Canopy arthropods community within and among oak species in central Mexico

Quercus rugosa and Q.laurina are species that presents a wide geographical distribution range in temperate forests of Mexico. Oak canopies contain a considerable portion of arthropod diversity and the arthropods fauna fulfill a wide variety of ecological roles. We examined the effect of oak species and seasonal changes on some community structure parameters (diversity, composition, similarity, biomass, rare species, and density of arthropod fauna) of canopy arthropods. In total, 40 oak canopies were fogged during rainy and dry season. A total of 614 identified arthropod morphospecies were recognized belonging to 22 orders associated with tree canopies. A separation of host tree species during both seasons, suggesting a different community structure on host plants species was demonstrated by the principal component analyses (PCA), therefore, differences between oak species results in phenotypes that structure the composition of the arthropod community. Q.laurina registered the highest densities, diversity index and number of rare species in comparison with Q.rugosa. While arthropod biomass showed an inverse pattern. Trees more close to one another (spatial distance) register a more similar canopy arthropod fauna. This study suggests that the trees of Q.laurina could act as a center of biodiversity by the accumulation of arthropod fauna with a considerable number of rare species, which presents wide ecological roles or is involved in critical processes that maintain forest ecosystems[Current Zoology 55(2):132-144,2009].     

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.     

Canopy structure,light microclimate and leaf gas exchange of Quercus coccifera L. in a Portuguese macchia: measurements in different canopy layers and simulations with a canopy model

Summary The structural characteristics of a diverse array of Quercus coccifera canopies were assessed and related to measured and computed light attenuation, proportion of sunlit foliage, foliage temperatures, and photosynthesis and diffusive conductance behavior in different canopy layers. A canopy model incorporating all components of shortwave and longwave radiation, and the energy balance, conductance, and CO₂ and H₂O exchanges of all leaf layers was developed and compared with measurements of microclimate and gas exchange in canopies in four seasons of the year. In the denser canopies with a leaf area index (LAI) greater than 5, there is little sunlit foliage and the diffuse radiation (400–700 nm) is attenuated to 5% or less of the global radiation (400–700 nm) incident on the top of the canopy. Foliage of this species is nonrandomly distributed with respect to azimuth angle, and within each canopy layer, foliage azimuth and inclination angles are correlated. A detailed version of the model which computed radiation interception and photosynthetic light harvesting according to these nonrandom distributions indicated little difference in whole-canopy gas exchange from calculations of the normal model, which assumes random azimuth orientation. The contributions of different leaf layers to canopy gas exchange are not only a function of the canopy microclimate, but also the degree to which leaves in the lower layers of the canopy exhibit more shade-leaf characteristics, such as low photosynthetic and respiratory capacity and maximal conductance. On cloudless days, the majority of the foliage in a canopy of 5.4 LAI is shaded —70%–90% depending on the time of year. Yet, the shaded foliage under these conditions is calculated to contribute only about one-third of the canopy carbon gain. This contribution is about the same as that of the upper 13% of the canopy foliage. Computed annual whole-canopy carbon gain and water use are, respectively, 60% and 100% greater for a canopy of 5 LAI than for one of 2 LAI. Canopy water-use efficiency is correspondingly less for the canopy of 5 LAI than for that of 2 LAI, but most of this difference is apparent during the cool months of the year, when moisture is more abundant.     

Drought acclimation of two deciduous tree species of different layers in a temperate forest canopy

Water-use strategies of Populus tremula and Tilia cordata, and the role of abscisic acid in these strategies, were analysed. P. tremula dominated in the overstorey and T. cordata in the lower layer of the tree canopy of the temperate deciduous forest canopy. Shoot water potential (), bulk-leaf abscisic acid concentration ([ABA]_leaf), abscisic acid concentration in xylem sap ([ABA]_xyl), and rate of stomatal closure following the supply of exogenous ABA (v) decreased acropetally through the whole tree canopy, and foliar water content per area (w), concentration of the leaf osmoticum (c), maximum leaf-specific hydraulic conductance of shoot (L), stomatal conductance (g_s), and the threshold dose per leaf area of the exogenous ABA (d_a) required to reduce stomatal conductance increased acropetally through the tree canopy (from the base of the foliage of T. cordata to the top of the foliage of P. tremula) in non-stressed trees. The threshold dose per leaf dry mass of the exogenous ABA (d_w) required to reduce stomatal conductance, was similar through the tree canopy. After a drought period (3 weeks), the , w, L, g_s, d_a and d_w had decreased, and c and v had increased in both species. Yet, the effect of the drought period was more pronounced on L, g_s, d_a, d_w and v in T. cordata, and on , w and c in P. tremula. It was concluded that the water use of the species of the lower canopy layer—T. cordata, is more conservative than that of the species of the overstorey, P. tremula. [ABA]_leaf had not been significantly changed in these trees, and [ABA]_xyl had increased during the drought period only in P. tremula. The relations between [ABA]_leaf, [ABA]_xyl and the stomatal conductance, the osmotic adjustment and the shoot hydraulic conductance are also discussed.     

泰山4种优势造林树种叶片凋落物分解对凋落物内细菌群落结构的影响

为研究泰山不同造林树种凋落物叶分解对细菌群落的影响。以泰山4种主要优势造林树种刺槐(Robinia pseucdoacacia)、麻栎(Quercus acutissima)、油松(Pinus tabulaeformis)和赤松(Pinus densiflora)为研究对象,采用凋落物分解袋法及Illumina Miseq测序平台对细菌16S rDNA V4—V5区扩增产物进行双端测序,分析了4种树种叶片凋落物分解对细菌群落结构及多样性的影响。结果表明:(1)4种树种叶片分解速率差异显著(P0.05),刺槐分解速率显著高于其他3个树种(P0.05),表现为刺槐赤松油松麻栎。(2)4种叶凋落物分解一年后化学元素含量与初始化学元素相比均存在显著差异。C、木质素含量均显著降低(P0.05);N、P含量显著升高(P0.05)。(3)所有样品一共获得643440条有效序列,分属于35门,92纲,121目,246科,410属,206种。细菌群落NMDSβ-多样性分析显示除油松和赤松间差异较小外,其他树种间差异程度均较大。其中,细菌群落相对丰度在5%以上的优势类群是变形菌门、放线菌门、拟杆菌门、酸杆菌门,且在4种处理之间差异显著(PSymbol|@@0.05)。在纲水平上,α-变形菌纲、β-变形菌纲、不明放线菌纲、鞘脂杆菌纲、γ-变形菌纲、δ-变形菌纲为主要的优势纲,其中不明放线菌纲和鞘脂杆菌纲差异显著(PSymbol|@@0.05)。在种水平上,Bradyrhizobium elkanii和Luteibacter rhizovicinus在4个处理中都为优势种,每个处理也都有自己所特有的优势种。(4)4个处理细菌丰富度(OUT、观测到的物种数和ACE指数)和系统发育多样性(PD指数)之间差异显著(PSymbol|@@0.05),且阔叶树种刺槐和麻栎显著高于针叶树种赤松和油松。(5)叶片凋落物性状和细菌群落NMDS分析表明,细菌群落多样性受到凋落物化学性质的影响,尤其是凋落物初始C/N比和木质素/N比。此外,在细菌群落多样性和叶片凋落物化学性质两个因素中,分解速率与凋落物化学性质相关性更大。研究结果有助于理解细菌群落结构和多样性对森林生态系统叶片凋落物分解的影响。     

Plastic changes of leaf mass per area and leaf nitrogen content in response to canopy openings in saplings of eight deciduous broad-leaved tree species

Leaf nitrogen content per area (N_area) is a good indicator of assimilative capacity of leaves of deciduous broad-leaved trees. This study examined the degrees of increase in N_area in response to canopy openings as leaf mass per area (LMA) and leaf nitrogen content per mass (N_mass) in saplings of eight deciduous broad-leaved tree species in Hokkaido, northern Japan. Five of the species were well-branched species with a large number of small leaves (lateral-growth type), and the other three species were less-branched species with a small number of large leaves (vertical-growth type). The degrees of increase in N_area were compared between the two crown types. In closed-canopy conditions, leaves of the vertical-growth species tended to have a lower LMA and higher N_mass than those of the lateral-growth species, which resulted in similar N_area for both. LMA increased in canopy openings in the eight species, and the degrees of increase were not largely different between the lateral- and vertical-growth species. On the contrary, N_mass was unchanged in canopy openings in the eight species. As a result, N_area of each species increased in canopy openings in proportion to the increase in LMA, and the degrees of increase in N_area were similar in the lateral- and vertical-growth species. Therefore, this study showed that the degrees of increase in N_area were not correlated with the crown architecture (i.e., the lateral- and vertical-growth types).     

Annual photosynthetic activities of temperate evergreen and deciduous broadleaf tree species with simultaneous and successive leaf emergence in response to altitudinal air temperature

To explain why the composition of evergreen and deciduous forests changes along air temperature gradients, we measured several traits of single leaves from temperate deciduous and evergreen broadleaf trees with simultaneous and successive leaf emergence growing at different altitudes in the field. The parameters included seasonal net photosynthetic rate, longevity, mass per area, nitrogen content, and photosynthetic nitrogen-use efficiency. With decreasing altitude, the leaf longevity of deciduous broadleaf trees increased, whereas the maximum net photosynthetic rate decreased. In contrast, leaf longevity of evergreen broadleaf trees decreased, whereas the minimum net photosynthetic rate in winter increased. Along the air temperature gradient, the annual production of deciduous trees with simultaneous leaf emergence may be constant, because the integrated lifetime net photosynthetic rate (ILNPR) of a single leaf changed little. In comparison, deciduous trees with successive leaf emergence may show enhanced annual production with increasing air temperature, by increasing the total leaf number per branch and tree under an extended growing season. Temperate evergreen broadleaf tree species may also show increased annual production with increasing air temperature by sufficiently raising the number of the first-year leaves to the total leaves of branch and tree, which is accelerated by raising the integrated first-year net photosynthetic rate of the single leaf, despite little change in the ILNPR. With increasing air temperature from cool-temperate to warm-temperate zones, evergreen broadleaf tree species gain an advantage of the annual production over deciduous broadleaf tree species with simultaneous leaf emergence.     

Population structure and succession in temperate forests of southwestern Japan

In the temperate forests of southwestern Japan, the population density of woody plants in the community increases in the early stage of secondary succession, reaches a peak in the old oak-chestnut forest, and decreases towards the climax beech forest. The species richness and diversity of woody plants also show a trend similar to that of the population density. The canopy-tree population decreases in the course of the succession while the basal area increases, showing a self-thinning process. The species richness, diversity and population density of herbaceous plants are much influenced by the dominance of the bamboo, Sasa palmata. The life-history traits of trees, lower trees and shrubs are discussed in relation to their shoot system, reproductive pattern and successional processes.     

Litter quality effects of beech and hornbeam on undergrowth species diversity in Luxembourg forests on limestone and decalcified marl

Question: Do beech and hornbeam differ in undergrowth species diversity, and could these differences be explained by mass of the organic layer, pH or soil moisture? Could species richness under beech and hornbeam be explained by differences in N dynamics? Location: Ancient forests dominated by beech and hornbeam in Luxembourg, on decalcified and lime‐rich parent material. Methods: Species composition and soil characteristics, i.e. mass of the organic layer, pH, moisture content, bulk density and air‐filled pore space, were determined under beech and hornbeam on decalcified marl and limestone, and analysed with cluster analysis, correspondence analysis and ANOVA, with tree species and parent material as independent factors. Net N mineralization, nitrification and N in soil and microbial biomass were determined in a laboratory incubation experiment over a pH gradient, and correlated with species richness. Results: Species richness was lower under beech than under hornbeam on both decalcified marl and on limestone. Mass of the organic layer was higher under beech in all cases, but pH and soil moisture only differed between tree species on decalcified marl. Species richness increased with soil N, microbial N and nitrification in the mineral topsoil, but was not correlated with net N mineralization, and was negatively correlated with nitrification in the organic layer and with efficiency of N mineralization per unit microbe. Conclusion: Litter quality affected species richness on both limestone and decalcified marl via the organic layer, but only on decalcified marl in terms of pH or soil moisture. Species richness did not seem to be stimulated by high overall net N mineralization or high nitrate availability in the organic layer, but only by high nitrate in the mineral topsoil.     

Adjustment of foliage structure and function to a canopy light gradient in two co-existing deciduous trees. Variability in leaf inclination angles in relation to petiole morphology

 Foliar inclination angles, petiole morphology and dry matter partitioning between assimilative and support biomass were studied in shade-intolerant Populus tremula L. and shade-tolerant Tilia cordata Mill. along a natural light gradient across the canopy. The leaves of sub-canopy species T. cordata were on average exposed to lower irradiances, and they were also more horizontal with greater blade inclination angles (ϕ_B, defined as the angle between the leaf fall-line and the horizon; ϕ_B was positive for the leaves inclined upwards, and negative for the leaves inclined downwards) than those in P. tremula. Seasonal average daily integrated quantum flux density (Q _int, mol m^–2 day^–1) and ϕ_B were not related in T. cordata, and only a weak negative effect of Q _int on ϕ_B was detected in P. tremula. Nevertheless, when both species were pooled, there was a strong negative relationship between Q _int and ϕ_B, implying that the leaves became progressively vertical with increasing height in the canopy. Interspecific differences in foliage inclination were mainly related to petiole morphology, in particular to petiole length, rather than to contrasting biomass investment patterns between assimilative and support tissues within the leaf. It was suggested that more horizontal leaves, resulting from the species-specific structure of petioles, partly explain the superior performance of shade-tolerant T. cordata in the understory and the sub-canopy. Received: 13 November 1997 / Accepted: 6 March 1998     

Tree species growth response to climate warming varies by forest canopy position in boreal and temperate forests

Reports of forest sensitivity to climate change are based largely on the study of overstory trees, which contribute significantly to forest growth and wood supply. However, juveniles in the understory are also critical to predict future forest dynamics and demographics, but their sensitivity to climate remains less known. In this study, we applied boosted regression tree analysis to compare the sensitivity of understory and overstory trees for the 10 most common tree species in eastern North America using growth information from an unprecedented network of nearly 1.5 million tree records from 20,174 widely distributed, permanent sample plots across Canada and the United States. Fitted models were then used to project the near-term (2041–2070) growth for each canopy and tree species. We observed an overall positive effect of warming on tree growth for both canopies and most species, leading to an average of 7.8%–12.2% projected growth gains with climate change under RCP 4.5 and 8.5. The magnitude of these gains peaked in colder, northern areas for both canopies, while growth declines are projected for overstory trees in warmer, southern regions. Relative to overstory trees, understory tree growth was less positively affected by warming in northern regions, while displaying more positive responses in southern areas, likely driven by the buffering effect of the canopy from warming and climate extremes. Observed differences in climatic sensitivity between canopy positions underscore the importance of accounting for differential growth responses to climate between forest strata in future studies to improve ecological forecasts. Furthermore, latitudinal variation in the differential sensitivity of forest strata to climate reported here may help refine our comprehension of species range shift and changes in suitable habitat under climate change.     

Variations in resistance to canopy disturbances and their interactions with the spatial structure of major species in a cool‐temperate forest

Question: How does typhoon‐related disturbance (more specifically, disturbance in the understorey due to tree‐fall and branch‐fall) affect different species mortality rates in a vertically well‐structured forest community? Location: Cool‐temperate, old‐growth forest in the Daisen Forest Reserve, Japan. Methods: We investigated the canopy dynamics and mortality rate trends of trees ≥5 cm diameter at breast height in a 4‐ha study plot, and analysed the effects of tree diameter and spatial structure on the mortality risks for major tree species in the understorey. Results: Significant differences were found in the mortality rates and proportions of injured dead stems between census periods, which were more pronounced in the understorey than in the canopy. Acer micranthum, which showed increased mortality during typhoon disturbance periods, had a clumped distribution. In contrast, Acer japonicum and Viburnum furcatum, which showed similar mortality rates between census periods, had a loosely clumped spatial distribution and a negative association with canopy trees, respectively. In the understorey stems of Acanthopanax sciadophylloides and Fagus crenata, whose spatial distribution patterns depended on canopy gaps, significant increases in mortality rates were observed only during severe typhoon‐related disturbance periods. Conclusions: The sensitivity of trees to typhoon‐related canopy disturbance is more pronounced in the lower layers of vertically structured forest communities. Differences in mortality patterns generated through the combined effects of spatial variation in disturbance regime and species‐specific spatial distribution patterns (spatial aggregation, association with canopy trees, and canopy gap dependency) contribute to the co‐existence of understorey species in forest communities that are subject to typhoon‐related disturbance.     

秦岭皇冠暖温带落叶阔叶林灌木层结构与物种多样性

灌木是森林生态系统中的重要组成部分,探究灌木层的物种组成、结构和空间分布可以为进一步阐明森林物种共存、生物多样性维持和群落演替机制奠定基础。本研究分析了秦岭皇冠25 hm²森林样地灌木层的物种组成、空间分布以及物种多样性。结果表明: 样地内灌木独立个体共20716株(分枝10463个),隶属于28科45属54种。灌木层优势种的优势度不明显,重要值均＜10。灌木层植被的径级结构为倒“J”型;灌木在样地内呈聚集分布且空间分布存在明显的海拔差异。样地内个体数最多的9种灌木的径级结构和空间分布与灌木层整体情况一致。随着海拔升高,Shannon多样性指数(H)和Simpson优势度指数(D)变化不显著,而Pielou均匀度指数(E)降低。多样性指数H、D、E随乔木层优势种个数增加而降低,E随亚乔木层优势种个数增加而升高。多样性指数H、D、E与土壤全磷呈显著负相关;H与土壤pH呈显著正相关,E与土壤全钾呈显著正相关。本研究区灌木层物种丰富,结构稳定,更新良好;灌木层的物种多样性主要受到乔木层优势种个数和土壤pH、全磷的影响。     

Fire frequency and tree canopy structure influence plant species diversity in a forest-grassland ecotone

Disturbances and environmental heterogeneity are two factors thought to influence plant species diversity, but their effects are still poorly understood in many ecosystems. We surveyed understory vegetation and measured tree canopy cover on permanent plots spanning an experimental fire frequency gradient to test fire frequency and tree canopy effects on plant species richness and community heterogeneity within a mosaic of grassland, oak savanna, oak woodland, and forest communities. Species richness was assessed for all vascular plant species and for three plant functional groups: grasses, forbs, and woody plants. Understory species richness and community heterogeneity were maximized at biennial fire frequencies, consistent with predictions of the intermediate disturbance hypothesis. However, overstory tree species richness was highest in unburned units and declined with increasing fire frequency. Maximum species richness was observed in unburned units for woody species, with biennial fires for forbs, and with near-annual fires for grasses. Savannas and woodlands with intermediate and spatially variable tree canopy cover had greater species richness and community heterogeneity than old-field grasslands or closed-canopy forests. Functional group species richness was positively correlated with functional group cover. Our results suggest that annual to biennial fire frequencies prevent shrubs and trees from competitively excluding grasses and prairie forbs, while spatially variable shading from overstory trees reduces grass dominance and provides a wider range of habitat conditions. Hence, high species richness in savannas is due to both high sample point species richness and high community heterogeneity among sample points, which are maintained by intermediate fire frequencies and variable tree canopy cover.     

中国温带和亚热带8个树种叶和吸收根协同分解

叶和细根(2mm)是森林生态系统的分解主体,二者是否协同分解,将极大影响所属植物在生态系统碳(C)循环中的物种效应。已有研究显示,叶和细根的分解关系具有极大的不确定性,认为很大程度上归因于细根内部具有高度的异质性,导致叶和细根在功能上不相似。为此,使用末梢1级根和细根根枝作为研究对象,它们在功能上同叶类似,称为吸收根。通过分解包法,分别在黑龙江帽儿山和广东鹤山,研究了2个阔叶树种和2个针叶树种(共8个树种)的叶和吸收根持续2a多的分解。结果发现,分解速率k(a~(-1),负指数模型)在8个树种整体分析时具有正相关关系(P0.05),在相同气候带或植物生活型水平上是否相关,受叶的分解环境及吸收根类型的影响;N剩余量整体上并不相关,亚热带树种的叶和细根根枝的N剩余量在分解1a后高度显著正相关,温带树种的叶和1级根的N剩余量在分解2a后显著高度正相关。本研究中,根-叶分解过程是否受控于相同或相关的凋落物性质是决定根-叶分解是否相关的重要原因,其中分解速率与酸溶组分正相关、与酸不溶组分负相关。比较已有研究,总结发现,根-叶分解关系受物种影响较大,暗示气候变化导致物种组成的改变将极大影响地上-地下关系,也因此影响生态系统C循环。