh) and medium trees (20-40 cm dbh), to slightly negative for large trees (>40 cm dbh). Among the biotic factors, we found strong evidence for negative density and frequency dependence in this temperate forest, as indicated by negative effects of both total basal area of neighbors and the frequency of conspecific neighbors. Among the abiotic factors tested, soil nutrients tended to be more important in affecting tree survival than topographic variables. Abiotic factors generally influenced survival for species with relatively high abundance, for individuals in smaller size classes and for shade-tolerant species. Our study demonstrates that the relative importance of variables driving patterns of tree survival differs greatly among size classes, species guilds and abundance classes in temperate forest, which can further understanding of forest dynamics and offer important insights into forest management.     

Density dependence across multiple life stages in a temperate old-growth forest of northeast China

Recent studies on species coexistence suggest that density dependence is an important mechanism regulating plant populations. However, there have been few studies of density dependence conducted for more than one life-history stage or that control for habitat heterogeneity, which may influence spatial patterns of survival and mask density dependence. We explored the prevalence of density dependence across multiple life stages, and the effects of controlling for habitat heterogeneity, in a temperate forest in northeast China. We used generalized linear mixed-effects models to test for density-dependent mortality of seedlings and spatial point pattern analysis to detect density dependence for sapling-to-juvenile transitions. Conspecific neighbors had a negative effect on survival of plants in both life stages. At the seedling stage, we found a negative effect of conspecific seedling neighbors on survival when analyzing all species combined. However, in species-level analyses, only 2 of 11 focal species were negatively impacted by conspecific neighbors, indicating wide variation among species in the strength of density dependence. Controlling for habitat heterogeneity did not alter our findings of density dependence at the seedling stage. For the sapling-to-juvenile transition stage, 11 of 15 focal species showed patterns of local scale (<10 m) conspecific thinning, consistent with negative density dependence. The results varied depending on whether we controlled for habitat heterogeneity, indicating that a failure to account for habitat heterogeneity can obscure patterns of density dependence. We conclude that density dependence may promote tree species coexistence by acting across multiple life-history stages in this temperate forest.