Behaviour and ecology of birds in tropical rain forest canopies

The canopy of rain forests exhibits several distinct features that determine avian adaptations. Light intensity and colour, wind, and temperature are the most important components of the physical environment. Tree architecture poses many challenges for the evolution of avian morphological features, and resource availability varies greatly. The canopy is easily accessible for birds that are equipped to travel over considerable distances. Given these conditions, high behavioural flexibility, and adaptations in behaviour and morphology related to a nomadic life-style, should prevail among canopy birds. The easy access for certain groups of birds also has consequences for the historical processes that shape canopy bird communities. Birds may contribute greatly to forest dynamics and composition, mainly due to their role as seed dispersers. We elaborate on these points with behavioural, morphological, and ecological data collected at our Neotropical study site in southern Venezuela (Surumoni Canopy Crane Project). We discuss how behaviour and morphological features reflect the ecological demands in the canopy. We contrast the specific behavioural and ecological characteristics of canopy birds with those of species found in the other tiers of the rain forest. It is pointed out that much of our knowledge is biased towards the Neotropics, and that better knowledge of the larger canopy species is needed for conservation.     

Microgeographical variation in thermal preference by an amphibian

Ectotherms use behaviour to buffer effects of temperature on growth, development and survival. While behavioural thermoregulation is widely reported, localized adaptation of thermal preference is poorly documented. Larval amphibians live in wetlands ranging from entirely open to heavily shaded by vegetation. We hypothesized that populations undergo localized selection leading to countergradient patterns of thermal preference behaviour. Specifically, we predicted that wood frog (Rana sylvatica) larvae from closed canopy ponds would be more strongly temperature selective and would prefer higher temperatures than conspecifics from populations found in open canopy ponds. In a study of six breeding ponds in north‐eastern Connecticut, USA, these predictions were upheld. The countergradient, microgeographical variation in thermal preference documented here implies that wood frog populations may have diverged rapidly in the face of contrasting selection pressures. Rapid, behaviourally mediated responses to changing thermal environments have important implications for understanding population responses to climate change.     

A bird's-eye view: Evaluating drone imagery for the detection and monitoring of endangered and invasive day geckos

Species monitoring can be strongly limited by terrain accessibility, impeding our understanding of species ecology and thus challenging their conservation. This is particularly true for species living in the canopy, on cliffs or in dense vegetation. Remote sensing imagery may fill this gap by offering a cost-effective monitoring approach allowing to improve species detection in inaccessible areas. We investigated the applicability of drone-based monitoring for a Critically Endangered insular gecko (Phelsuma inexpectata) and two invasive alien gecko species representing a risk for the former (P. grandis and P. laticauda). We determined the approach distance before inducing behavioural response caused by the drone's presence. All three study species showed no escape behaviour to the drone's presence until very close distances (mean distance for P. inexpectata: 33.8 cm; P. grandis: 21.9 cm; P. laticauda: 26.4 cm). We then performed horizontal and vertical approaches with the drone, taking photos every meter starting at 10 m away from the canopy edge to determine an optimal distance for detection while ensuring species-level identification. We examined a total of 328 photos. We found a bimodality in the number of detected geckos for two species, with different individuals recorded between short and intermediate distances. Therefore, we recommend taking photos at two distances of 2–2.5 and 5 m away from the canopy, ideally facing away from the sun and in low wind conditions. We encourage the application of our methodology for Phelsuma spp., but also for other species of similar size and ecology to improve detection in inaccessible areas.     

Relationships between wild bee abundance and fruit set of Camellia oleifera Abel

Camellia oleifera Abel. is an economically important plant for edible oils. To investigate the importance of wild bee abundance at different canopy and spatial scale for fruit set of Camellia oleifera, we evaluated the relationship of bee abundance with fruit set using yellow pan trap, with additional focus on distance and wild bee abundance counts at a farm in China for 2 years. We found that yellow traps collected 95% more individual bees than the second-best (pink) trap colour. Looking across canopy positions, the number of wild bees (including Colletes gigas, Andrena camellia, Andrena striata, Andrena hunanensis and Andrena chekiangensis) varied significantly in each year. The fruit set also varied significantly based on canopy position. In 2018, there was with an average fruit set of 34.13% in the lower canopy, 54.30% in the middle canopy and 47.41% in the upper canopy. In 2019, fruit set was 29.67% in the lower canopy, 54.15% in the middle canopy and 47.36% in the upper canopy. Comparing inter-annual changes, only the fruit set and number of bees in the low canopy showed significant differences between 2018 and 2019. Fruit set and wild bee abundance were positively correlated in all canopy layers in each year, and there was an interactive effect between canopy and bee abundance on fruit set. Across 205 trees, growth indicators, crown size and tree height, increased significantly from 2018 to 2019, but the fruit set did not increase with the significant increase in the number of flowers. From the spatial perspective, wild bee abundance and fruit set were positively correlated, but fruit set was negatively correlated with distance from a large bee nesting aggregation nearby. Among the variables investigated, bee abundance seems to have the strongest influence on fruit set in C. oleifera.     

Frankia and Alnus rubra Canopy Roots: An Assessment of Genetic Diversity, Propagule Availability, and Effects on Soil Nitrogen

The ecological importance of microbial symbioses in terrestrial soils is widely recognized, but their role in soils that accumulate in forest canopies is almost entirely unknown. To address this gap, this study investigated the Frankia–Alnus rubra symbiosis in canopy and forest floor roots at Olympic National Park, WA, USA. Sixteen mature A. rubra trees were surveyed and Frankia genetic diversity in canopy and forest floor nodules was assessed with sequence-based nifH analyses. A seedling bioassay experiment was conducted to determine Frankia propagule availability in canopy and forest floor soils. Total soil nitrogen from both environments was also quantified. Nodules were present in the canopies of nine of the 16 trees sampled. Across the study area, Frankia canopy and forest floor assemblages were similar, with both habitats containing the same two genotypes. The composition of forest floor and canopy genotypes on the same tree was not always identical, however, suggesting that dispersal was not a strictly local phenomenon. Frankia seedling colonization was similar in canopy soils regardless of the presence of nodules as well as in forest floor soils, indicating that dispersal was not likely to be a major limiting factor. The total soil nitrogen of canopy soils was higher than that of forest floor soils, but the presence of Frankia nodules in canopy soils did not significantly alter soil nitrogen levels. Overall, this study indicates that the Frankia–A. rubra symbiosis is similar in canopy and forest floor environments. Because canopy roots are exposed to different environmental conditions within very small spatial areas and because those areas can be easily manipulated (e.g., fertilizer or watering treatments), they present microbial ecologists with a unique arena to examine root–microbe interactions.     

冠层高度对毛竹叶片光合生理特性的影响

借助LI-6400便携式光合作用系统,研究了冠层高度对不同林龄毛竹(Phyllostachys pubescens)叶片光合生理特性和水分利用效率(WUE)的季节性影响,为促进毛竹林碳汇能力和生产力提升的林分结构调整等可持续栽培技术提供理论依据。结果表明:(1)出笋期,不同竹龄毛竹叶片净光合速率(Pn)和蒸腾速率(Tr)的日均值呈现出冠层上部小于冠层下部的梯度变化趋势,且2a生毛竹不同冠层Pn日均值大于3a生毛竹;孕笋行鞭期,不同林龄毛竹各时间点Pn值和日均值、以及2年生毛竹各时间点的Tr值均为冠层上部大于冠层下部。各生长季节,不同林龄毛竹个体叶片的气孔导度(Gs)均与Tr的变化趋势一致。(2)2年生毛竹各季节仅冠层上部叶片会出现"光合午休",而3年生毛竹仅于出笋期时各冠层叶片出现"光合午休"现象。(3)出笋期毛竹叶片WUE日均值随着冠层高度增加而增加,这种变化趋势不受竹龄影响;而孕笋行鞭期,仅2年生毛竹叶片WUE日均值随着冠层高度增加而下降。不同冠层高度的孕笋行鞭期毛竹叶片WUE日均值都显著高于出笋期;冠层高度对毛竹叶片气体交换特性和WUE的影响受生长发育关键期的季节因素影响,且毛竹叶片WUE与Gs之间存在负相关关系,其不受毛竹个体年龄和叶片冠层高度影响。(4)不同生长季节各冠层叶绿素a/b值均随着冠层高度下降而降低,不同林龄毛竹叶片叶绿素含量基本随着冠层自上而下呈逐渐增加的趋势。各生长季节,不同林龄个体叶片氮素含量、比叶重随冠层高度垂直变化趋势与叶片Pn日均值的垂直变化趋势一致。研究认为,毛竹不同冠层部位叶片通过改变形态、氮素含量来适应不同生长季节生长环境的变化,以便充分利用光能提高光合能力。     

Spatial dynamics of regeneration in a conifer/broad‐leaved forest in northern Japan

Abstract. This study deals with stand dynamics over a 6‐yr period in a conifer/broad‐leaved mixed forest in Hokkaido, northern Japan. The annual rates of gap formation and recovery were 81.3 m²/ha and 66.7 m²/ha, respectively and turnover time of the canopy was 125 yr. The recruitment processes of the component species in this cool‐temperate forest were governed by different canopy types: gap, canopy edge and closed canopy. Magnolia obovata regenerated in canopy edges, and Acer mono and Prunus ssiori regenerated in canopy edges and gaps. The results suggested that the mosaic structure made up of closed canopy, canopy edge and gap was related to various regeneration niches. Abies sachalinensis had high mortality rates, initiating gap expansion. The transition probabilities from closed canopy or canopy edge to gap for deciduous broad‐leaved trees were lower than for A. sachalinensis, which implies that the difference in degeneration patterns of conifer and broad‐leaved canopies contributes to the heterogeneity of spatial structure in the mixed forests. Spatial dynamics were determined by a combination of gap expansion by A. sachalinensis (neighbour‐dependent disturbance) and gap formation by deciduous broad‐leaved trees (random disturbance).     

Differences between understorey and canopy in herbivore community composition and leaf quality for two oak species in Missouri

1. From July 1994 to September 1995, at six censuses, the herbivore community associated with understorey (< 2.5 m height) and canopy (15–20 m) leaves of Quercus alba and Q. velutina was sampled in south-eastern Missouri, U.S.A. 2. Across all censuses, herbivore densities were not significantly different between canopy and understorey for Q. alba and Q. velutina, except in August 1994 when herbivore densities were 60% higher in the canopy on Q. alba. Little significant spatial variation in herbivore densities or community composition was found during the study years. 3. The herbivore community was diverse, consisting of 138 species of leaf-chewing insects. Species richness was significantly greater (by 5–20%) in the understorey than in the canopy for both tree species, and the relative abundance of the main families, different feeding guilds, and most common species differed significantly between understorey and canopy. 4. To determine the extent to which leaf quality explained the observed patterns, percentage nitrogen and protein binding capacity were measured in canopy and understorey leaves of Q. alba and Q. velutina. Per cent nitrogen was higher in canopy leaves for Q. velutina while protein binding capacity was higher in canopy leaves for Q. alba. 5. These results suggest that the herbivore community associated with these two species of Quercus comprises species that appear to respond individually to environmental and biological conditions encountered in the understorey and the canopy.     

华北落叶松人工林蒸散及产流对叶面积指数变化的响应

定量评价林地蒸散和产流等水文过程对冠层叶面积指数(LAI)的响应,对于深入认识森林植被的生态水文过程及其发生机制,实现半干旱区林水综合管理和区域可持续发展是非常必要的。应用集总式生态水文模型BROOK90,模拟分析了不同降水年型(丰水年、平水年、枯水年)下,位于半干旱区的宁夏六盘山叠叠沟小流域内华北落叶松(Larix principis-rupprechtii)人工林的水文过程对冠层LAI变化的响应关系。结果发现:林分总蒸散量、冠层截留量、蒸腾量与LAI都呈显著的正相关关系(R~20.99,P0.01),而土壤蒸发量、产流量则与LAI均呈显著的负相关关系(R~20.99,P0.01);在不同的降水年型下,各水文过程变量与LAI的关系都可以很好地用指数函数来表达,且都存在着一个LAI阈值。当LAI低于阈值时,各水文过程变量随LAI的变化幅度较大;但高于阈值时,各变量的变化十分缓慢并趋于稳定。在不同降水年型下,各变量LAI阈值之间存在着一定的差异。一般地,丰水年各变量的LAI阈值要大于枯水年,尤其是冠层截留和土壤蒸发。在丰水年,各水文过程变量随LAI增加而变化的速率要比在平水年、枯水年更快,说明在水分充足年份中各变量的波动更多取决于LAI变化,而在水分亏缺的年份中则可能更多地受到水分条件的限制。模拟结果表明,通过减少冠层LAI(如间伐)导致的林分的降低蒸散耗水和增加产流的作用是有限的,这是由于林分蒸散降低的幅度要比LAI降低的幅度小。例如,在平水年,当LAI从4.2变为2.0(减少幅度52.4%)时,林分年蒸散仅从357.2 mm减少至333.9 mm(减少幅度6.5%)。     

大兴安岭北部兴安落叶松(Larix gmelinii)林下穿透雨空间分布特征

森林冠层对降雨的水量和水质再分配是生态水文学研究的热点问题之一。为了研究兴安落叶松林下穿透雨的空间分布规律,探究森林冠层结构对穿透雨影响的生态机制,利用在兴安落叶松林下布设38个雨量筒,测定19场不同降雨事件的穿透雨数据(2013年7—8月),通过统计学方法分析冠层结构各因子与穿透雨的空间变异性规律,结果表明:观测期间,兴安落叶松林穿透雨量为148.3 mm,占同期大气降雨量的80.62%,穿透雨率随着降雨量的增加呈增加趋势;兴安落叶松林下穿透雨具有较大空间异质性,其变异程度随降雨量的增加而减小,以对数方程拟合较好(P0.01);冠层结构特征是影响穿透雨空间变异的重要因素,冠层复杂程度与穿透雨量呈负相关关系(P0.01);距树干距离、冠层厚度、叶面积指数等因素均可影响穿透雨的空间分布,以距树干距离影响最大,其与穿透雨率呈正相关关系(P0.01),而冠层厚度、叶面积指数则均与穿透雨率呈负相关关系(P0.01),但拟合效果不佳;从影响穿透雨的生态学机制来考虑,在冠层结构特征因子中,冠层厚度是决定穿透雨空间分布的最主要因素。     

Distribution and regeneration strategies of major canopy dominants in species-rich subtropical/warm temperate rainforests in south-western Japan

The distribution and regeneration strategy of the major canopy dominants in species-rich subtropical/warm temperate rainforests in south-western Japan was studied in a hilly zone below 1100 m a.s.l. Using the patch sampling method, four dominance-community types were numerically identified and they corresponded to four habitats which represented a combination of topography and altitude (i.e. ridgesvs slopes, and belowvs above 500 m a.s.l.) Seven major canopy dominants had their respective distributional core in one of these topo-altitudinal habitats. The seven canopy dominants could be classified into three species groups according to their size structure and growth habits. Group A (Tsuga sieboldii andQuercus gilva) and group B (Quercus acuta, Quercus salicina, Machilus thunbergii) were restricted to the emergent and canopy layer, respectively. These two groups had only a few subcanopy trees and saplings. Saplings of group B showed a rapid growth rate in canopy gaps. Group C (Distylium racemosum) was characterized by many subcanopy trees and saplings that grew steadily under the closed canopy.Castanopsis sieboldii showed intermediate characteristics between group B and C in the size structure and growth habit of its saplings. The density ofD. racemosum canopy trees was markedly reduced on ridges and slopes above 500 m and on slopes below 500 m. In these marginal habitats, the three species groups coexisted by sharing different strata within a community. This situation was possible due to the differences in regeneration strategies among the canopy species.     

Relationship of Heliothis zea predators, parasitoids and entomopathogens to canopy development in soybean as affected by Heterodera glycines and weeds

The influence of canopy development in soybean on the survival of corn earworm, Heliothis zea (Boddie) (Lepidoptera: Noctuidae), egg and larval stages and population dynamics of arthropod fauna were evaluated in field trials during 1986–88 in eastern North Carolina. Soybean canopy size decreased as soybean cyst nematode, Heterodera glycines Ichinohe (Nematoda: Heteroderidae), initial population densities increased. Plant species composition of the soybean canopy was affected by weed population densities. Mortality of H. zea larvae due to parasitism and infection with entomopathogens was greater in closed canopy and (or) weedy soybeans than in very open and (or) weed free soybeans. Predation and parasitism of corn earworm eggs were similar across nematode and weed density treatments. Natural enemy populations increased to highest levels during July in closed canopy and (or) weedy soybeans, coinciding with availability of largest prey population reservoirs. A delay in colonization of very open and (or) weed free soybeans by beneficial arthropods until mid to late August allowed greater H. zea larval survival than in closed canopy and (or) weedy soybeans. Arthropod species richness was generally greatest in closed canopy and (or) weedy soybeans during mid to late July, with differences becoming nonsignificant in August and early September. Mean and maximum ambient temperatures were higher and relative humidities lower in open canopy than in closed canopy plots. These conditions were less favorable for development of pathogens and natural enemies.     

河西走廊沙漠泡泡刺群体光合特征及其与环境因子的关系

以河西走廊沙漠典型的荒漠植物泡泡刺群落为材料,运用LI-8100土壤碳通量测定系统与改进的同化箱联合对其7月和8月中旬的群体光合作用进行了测定,分析群体光合速率与立地环境因子的关系,探讨泡泡刺群体对干旱荒漠环境的光合适应机制。结果表明,7月中旬泡泡刺的土壤呼吸速率和蒸发速率均高于8月中旬,而其同期群体光合速率和水分利用效率也显著高于8月中旬。两时期的光合有效辐射是影响泡泡刺群体光合速率的主要因子,于7、8月中旬对泡泡刺群体的光合速率起着直接作用;同时,实验地8月中旬的空气相对湿度也能通过与大气温度以及光合有效辐射的相互作用对泡泡刺群体的光合速率产生较大影响。在7月中旬高温强光环境下,泡泡刺的群体光合速率高于8月中旬,从而说明泡泡刺对高光强、高温的荒漠环境具有较好的适应性。     

Patterns of cytokinins and leaf gas exchange among canopy layers of a mature sugar maple (<Emphasis Type="Italic">Acer saccharum</Emphasis>) stand

Leaf cytokinins (CKs) were profiled within four locations throughout the inner and outer layers of a mature sugar maple (Acer saccharum) canopy. Leaf CK was associated with leaf gas exchange activity and some corresponding microclimate variables. Both inner and outer layers in the upper canopy had higher concentrations of leaf CKs than the lower canopy layers and the difference was comprised primarily by riboside forms of CK. Transpiration (E) showed a similar pattern to leaf CK content, with significantly higher rates in the upper canopy. There was, however, no clear pattern discernable in stomatal conductance (gs), other than it tended to be higher in the outer canopy layers. The upper/outer canopy showed a significantly different environment than all other canopy positions with higher photosynthetically active radiation (PAR), ultra-violet light (UV-B) and leaf temperature. Simple linear regression analysis showed that the nucleotide CK group (including iPNT, cis- and trans-[9RMP]Z, [9RMP]DZ) was positively related to PAR. Exogenous applications of benzylaminopurine (BAP), showed that low concentrations of BAP reduced E and g _s, and indicated that CK may help regulate stomatal aperture. The similar patterns in E and CK content suggest that CKs and leaf gas exchange are functionally connected.     

Canopy dynamics estimated from shoot morphology in an evergreen broad-leaved forest

Summary A forest canopy structure may be defined as the spatial distribution pattern of foliage density, and dynamics of canopy can be considered as changes of spatial distribution of foliage density. To study this process, the annual intrinsic growth factor (r) of foliage and the speed with which foliage shifts its position were estimated from shoot branching and shoot length. The spatial distributions of these parameters were obtained from a profile of evergreen broad-leaved forest. r was large in the upper canopy layer and canopy gap; this indicated the active development of foliage. This phenomenon may be a major reason for the existence of dense foliage in the upper canopy. The speed with which foliage shifts its position was high in the canopy gap. For dominant species, light conditions affected positively on the distribution of r.     

Tree species regeneration in a mid-elevation,temperate rain forest in Isla de Chiloé, Chile

The regeneration of canopy and subeanopy species in a mid-elevation, primary rain forest in the Coastal Range of Isla de Chiloé (42°30S), in the cold-temperate region of Chile, was studied by comparing seedling and sapling abundances under the forest canopy, and within 36 tree-fall gaps. The forest was dominated byAmomyrtus luma andLaurelia philippiana (33 and 32% of the main canopy individuals), and two subcanopy species (Myrceugenia ovata, andMyrceugenia planipes) were also important. Uncommon species in the canopy wereDrimys winteri, Amomyrtus meli, andRaphithamnus spinosus. Tree-fall gaps were created generally by the fall of several trees, and the main canopy species were the principal gap-makers. Gap sizes varied between 28 and 972 m², with a mean of 197 m². Seedling and sapling abundances indicate that the dominant species are capable of regenerating below the canopy, but they also germinate and show enhanced growth within small light gaps. For one of the common subcanopy species (M. planipes) and the two infrequent canopy species (D. winteri, andA. meli) regeneration seems to depend entirely on tree-fall gaps. Thus, in this forest, light gaps allow the persistence of infrequent canopy species, but seem less important for the regeneration and maintenance of dominant canopy species.     

Shoot growth and tree architecture of saplings of the major canopy dominants in a warm-temperate rainforest

Sapling density, shoot growth, and sapling architecture were studied in five major canopy dominants both under closed canopy and gaps in a warm-temperate rainforest. The five species showed wide variations in distribution, shoot growth, and sapling architecture. Distylium racemosum and Quercus acuta had significantly higher sapling densities under closed canopy than in gaps. Castanopsis sieboldii and Machilus thunbergii had significantly higher sapling densities in gaps than under closed canopy. Quercus salicina showed no significant difference in sapling density between the two habitats. Under closed canopy, C. sieboldii and M. thunbergii had wider crowns than the other species. Distylium racemosum had the greatest number of terminal shoots among the species. Quercus acuta had a branchless small crown. Quercus salicina showed intermediate values in crown width, depth and the number of terminal shoots among the species. Distylium racemosum showed the greatest height-growth rate among the species under the closed canopy, but was the slowest in gaps. Castanopsis sieboldii and M. thunbergii showed the greatest height-growth rates among the species in gaps. Quercus salicina showed the slowest height-growth rates both under closed canopy and gaps. All of the five species showed low mortality under closed canopy. For the major canopy dominants: (i) sapling architecture may not be an important factor in determining mortality but it may be important for height-growth rate; and (ii) sprouting helps saplings to survive until gap formation and facilitates rapid growth in the gaps.     

Stem growth and canopy dynamics in a world-wide range of Fagus forests

Abstract. Forests dominated by Fagus (beech) occur widely in the Northern Hemisphere. Tree species dominant together with beech vary in tolerance of understorey conditions. They are deciduous broad-leaved, evergreen broad-leaved or evergreen coniferous. The frequency and intensity of events that reduce the forest canopy cover are important determinants of the ratio of beech to other species in the canopy. For trees in the understorey and the canopy, stem diameter growth rate is determined by light regime and growing space which in turn are determined by canopy cover. We evaluated increase in stem diameter growth rate as an indicator of sudden reductions in canopy cover and canopy dynamics. We used tree-ring chronologies and calculated an index of growth rate increase (GI) to compare the canopy dynamics of 11 natural beech forests. Per site, the annual average value of GI poorly reflected the effects of dry or cool summers, and it clearly reflected events like tornados and hurricanes that removed substantial canopy cover. Among groups of sites average values of GI were significantly different. In the sites with a lower level of average GI, the establishment of the more shade tolerant tree and shrub species in the understorey was favoured, and subcanopy layers became more dense. On the other hand, higher levels of average GI allowed for more light demanding tree species to reach the canopy.     

Dendrochronological analysis of the canopy history of two Ohio old-growth forests

This study examined the temporal patterns of establishment, suppression, and release of major tree species in two old-growth Ohio forest remnants as a means to determine the past disturbance history of these forests. Increment cores were taken from a total of 154 trees from two well-drained, upland plots and two poorly-drained, bottomland plots in each of the two forested areas. Acer saccharum and Fagus grandifolia exhibited multiple episodes of suppression and release prior to becoming canopy trees, and could tolerate suppressions as long as 84 years. In contrast, Quercus macrocarpa, Q. muehlenbergii, Prunus serotina, and Acer saccharinum rerely exhibited any tolerance to suppression and appeared to have entered the canopy after single disturbances had opened large areas of canopy. There was clear synchrony in the temporal pattern of establishment and final release from suppression among trees from bottomland plots scattered throughout the stands, indicating that relatively large disturbances were important in these poorly-drained areas. In contrast, there was little synchrony among trees from well-drained upland plots, except in a single instance where selective cutting of Quercus trees opened the canopy. Thus, the canopy of upland site was likely subjected only to small disturbances resulting from the death of one or a few trees. At the whole of forest level, there was evidence of episodic recruitment of canopy trees in both forests. Establishment of Fraxinus spp. and Quercus spp. were particularly episodic, and few Fraxinus or Quercus trees alive today established during the last century. These data suggest that large disturbances have affected canopy dynamics of both upland and bottomland areas prior to 1900 and in bottomland forests through this century. In contrast, disturbances in upland areas during this century have been restricted to small, treefall-generated canopy gaps.     

Demographic traits of understory trees and population dynamics of aPicea-Abies forest in Taisetsuzan National Park, northern Japan

The size structure transition matrices ofPicea jezoensis, Picea glehnii andAbies sachalinensis of a sub-boreal forest in Hokkaido, northern Japan were constructed based on the demography of each species (Picea jezoensis andPicea glehnii were dealt with together asPicea) during a 4-year period. Two types of matrices, density-independent and density-dependent population dynamics models, were investigated for evaluating the ‘waiting pattern’ betweenPicea spp. andA. sachalinensis. For the density-dependent model, it was assumed that the demographic traits of understory trees, the recruitment rate, the understory mortality rate and the transition probability from the understory to canopy stages, were regulated by the one-sided competitive effect of canopy trees. The observed size structure ofPicea was almost consistent with the stationary size structure obtained in both the density-independent and the density-dependent models, whereas the observed size structure ofA. sachalinensis was not realized in the two models. The effects of both the transition probability from the understory to canopy stages and the recruitment rate on the dynamics of canopy trees were investigated. ForPicea, two parameters—recruitment rate (e _i ) and transition probability from the understory to canopy stages-exponentially affected the dynamics of canopy trees. In contrast, forAbies sachalinensis, the two parameters affected linearly the dynamics of canopy trees. In conclusion, the population dynamics ofPicea andA. sachalinensis was determined by the parameters of the recruitment rate and the transition probability from the understory to canopy stages, relating to waiting patterns of understory trees for future gap formation. InPicea, the demographic parameters of understory trees intensively regulated the dynamics of canopy trees if compared withA. sachalinensis, suggesting that the performance of understory trees plays a key role in the population dynamics ofPicea. This reflects the growth pattern of understory trees in the regeneration of the two species.