Seed germination in temperate rain forest species of southern Chile: chilling and gap-dependency germination

Canopy gap and chilling requirements for seed germination were assessed in 61 and 44 species, respectively, in the temperate rain forest of southern Chile. Germination assays within canopy gap and understorey were carried out under natural conditions. Germination tests in cold stratified and non-stratified seeds were performed under laboratory conditions. Seeds were collected of common trees, shrubs, vines and perennial herbs of forests in southern Chile. Final percent germination was significantly enhanced under canopy gap conditions in 19 species, and significantly reduced in 11 species. Germination proved indifferent under gap vs. understorey conditions in half the species tested. Cold stratification affected germination in 11 species: significantly increasing final germination of four species, and significantly decreasing final percent germination of seven species. A Principal Components Analysis (PCA) was applied with the purpose of identifying groups of species with similar germination strategies. Four attributes were included in the PCA: final germination percentage in canopy gap, germination rate in the laboratory, and gap-and chilling-dependency indices (EGAP and STRAT, respectively). The first axis separated species mainly on EGAP variation while the second axis separated them mainly according to STRAT variation; the two axes together explaining 73% of the among-species variation. A small group of trees and vines germinating best in the understorey and neutral to chilling could clearly be distinguished from the remaining species analyzed. Multifactorial ANOVAs were used for evaluating the combined effect of successional status, seed mass, dispersal period, life form, phylogenetic categories, and dispersal syndrome on EGAP and STRAT variation. The EGAP value of secondary successional species proved significantly greater than that of primary successional species and species with endozoochorous seed dispersal were significantly less dependent on chilling (according to STRAT values) compared to species with mainly wind-dispersed seeds. The possible implications of these results for seedling establishment are discussed. This revised version was published online in August 2006 with corrections to the Cover Date.     

Impacts of forest fragmentation on species composition and forest structure in the temperate landscape of southern Chile

Aim Few studies have explicitly examined the influence of spatial attributes of forest fragments when examining the impacts of fragmentation on woody species. The aim of this study was to assess the diverse impacts of fragmentation on forest habitats by integrating landscape‐level and species‐level approaches. Location The investigation was undertaken in temperate rain forests located in southern Chile. This ecosystem is characterized by high endemism and by intensive recent changes in land use. Method Measures of diversity, richness, species composition, forest structure and anthropogenic disturbances were related to spatial attributes of the landscape (size, shape, connectivity, isolation and interior forest area) of forest fragments using generalized linear models. A total of 63 sampling plots distributed in 51 forest fragments with different spatial attributes were sampled. Results Patch size was the most important attribute influencing different measures of species composition, stand structure and anthropogenic disturbances. The abundance of tree and shrub species associated with interior and edge habitats was significantly related to variation in patch size. Basal area, a measure of forest structure, significantly declined with decreasing patch size, suggesting that fragmentation is affecting successional processes in the remaining forests. Small patches also displayed a greater number of stumps, animal trails and cow pats, and lower values of canopy cover as a result of selective logging and livestock grazing in relatively accessible fragments. However, tree richness and β‐diversity of tree species were not significantly related to fragmentation. Main conclusions This study demonstrates that progressive fragmentation by logging and clearance is associated with dramatic changes in the structure and composition of the temperate forests in southern Chile. If this fragmentation process continues, the ability of the remnant forests to maintain their original biodiversity and ecological processes will be significantly reduced.     

Spatial dynamics of plant species in an agricultural landscape in the Netherlands

This study examined the changes in distribution patterns of 13 herbaceous plant species from 1998 to 2000 in ditch banks along the edges of arable fields in the Netherlands. The objective was to test if spatial dynamics could be related to spatial isolation and disturbance of habitat and to the dispersal and seed bank characteristics of the species. Knowledge of these relations should be used to increase the effectivity of agri-environmental schemes aiming at an increase of botanical diversity. All species frequently colonized empty patches and populations in occupied patches frequently went extinct. Most colonization events occurred within 50 m of conspecific source patches in the preceding year, but colonization events in patches at distances more than 200 m from conspecific source patches were also observed. The colonization probabilities decreased with isolation distance. For nine species this relation was statistically significant, after correction for year and habitat. The extinction probabilities increased with isolation. For only four species this relation was statistically significant. Both colonization and extinction probabilities were more often statistically significant related to isolation for species with transient seed banks than species with persistent seed banks. Implications for management options aiming at survival of plant species in fragmented landscapes are discussed.     

Forest species in an agricultural landscape in the Netherlands: Effects of habitat fragmentation

Abstract. For 312 forest patches on sandy soils in the Netherlands, effects of fragmentation are studied of forest habitat in the past on the present occurrence of forest plant species. Using regression techniques, the numbers of forest edge, interior, zoochorous and anemochorous species, as well as occurrence of 24 individual species were related to patch area and connectivity measures. Connectivity was defined as the amount of forest habitat around patches within three zones up to 1000 m. Plant categories were distinguished by habitat type and dispersal mechanism. The results showed that number of total species and number of species of all habitat and dispersal categories increased with area. The occurrence of ten individually studied species were also positively related to area. Most of them were interior species. The number of zoochorous species increased with increasing connectivity. Also occurrence of ten individually studied species were affected by connectivity. Interior zoochorous species showed the highest percentage of affected species. The relationship of interior, animal-dispersed plants to connectivity can be explained by the limited distances covered by their dispersal agents (forest birds and ants) in a non-forest habitat. Also, some anemochorous plants appeared to be affected by connectivity, especially those with heavy seeds and potentially short distance dispersal. As not all species within a certain dispersal or habitat category react similar to area or isolation, it is suggested that differences in underlying processes of fragmentation such as local extinction and colonization need more focus.     

长白山红松阔叶林主要树种高度生态位的研究

森林生态系统内主要乔木的高度生态位反映了不同树种在各个高度资源单位内资源利用的互补情况,同时也是表征森林生态系统演替的趋势及其森林循环过程主要特征之一．生物学特性和生态学特性接近的树种的高度生态位重叠较大,反之则较小．在Ｇａｐ内,色木槭和扭筋槭的高度生态位重叠最大,而青楷槭和千金榆、花楷槭和水曲柳、青楷槭和裂叶榆的高度生态位重叠较小;在非Ｇａｐ生态系统内,青楷槭和白牛槭的高度生态位重叠最大,青楷槭和裂叶榆、青楷槭和千金榆、花楷槭和水曲柳的高度生态位重叠较小．红松和阔叶树之间的高度生态位分离较大,利用资源的方式和能力差异较大,它们之间可以在不同空间的资源利用上达到互补,种间竞争较弱,从而能够保持红松和阔叶树之间长期而稳定的共存．但Ｇａｐ内红松和阔叶树的高度生态位重叠均大于非Ｇａｐ生态系统,可见Ｇａｐ的存在加大了红松和阔叶树之间的种间竞争,减小了红松和阔叶树之间的资源互补利用的能力．Ｇａｐ削弱了个体生态位重叠值较大的树种间的种间竞争,增强了个体生态位重叠值较小的树种间的种间竞争     

Flowering,fruiting and seed germination in Chilean rain forest myrtaceae: ecological and phylogenetic constraints

Phenological studies in plant communities have generally focused on taxonomically heterogeneous species assemblages, and have only occasionally examined the evolutionary and ecological constraints on the phenological patterns of species within a single family or a genus. Here, we determine relative importance of phylogenetic versus other constraints on the flowering and fruiting periods of 12 species and the germination ecology of 10 species of Myrtaceae sympatric to the temperate rainforest of Isla Grande de Chiloé (42 °Cs 30 S), in southern Chile.We found that, for most species in the family Myrtaceae, flowering was strongly aggregated in January and February. Although this pattern is consistent with the expectation of the 'facilitation' hypothesis (i.e., interspecific overlaps are maximized to attract pollinators), available evidence suggests that pollinators, mainly hymenopterans and dipterans, do not limit fruit production in these species of Myrtaceae in the temperate rainforest. In contrast to flowering, fruiting occurred all-year-round, showing greater segregation in time among the species. According to the their temporal patterns of fruit ripening, two functional groups were defined within the Myrtaceae: those that ripen their fruits immediately after flowering (species in the subtribe Myrtinae) and those in which green fruit develops slowly for several months before ripening (subtribe Myrciinae). Seed germination in the field occurred mainly between August and October. Lab assays showed that the species of Myrtaceae, subtribe Myrtinae, exhibited a long seed dormancy (>40 days), while the seeds of species in the subtribe Myrciinae often germinated within one week after harvesting. The analysis of the phenology of reproductive events in the species of Myrtaceae in this rainforest suggests that: (1) flowering periods patterns are constrained mainly by phylogenetic inertia at the family level, and (2) differences in fruiting patterns and dormancy periods are determined mainly by fruit and seed size, which in turn are associated primarily with phylogenetic closeness within the family, and secondarily with the activity of vertebrate seed dispersers.     

雾灵山国家自然保护区森林群落物种多样性研究

在样方法获取全面资料并用数量方法确定森林群落类型的基础上,对雾灵山国家自然保护区各森林群落的物种多样性(D)、生态优势度(C)、群落均匀度(J)等指标进行了计测;并着重对物种多样性进行评价,分析了物种多样性变化的时空规律,就物种多样性的保护与持续利用进行探讨。分析结果表明：雾灵山国家自然保护区森林群落物种多样性比较丰富,其D值在2.37～4.17之间;在特定高度范围内,随海拔增高D值增大,C值变小;多数森林群落灌木层的D值高于乔木层,C值低于乔木层;地带性群落的D值大于非地带性群落,C值相反。     

Sunken roads as habitats for forest plant species in a dynamic agricultural landscape: effects of age and isolation

Aim This study tests the hypothesis that linear, woody habitat patches surrounding small, sunken rural roads not only function as an unstable sink but also as a true, sustainable habitat for forest plants. Furthermore, factors affecting the presence of forest plant species in sunken roads are determined. Finally, the implications of these findings for the overall metapopulation dynamics of forest plant species in fragmented agricultural landscapes are assessed. Location The study area, c. 155 km² in size, is situated in a fragmented agricultural landscape within the loamy region of central Belgium. Methods Forest species presence–absence data were collected for 389 sunken roads. The effect of area, depth, age and isolation on sunken road species richness was assessed using linear regression and analysis of variance (anova ). Analysis of covariance was employed to study the interaction between age and isolation. Differences in plant community dispersal spectra in relation to sunken road age and isolation were analysed by means of linear regression and anova . Results Sunken roads proved to function as an important habitat for forest plants. The sink‐hypothesis was falsified by a clear species accumulation in time: sunken road species richness significantly increased with the age of the elements. Sunken road age mainly affected species richness through effects on both area and depth, affecting habitat quality and diversity. Furthermore, sunken road isolation had a significant impact on species richness as well, with the number of forest species decreasing with increasing isolation of the elements, indicating dispersal limitation in sunken road habitats. Moreover, a significant age × isolation interaction effect was demonstrated. Differences in regression slopes for isolation between age classes revealed that the effect of isolation intensified with increasing age of the elements. Differential colonization in relation to forest species dispersal capacities probably account for this, as confirmed by the analysis of sunken road plant community dispersal spectra, with the fraction of species with low dispersal capacities increasing with increasing age and decreasing isolation of the elements. Main conclusions During sunken road development, area and depth increase and, gradually, suitable habitat conditions for forest plant species arise. Depending on their ecological requirements and dispersal capacities, forest species progressively colonize these habitats as a function of the element's isolation. The functioning of sunken roads as a sustainable habitat for forest species enhances the metapopulation viability of forest plants in agricultural landscapes and has important consequences for forest restoration practices. Moreover, the results of this work call for integrating the presence of forest species in small‐scaled linear habitat patches in forest fragmentation studies.     

长白山红松阔叶混交林林冠空隙树种更新动态规律的研究

分析了长白山红松阔叶混交林林冠空隙(Gap)内更新树种的数量特征.结果表明, 在34个林冠空隙中, 更新乔木树种出现频度为7.27%, 更新灌木树种出现频度为21.02%, Gap形成后, 阳性先锋树种首先侵入, 出现频度较高, 占据较宽的生态位, 随着Gap年龄的增加, 树种间的竞争逐渐增大, 阳性树种的生长逐渐受到限制, 阴性树种逐渐增多;Gap年龄与Gap郁闭度和Gap面积呈负相关, Gap面积与树种出现频度呈负相关;Gap年龄主要分布在5～15a和46～55a两段期间, 树种出现频度与Gap年龄存在着密切关系, 当Gap年龄小于25a时, 二者呈正相关;当Gap年龄在25～40a时, 二者呈负相关;当Gap年龄大于40a时, 树种出现频度趋于稳定;34个Gap内4种主要乔木更新树种和4种主要灌木更新树种的胸径均基本上呈正态分布, 乔木更新树种的最高点胸径为4～6cm径级, 灌木更新树种的最高点胸径为1.2cm径级.     

不同季节亚热带常绿阔叶林6个树种凋落叶钙、镁、锰的释放特征

采用凋落物分解袋法,研究了亚热带常绿阔叶林区马尾松、柳杉、杉木、香樟、红椿、麻栎6个典型树种凋落叶的Ca、Mg、Mn在第一个分解年不同雨热季节的释放特征.结果表明：经历1年的分解,6种凋落叶Ca、Mg、Mn释放率分别为-13.8%～92.3%、4.0%～64.8%和41.6%～81.1%.马尾松和香樟凋落叶Ca释放动态呈现富集-释放模式,其余4种凋落叶整体上呈现释放的模式;香樟凋落叶Mg释放动态呈现富集-释放模式,其余5种凋落叶呈现直接释放模式;柳杉和红椿凋落叶Mn释放动态呈现富集-释放模式,其余4种凋落叶呈现直接释放模式.凋落叶Ca、Mg、Mn的释放明显受到季节性降雨的影响,且因凋落叶种类不同而有差异.Ca、Mg、Mn在雨季的释放率和释放量均高于旱季.初始养分含量和降雨量是影响凋落物分解过程元素释放的重要因子.     

Soil seed bank composition and diversity in a managed temperate deciduous forest

Little is known about the influence of forest management on the interaction between seed bank and aboveground vegetation. We surveyed seed banks and vegetation in 10 forest stands under similar abiotic conditions but submitted either to a coppice-with-standards treatment (n=5) or to a selective-cutting system (n=5). We analyzed species composition and diversity, community ecological profile, and distribution of taxa among different life forms, strategy, morphology and functional type categories. A total of 2085 seedlings (8296 seedsm^–2) germinated-corresponding to 28 species, among which Juncus effusus was the most abundant. Fifty-seven percent of the species were also recorded in the aboveground vegetation, the dominant species being Rubus fruticosus agg., but only 28% of the aboveground species were present in the seed bank. Our results suggest that (1) vernal geophytes and shade-tolerant perennials, which group most true forest species, are not incorporated in the seed bank, (2) parent plants of most seeds were present either in the stand in an earlier dynamic stage or apart from the stand and long-distance dispersed, (3) as expected, early-successional species are well represented in the seed bank, (4) forestry vehicles seem to be a major means of dispersion for stress-tolerant species normally found in forest lanes and wheel tracks. We conclude that seed banks contain species that have a potentially negative impact on the true forest flora and, thus, forest management should minimize soil disturbance and retain remnants of old-coppice woods to conserve disturbance-sensitive true forest species.     

Gap-phase regeneration in a tropical montane forest: the effects of gap structure and bamboo species

To study the influence of gap structure and bamboo species on the regrowth of montane Atlantic forest, colonization by plants was characterized in 30 treefall gaps (30.3–500.5 m²). The study was conducted at Santa Virgínia (45°30 W, 23°17 S), a 4970-ha reserve of Atlantic montane forest in southeastern Brazil. Area covered by bamboos ranged from 0% to 100% of gap area. Average height of surrounding canopies ranged from 12 to 30 m. As gap are covered by bamboo and average height of surrounding canopies increased, both density and richness of pioneer woody species decreased. Density and richness of shade-tolerant species were negatively influenced by gap area. Low-light-demanding species of Miconia, Leandra and Rapanea accounted for the majority of both pioneer species and individuals sampled, whereas high-light demanding pioneers of Cecropia, Alchornea and Tibouchina were poorly represented. We suggest that in the Atlantic montane forest bamboo species compete for gaps, excluding other light-demanding pioneers. This results in an overall reduction of pioneer species richness in the Atlantic forest.     

应用指示种预测森林管理对物种多样性及群落组成的影响

采用指示种分析方法,研究了会同亚热带森林物种多样性和群落组成对森林管理的响应.从357个林下种中鉴定出显著性指示种94个,并构造新的指示种数据集,检验指示种数据集和源群落数据集之间的关联,评估指示种对林下植被管理效应的预测潜能.结果表明:指示种数据集和源群落数据集之间存在极显著的关联(Mantel r=0.898),指示种数据集很好地预测了生物多样性的变化(回归分析,R2>0.74);指示种很好地预测了群落组成对森林管理的响应(ANOVA,F>16.79);非度量多尺度排序(NMDS)以及K-means聚类分析表明,对于不同森林管理的样地类型,指示种数据集的识别能力和源群落数据集是一致的.从物种多样性、群落组成以及在森林类型的识别上,指示种数据集和源群落数据集有一致性规律,作用几乎相同,因此森林评估可以利用指示种代替源群落预测森林管理效应,以减少森林全面调查的成本.     

Effects of environment,habitat configuration and forest continuity on the distribution of forest plant species

In present day European landscapes many forest plant species are restricted to isolated remnants of a formerly more or less continuous forest cover. The two major objectives of this study were (1) to determine the relative importance of habitat quality (mainly in terms of soil parameters), habitat configuration (patch size and isolation) and habitat continuity for the distribution of herbaceous forest plant species in a highly fragmented landscape and (2) to examine if groups of species with different habitat requirements are affected differently. Deciduous forest patches in northwestern Germany were surveyed for the presence of a large set of forest species. For each patch, habitat quality, configuration and continuity were determined. Data were analysed by Redundancy Analysis with variation partitioning for effects on total species composition and multivariate logistic regression for effects on individual species, for two different data sets (base‐rich and base‐poor forest patches). Overall, we found strong effects of habitat quality (particularly of soil pH, water content and topographic heterogeneity in the base‐rich forest patches; and of calcium content and disturbance in the base‐poor patches), but only relatively weak effects of habitat configuration and habitat continuity. However, a number of species were positively affected by patch area and negatively affected by patch isolation. Furthermore, the relative importance of habitat configuration tended to be higher for species predominantly growing in closed forests compared to species occurring both in the forest and in the open landscape.     

人为干扰对闽北森林群落物种多样性的影响

本文从群落及其组成结构等方面分析了闽北长期封禁保护的森林（ｃｏｎｓｅｒｖｅｄｆｏｒｅｓｔ,ＣＦ）和人为干扰后的退化森林（ｄｅｇｒａｄｅｄｆｏｒｅｓｔ,ＤＦ）群落物种多样性,结果认为,ＤＦ群落的植物物种丰富度和ＳｈａｎｎｏｎＷｉｅｎｅｒ指数与ＣＦ群落相近或略高,但ＤＦ群落植物种类组成和层次结构的物种多样性发生明显变化,群落物种种群基础已改变,稀有种和渐危种缺乏,这正是ＤＦ与ＣＦ群落物种多样性的本质差异。     

The impact of forest continuity and management on forest floor vegetation evaluated by species traits

The distribution of the flora with different species attributes are investigated in two studies of forest ecosystems in Denmark, One study compared 17 forests with different degrees of management, the other study compared the flora of 25 old forests with that of 6 new forests. No effect of forest management could be detected in the distribution of species attributes in the flora, whereas the forest continuity was correlated with almost all of the 14 traits investigated. Species with heavy seeds, transient seedbanks, ant dispersed seeds, early and short flowering time, low stature and high extent of lateral spread were more common in old forests, while the proportion of species with small, short-lived seeds, epizoochorous dispersal, little lateral spread and later and longer flowering period was higher in new forests. In particular, Raunkiaers therophyles, and ruderal species were more common in new forests than in old.
Differences in the effects of the two types of disturbance - forest management, and breaks in forest eontinuity - are discussed in relation to survival strategies of the forest flora. The importanee of persistence ability in contrast to colonisation ability is stressed for many forest plants.     

常绿阔叶林片段中木荷凋落叶分解速率及中小型土壤节肢动物群落的结构动态

2004年6月-2006年4月,选择浙、闽、赣交界山地的4个森林片段作为研究地点,并以浙西的古田山国家级自然保护区作为连续森林对照,采用尼龙网袋法,研究了不同常绿阔叶林群落优势种木荷凋落叶的分解速率及土壤节肢动物群落的结构和动态.结果表明：5个研究地共获土壤节肢动物1050个,分属8纲23目,其中鳞翅目、膜翅目、弹尾目和双翅目占个体总数的10%以上,为该地区的优势类群.片段化对群落物种组成的影响主要体现在稀有类群的差异上.不同分解阶段各种土壤节肢动物类群优势度因其功能的不同而各异.经过2年的分解,各研究地木荷凋落叶损失量为60%～70%,研究地凋落物中土壤节肢动物的各项多样性指标均有一定的变化,且与连续森林的变化规律不一致.     

Variation in tree growth,mortality and recruitment among topographic positions in a warm temperate forest

Questions: Do the population dynamics of trees differ among topographic positions and, if so, how does topographic position affect the population dynamics of species that are distributed in a topography‐specific manner? Which is the most important life stage in determining vegetation patterns? Location: Primary and secondary warm temperate evergreen broad‐leaved forest (40 ‐ 280 m a.s.l.) on the western part of Yakushima Island, Japan. Methods: Mortality, recruitment, DBH growth and distribution of stems (= 5 cm DBH) in a 2.62‐ha plot were surveyed in 1992 and 2002 to determine the relationships between population parameters and (1) topography and (2) distribution patterns of 17 common tree species. Results: Common species (n = 17) were classified into three distribution pattern groups: group A, distributed mainly on convex slopes; group B, on concave slopes, and group C, not aggregated with respect to topographic position. Stem mortality, recruitment and DBH growth were greater in group A than in group B within each topographic class. The hierarchy of stem mortality among topographic classes for groups A and B was convex > planar > concave. Stem recruitment density was relatively high on the convex and concave slopes, respectively, for groups A and B. Conclusions The topographical positions of adult trees were not always most suited for adult survival and growth. For group A, the distribution pattern of adults was determined in the juvenile stage, while this was not the case for group B. Studies of juvenile stages are important for understanding the demographic basis of vegetation distribution patterns.     

浙江天台山常绿阔叶林优势种群结构及种间联结性研究

以浙江天台山常绿阔叶林为研究对象 ,用“空间序列代替时间变化”的方法 ,对该群落优势种群的结构和动态进行了分析 ,结果是 :甜槠、木荷、东南石栎种群的存活曲线均表现为极显著的线性关系 ,种群的年龄结构为稳定型或增长型 ,马尾松、短柄种群的年龄结构为衰退型 ,小叶青冈种群的年龄结构由稳定型趋向衰退型。用方差比率 (VR)和通过 2× 2联列表进行 X2 检验的方法 ,检验了常绿阔叶林乔木层 2 0个主要树种、灌木层 2 2个主要种类的总体相关性和各种对间的联结性。结果是乔木层、灌木层主要种类间在总体上均呈显著的正相关 ,正联结的种对多于负联结的种对。