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。     

密度制约对宝天曼落叶阔叶林锐齿栎死亡前后分布格局的影响

为了在温带和亚热带的过渡带中验证森林树木死亡是否受密度制约的影响, 我们在宝天曼国家级自然保护区选择1个100 m×100 m的暖温带落叶阔叶林样地, 用双变量函数g(r)(the pair-correlation function)研究了锐齿栎(Quercus aliena var. acuteserrata)死亡前(活树和枯木统称为死亡前树木)和死亡后(活树为死亡后树木)的分布格局。把样地中的锐齿栎分为幼树(1 cm ≤ DBH<10 cm)、小树(10 cm ≤ DBH<20 cm)和成年树(DBH≥20 cm)3个不同的生长阶段来研究密度制约对空间分布格局的影响机制。结果表明: (1)死亡前锐齿栎在r>5 m尺度呈聚集分布, 死亡后幼树(1 cm≤DBH<10 cm)和成年树(DBH≥20 cm)在1-25 m尺度呈现随机分布, 死亡后小树(10 cm≤DBH< 20 cm)在r<1.5 m和2.5-4.5 m的尺度为随机分布, 在r>5 m的尺度呈聚集分布; (2)采用随机标签零模型和案例-对照设计的方法, 排除生境异质性影响后, 将幼树和小树的分布格局作为案例, 将成年树的分布格局作为对照, 并代表生境异质性的作用, 通过小径级树木与成年树分布格局的对比发现, 密度制约效应对死亡前后的锐齿栎分布格局均具有影响; (3)幼树和小树在成年树周围的分布死亡前为显著聚集分布格局, 死亡后剩余树木的聚集强度下降; 随着与成年树之间距离的增加, 死亡后的幼树(仅包括现存的活树)逐渐向随机分布格局演替。本研究初步表明锐齿栎空间格局受生境异质性的影响并呈现出显著的聚集效应, 排除生境异质性影响后, 锐齿栎死亡前后的空间格局受到密度制约的影响, 这一结果为Janzen-Connell假说提供了支持。     

密度制约效应对啮齿动物繁殖的影响

啮齿动物种群动态调节是生态学中关键的问题之一,它与制定鼠害防治策略密切相关。目前,国内外学者已在相关领域取得大量的研究成果。整理了密度制约效应对啮齿动物繁殖影响的文献,包括在性器官、性成熟、性周期、繁殖指数、激素和行为的调节作用,同时分析该领域的研究途径及研究前景,旨在为今后的研究提供更多的思路。     

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。     

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。     

东方田鼠种群密度制约的迟滞效应

本研究中用成年东方田鼠不同时间不同密度笼养后（笼内雌雄各半）,低密度配对饲养,观察各阶段东方田鼠繁殖指标的差异。试鼠144只,处理时分为LL（低密度长时间）（2只/笼,共20笼,90d）、HL（高密度长时间）（8只/笼,共5笼,90d）、HM（高密度中等时间）（8只/笼,共4笼,20d）与HS（高密度短时间）（8只/笼,共4笼,10d）4组。处理后在低密度条件下观察繁殖情况（胎仔数、怀孕率、分娩频率以及产仔间隔等）,直至180 d。实验数据按时间划分为3个部分统计：0~90 d为第一阶段(不同密度处理期),90~109 d为过渡阶段（处理后的过渡期）,109 d以后为第二阶段（正常低密度配对繁殖期）。结果表明：经过长时间高密度处理后,平均产仔数显著减少,组间平均分娩频率无显著差异。怀孕率由HL组到HM组,再到LL和HS组显著增加。在第一阶段,除了LL组之外,其余各组都未发现繁殖。过渡阶段的怀孕率组间差异显著,LL组以及HM组显著高于HL组。第二阶段的怀孕率以及胎仔数的组间差异显著,都为LL、HS两组较高,HL、HM两组较低。平均分娩频率、平均胎仔数以及产仔间隔各组间无显著差异。结论：不同持续时间的密度效应有较大差别。对于东方田鼠,20d的高密度处理相比于10d更能对其繁殖起到抑制作用。恢复低密度后,存在种群的迟滞性密度制约。     

常绿阔叶林谱系多样性对幼苗存活率的影响

密度制约机制对于维护生物多样性有非常重要的作用。随着对密度制约机制的深入研究, 人们逐渐认识到: 不仅在种内存在密度制约效应, 亲缘关系相近的物种之间也可能表现出密度制约效应。Webb在2000年提出的NRI (净种间亲缘关系指数)、NTI (净最近种间亲缘关系指数)考虑了比较全面的谱系信息, 获得了广泛的应用。该文采用NRI、NTI来代表种间关系, 并用Logistic回归模型来分析了谱系因子对浙江省开化县古田山自然保护区24 hm²永久监测样地中501个幼苗样方死亡率的影响。通过对6次幼苗调查数据的分析表明: 不仅相同物种密度对于幼苗的死亡率有显著影响, 当密度达到一定水平时, 谱系因子同样也对幼苗的死亡率有显著影响——苗区中个体间的亲缘关系越近, 幼苗个体的死亡率越高。     

种群密度效应对成年东方田鼠内脏器官的影响

为了研究种群密度对东方田鼠(Microtus fortis)内脏器官(心、肺、肝、脾、肾、肾上腺)的影响,随机选取稳定的同性别饲养群的成年东方田鼠258只,雌雄各半,分A组30笼:2只/笼,B组15笼:4只/笼,C组11笼:6只/笼和D组9笼:8只/笼共4个密度组,饲养90d后解剖,取内脏器官,称重以及测定血浆皮质醇浓度并进行比较。结果表明:在相同密度内,除B组孕鼠的肝脏重较未孕鼠有极显著差异外,其它各组(A、C、D)怀孕鼠和未孕鼠的各内脏器官重均无显著差异。在不同密度间,怀孕鼠的肝脏重有极显著差异,孕鼠与未孕鼠的其余各内脏器官均无显著差异;雄鼠的肺重、脾脏重和雌鼠的脾脏重、肾上腺重差异显著,其余各器官重在不同密度间虽未表现出显著差异,但均具有随着密度增加而器官重量逐渐增大的趋势。各个密度组中的雄性个体血浆皮质醇浓度分别显著高于雌性,且差异显著;同一性别比较,不同密度间的雌性个体血浆皮质醇浓度差异不显著,而雄性个体血浆皮质醇浓度差异则显著,随密度增大而明显增加。可见高密度压力导致内脏器官重量上有所调节,但不同脏器受密度因素影响程度不同。     

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。     

天山雪岭云杉森林群落的密度制约效应

密度制约是维持自然森林群落物种共存的重要机制之一。雪岭云杉(Picea schrenkiana)是天山森林群落的单优种, 在我国温带森林中占有重要地位。本文基于对天山雪岭云杉8 ha森林动态监测样地的两次调查结果, 分析了该样地群落物种组成的动态变化, 运用点格局方法分析了雪岭云杉个体在不同龄级的空间分布格局及其关联性, 探讨了该群落的密度制约效应。结果表明: (1)该样地内, DBH ≥ 1 cm的木本植物在2009年有11,835株, 2014年为11,050株, 5年间的个体死亡率为8.82%, 补员率为2.19%, 个体株数净减少6.63%。(2)使用双关联函数g(r)分析了不同龄级雪岭云杉的空间分布格局, 发现幼龄树和中龄树在0-40 m尺度上主要呈现聚集分布, 但随着尺度增大聚集强度逐渐减小; 老龄树在0-2 m及4 m尺度上呈随机分布, 在其他尺度上呈聚集分布, 但聚集程度较小。(3)用“案例-对照”设计的方法, 排除生境异质性的影响后, 将幼龄树和中龄树作为案例, 老龄树作为对照, 对比幼龄树、中龄树与老龄树的分布格局, 发现雪岭云杉幼龄树和中龄树在0-40 m尺度上呈现额外的聚集, 随着径级的增大, 这种额外的聚集强度逐渐减小, 即表现出密度制约效应。(4)老龄树与中龄树、幼龄树的空间关联性相同, 在0-40 m尺度上均呈明显负关联。     

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.     

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.     

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.     

Elevational variation in density dependence in a subtropical forest

Density‐dependent mortality has been recognized as an important mechanism that underpins tree species diversity, especially in tropical forests. However, few studies have attempted to explore how density dependence varies with spatial scale and even fewer have attempted to identify why there is scale‐dependent differentiation. In this study, we explore the elevational variation in density dependence. Three 1‐ha permanent plots were established at low and high elevations in the Heishiding subtropical forest, southern China. Using data from 1200 1 m² seedling quadrats, comprising of 200 1 m² quadrats located in each 1‐ha plot, we examined the variation in density dependence between elevations using a generalized linear mixed model with crossed random effects. A greenhouse experiment also investigated the potential effects of the soil biota on density‐dependent differentiation. Our results demonstrated that density‐dependent seedling mortality can vary between elevations in subtropical forests. Species found at a lower elevation suffered stronger negative density dependence than those found at a higher elevation. The greenhouse experiment indicated that two species that commonly occur at both elevations suffered more from soilborne pathogens during seed germination and seedling growth when they grew at the lower elevation, which implied that soil pathogens may play a crucial role in density‐dependent spatial variation.     

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.     

Effects of size,neighbors, and site condition on tree growth in a subtropical evergreen and deciduous broad‐leaved mixed forest,China

Successful growth of a tree is the result of combined effects of biotic and abiotic factors. It is important to understand how biotic and abiotic factors affect changes in forest structure and dynamics under environmental fluctuations. In this study, we explored the effects of initial size [diameter at breast height (DBH)], neighborhood competition, and site condition on tree growth, based on a 3‐year monitoring of tree growth rate in a permanent plot (120 × 80 m) of montane Fagus engleriana–Cyclobalanopsis multiervis mixed forest on Mt. Shennongjia, China. We measured DBH increments every 6 months from October 2011 to October 2014 by field‐made dendrometers and calculated the mean annual growth rate over the 3 years for each individual tree. We also measured and calculated twelve soil properties and five topographic variables for 384 grids of 5 × 5 m. We defined two distance‐dependent neighborhood competition indices with and without considerations of phylogenetic relatedness between trees and tested for significant differences in growth rates among functional groups. On average, trees in this mixed montane forest grew 0.07 cm year^?1 in DBH. Deciduous, canopy, and early‐successional species grew faster than evergreen, small‐statured, and late‐successional species, respectively. Growth rates increased with initial DBH, but were not significantly related to neighborhood competition and site condition for overall trees. Phylogenetic relatedness between trees did not influence the neighborhood competition. Different factors were found to influence tree growth rates of different functional groups: Initial DBH was the dominant factor for all tree groups; neighborhood competition within 5 m radius decreased growth rates of evergreen trees; and site condition tended to be more related to growth rates of fast‐growing trees (deciduous, canopy, pioneer, and early‐successional species) than the slow‐growing trees (evergreen, understory, and late‐successional species).