Formation and closure of canopy gaps in the rain forest at Nouragues, French Guiana

Formation and closure of canopy gaps was monitored for three years in 12 ha of primary rain forest at Nouragues, French Guiana. At the first inventory, in April 1991, 74 openings in the canopy > 4 m² (sensu Brokaw 1982a) were located; 60 of these gaps were formed before January 1990. Between January 1990 and December 1993, 5 to 15 gaps were annually formed, opening 0.64–1.33% of the forest canopy each year. Of all gaps, 41% were created by a falling, snapped tree, 34% by a falling, uprooted tree, 22% by a falling branch, and 3% by a falling dead stem. A refined nearest neighbour analysis showed that gaps formed after January 1990 were clustered: uprooting of trees seemed to be related to shallow soils, and relatively many other trees fell when a tree uprooted, independent of the dbh of the uprooted tree. In 37 gaps, canopy openness in the gap centre (determined by hemispherical photographs) was monitored over three years. In 54% of the gaps, canopy openness increased in two successive years. It is reasoned that edges of especially large gaps may frequently be re-disturbed by falling trees or branches. Results suggest that gaps have closed after around 15 years. More data are needed to verify this.     

林窗几何特征的测定方法

林窗面积、形状及边界木高是决定林窗环境异质性的3个林窗几何特征,影响林窗内植物更新。林窗几何特征的快速测量方法是林窗研究的基础,测量方法可分为2类:基于地面实际测量的地面法和基于林窗林冠照片的相片法。地面法费时费力,受人为因素影响大,可测量林冠林窗和扩展林窗的面积,但不能测量林窗形状和边界木高。相片法具有简单、客观、可重复的优点,但仅适用于林冠林窗。相片法共有5种:"平面相片法"、"航片法"、"半球面影像法"、"双半球面影像法"和"改进的半球面影像法"。前3种测量方法只能测量林冠林窗面积;"改进的半球面影像法"可测量林冠林窗面积和形状,且精度高于前3种相片法,但所需参数最多;"双半球面影像法"可测量林窗面积、形状及边界木高这3个林窗几何特征,且精度较高,但拍摄要求较高。     

Crown development in tropical rain forest trees in gaps and understorey

Emergence and establishment of Tsuga sieboldii seedlings were monitored for six years in an old-growth mixed forest in Japan, to clarify the reason why adult T. sieboldii trees are distributed only on convex slopes. T. sieboldii seedlings germinated on exposed soils without broad-leaved litter cover on both convex and concave slopes, but they died out from concave slope during the first two years after germination. Along a gradient from convex to concave slope, litter cover became higher and surface soil erosion became severer, and T. sieboldii seedlings died at the plots with severe surface soil erosion. Other environmental variables, such as relative illuminance, vegetation cover of seedling layer and soil water potential were not significantly different between convex and concave slopes. We concluded that the restricted seedling establishment of T. sieboldii to convex slope is determined by stability of surface soils and broad-leaved litter cover.     

Size distribution and expansion of canopy gaps in a northern Appalachian spruce-fir forest

Canopy gap area/age distributions and growth mechanisms were examined in a virgin subalpine forest in the White Mountains, New Hampshire, USA. The gap area distribution was negative exponential in form. Whithin gap tree ages varied widely in response to stepwise gap expansion caused by windthrow of peripheral trees or death of standing mature Picea rubens at gap edges. As a consequence, the density of small gaps may have been underestimated and the density of large gaps overestimated. The estimates of canopy turnover time, 303 yr, and of patch birth rate on an area basis, 3.3×10^-3 ha new patches/ha land area/yr, were not affected by the gap expansion phenomenon. However, any estimate of patch birth rate as numbers of new patches formed per year would have been too low. Because of increasingly widespread Picea death, the patch area/age distribution of this forest may not currently be in steady-state.     

The role of canopy gaps in the regeneration of coastal dune forest

In regenerating coastal dune forest, the canopy consists almost exclusively of a single species, Acacia karroo. When these trees die, they create large canopy gaps. If this promotes the persistence of pioneer species to the detriment of other forest species, then the end goal of a restored coastal dune forest may be unobtainable. We wished to ascertain whether tree species composition and richness differed significantly between canopy gaps and intact canopy, and across a gradient of gap sizes. In three known‐age regenerating coastal dune forest sites, we measured 146 gaps, the species responsible for gap creation, the species most likely to reach the canopy and the composition of adults, seedlings and saplings. We paired each gap with an adjacent plot of the same area that was entirely under intact canopy and sampled in the same way. Most species (15 of 23) had higher abundance in canopy gaps. The probability of self‐replacement was low for A. karroo even in the largest gaps. Despite this predominance of shade‐intolerant species, regenerating dune forest appears to be in the first phase of succession with ‘forest pioneers’ replacing the dominant canopy species. The nature of these species should lead to successful regeneration of dune forest.     

Seasonal diets of insectivorous birds using canopy gaps in a bottomland forest

ABSTRACT.   Little is known about how insectivorous bird diets are influenced by arthropod availability and about how these relationships vary seasonally. We captured birds in forest-canopy gaps and adjacent mature forest during 2001 and 2002 at the Savannah River Site in Barnwell County, South Carolina, and flushed their crops to gather information about arthropods eaten during four periods: spring migration, breeding, postbreeding, and fall migration. Arthropod availability for foliage- and ground-gleaning birds was examined by leaf clipping and pitfall trapping. Coleopterans and Hemipterans were used by foliage- and ground-gleaners more than expected during all periods, whereas arthropods in the orders Araneae and Hymenoptera were used as, or less than, expected based on availability during all periods. Ground-gleaning birds used Homopterans and Lepidopterans in proportions higher than availability during all periods. Arthropod use by birds was consistent from spring through fall migration, with no apparent seasonal shift in diet. Based on concurrent studies, heavily used orders of arthropods were equally abundant or slightly less abundant in canopy gaps than in the surrounding mature forest, but bird species were most frequently detected in gaps. Such results suggest that preferential feeding on arthropods by foliage-gleaning birds in gap habitats reduced arthropod densities or, alternatively, that bird use of gap and forest habitat was not determined by food resources. The abundance of arthropods across the stand may have allowed birds to remain in the densely vegetated gaps where thick cover provides protection from predators.     

Factors influencing sapling composition in canopy gaps of a temperate deciduous forest

To detect the factors that affect sapling species composition in gaps, we investigated 55 gaps in an old-growth temperate deciduous forest in Ogawa Forest Reserve, central Japan. Gap size, gap age, gap maker species, topographic location, adult tree composition around gaps, and saplings of tree species growing in the gaps were censused. For gaps 5 m², mean gap size was 70 m² and the maximum was 330 m². Estimated ages of gaps had a tendency to be concentrated in particular periods relating to strong wind records in the past. The sapling composition in gaps was highly and significantly correlated to that under closed canopy, indicating the importance of advance regeneration in this forest. However, some species showed significant occurrence biases in gaps or under closed canopy, suggesting differences in shade tolerance. The result of MANOVA showed that gap size and topography were important factors in determining the sapling composition in gaps. Species of gap makers affected the sapling composition indirectly by influencing gap size. The existence of parent trees around gaps had effects on sapling densities of several species. Gap age did not have clear influences on sapling composition. Variations in gap size and topography were considered as important factors that contribute to maintenance of species diversity in this forest.     

Community dynamics of evergreen broadleaf forests in southwestern Japan. I. Wind damaged trees and canopy gaps in an evergreen oak forest

The rates of treefall and canopy opening in the evergreen oak forest in southwestern Japan were determined by studying the number and size distribution of overstory trees, wind damaged trees, and canopy gaps in a belt transect in the Kasugayama Forest Reserve in Nara City. Thirty three percent of the overstory trees wereCastanopsis cuspidata. The total area of canopy gaps was about 20% of the total land area in the study area. The ages of the gaps were determined by counting the annual rings of various kinds of trees growing in gaps. By comparing gap ages with meteorological data, it became evident that gap formation was mainly caused by strong typhoons. The mean time interval between strong typhoons visiting the forest reserve, 6.57 years, was determined by applying the MNY method to the meteorological data. The treefall rate and the mean area of canopy openings per year were 0.84 overstory trees/ha·year and 55.6 m²/ha·year, respectively. The mean residence time of the forest canopy was about 180 years.     

Fungicides for forest trees,shade trees and forest products

The Botanical Review -     

Leaf dynamics of seedlings of rain forest species in relation to canopy gaps

Summary Leaf dynamics of eight tropical rain forest species seedlings was studied in three environments: the shaded forest understorey, a small gap of ±50 m², and a large gap of ±500 m². Leaf production rate and leaf loss rate were enhanced in gaps, and a large gap resulted in larger increases than a small gap. For most species net leaf gain rate was larger in gaps, although this rate was not always largest in the large gap. Leaf loss decreased, and leaf survival percentages increased with increasing shade tolerance of species, indicating a slower leaf turnover for more shade tolerant species. Leaf area growth rate was only partly determined by net leaf gain rate. Ontogenetic effects on leaf size were also important, especially in the large gap. Species which possessed leaves with high specific leaf weight (SLW) showed lower leaf loss rates and higher leaf survival percentages than species with low SLW leaves. Leaf life span seemed to be related to leafcost per unit area. The relation of specific patterns in leaf production and leaf loss to the regeneration mode of the species is briefly discussed.     

Logging leaves a fingerprint on the number,size, spatial configuration and geometry of tropical forest canopy gaps

Recent advances in remote sensing such as airborne laser scanning have revolutionized our ability to accurately map forest canopy gaps, with huge implications for tracking forest dynamics at scale. However, few studies have explored how canopy gaps vary among forests at different successional stages following disturbances, such as those caused by logging. Moreover, most studies have focused exclusively on the size distribution of gaps, ignoring other key features such as their spatial distribution and shape. Here, we test a series of hypotheses about how the number, size, spatial configuration, and geometry of gaps vary across a logging disturbance gradient in Malaysian Borneo. As predicted, we found that recently logged forests had much higher gap fraction compared to old-growth forests, a result of having both a greater total number of gaps and a higher proportion of large gaps. Regrowing forests, on the other hand, fell at the opposite end of the spectrum, being characterized by both fewer and smaller gaps compared to nearby old-growth forests. Across all successional stages gaps were found to be spatially clustered. However, logging significantly diluted the degree of spatial aggregation and led to the formation of gaps with much more complex geometries. Our results showcase how logging and subsequent regrowth substantially alter not just the number and size of gaps in a forest, but also their spatial arrangement and shape. Linking these emergent patterns to their underlying processes is key to better understanding the impacts of human disturbance on the structure and function of tropical forests.     

Tree species distribution in canopy gaps and mature forest in an area of cloud forest of the Ibitipoca Range,south-eastern Brazil

The tree community of both canopy gaps and mature forest was surveyed in a 5 ha plot of cloud forest in the Ibitipoca Range, south-eastern Brazil, aiming at: (a) comparing the tree community structure of canopy gaps with that of three strata of the mature forest, and (b) relating the tree community structure of canopy gaps with environmental and biotic variables. All saplings of canopy trees with 1–5 m of height established in 31 canopy gaps found within the plot were identified and measured. Mature forest trees with dbh 3 cm were sampled in four 40×40 quadrats laid on the four soil sites recognised in the local soil catena. All surveyed trees were identified, measured and distributed into three forest strata: understorey (<5 m of height), sub-canopy (5.1–15 m) and canopy (15.1–30 m). The following variables were obtained for each gap: mode of formation, age, soil site, slope grade, size, canopy openness and abundance of bamboos and lianas. A detrended correspondence analysis indicated that the tree community structure of gaps in all soil sites was more similar to that of the mature forest understorey, suggesting that the bank of immatures plays an important role in rebuilding the forest canopy and that gap phases may be important for understorey shade-tolerant species. There was evidence of gap-dependence for establishment for only one canopy tree species. Both canonical correspondence analysis and correlation analysis demonstrated for a number of tree species that the distribution of their saplings in canopy gaps was significantly correlated with two variables: soil site and canopy openness. The future forest structure at each gap is probably highly influenced by both the present structure of the adjacent mature forest and the gap creation event.     

The effect of canopy gaps on growth and morphology of seedlings of rain forest species

Summary Growth and morphology of seedlings of ten tropical rain forest species were studied at Los Tuxtlas, Mexico. Seedlings were grown in three environmental conditions: the shaded forest understorey (FU, receiving 0.9–2.3% of the daily photosynthetic photon flux, PF, above the canopy), a small canopy gap of approx. 50 m² (SG, receiving 2.1–6.1% of daily PF), and a large canopy gap of approx. 500 m² (LG, receiving 38.6–53.4% of daily PF). The growth of all species was enhanced in gaps, and in LG the effect was stronger than in SG. Plants grown in LG had a sunplant morphology, with a high root-shoot ratio (R/S), a high specific leaf weight (SLW) and a low leaf area ratio (LAR). Plants grown in SG or FU showed a shade-plant morphology, with a low R/S, a low SLW and a high LAR. Growth responses varied from species unable to grow in the shade but with strong growth in the sun, to species with relatively high growth rates in both shade and sun conditions. Shade tolerant species were able to grow in the shade because of a relatively high unit leaf rate. The pioneerCecropia had a high growth rate in LG because of a high LAR. Most species showed a complex growth response in which they resembled the shade intolerant extreme in some aspects of the response, and the shade tolerant extreme in other aspects.     

Spatial and temporal dynamics of forest canopy gaps following selective logging in the eastern Amazon

Selective logging is a dominant form of land use in the Amazon basin and throughout the humid tropics, yet little is known about the spatial variability of forest canopy gap formation and closure following timber harvests. We established chronosequences of large‐area (14–158 ha) selective logging sites spanning a 3.5‐year period of forest regeneration and two distinct harvest methods: conventional logging (CL) and reduced‐impact logging (RIL). Our goals were to: (1) determine the spatial characteristics of canopy gap fraction immediately following selective logging in the eastern Amazon; (2) determine the degree and rate of canopy closure in early years following harvest among the major landscape features associated with logging – tree falls, roads, skid trails and log decks; and (3) quantify spatial and temporal differences in canopy opening and closure in high‐ and low‐damage harvests (CL vs. RIL). Across a wide range of harvest intensities (2.6–6.4 felled trees ha^?1), the majority of ground damage occurred as skid trails (4–12%), whereas log decks and roads were only a small contributor to the total ground damage (<2%). Despite similar timber harvest intensities, CL resulted in more ground damage than RIL. Neither the number of log decks nor their individual or total area was correlated with the number of trees removed or intensity of tree harvesting (trees ha^?1). The area of skids was well correlated with the ground area damaged (m²) per tree felled. In recently logged forest (0.5 years postharvest), gap fractions were highest in log decks (mean RIL=0.83, CL=0.99) and lowest in tree‐fall areas (RIL: 0.26, CL: 0.41). However, the small surface area of log decks made their contribution to the total area‐integrated forest gap fraction minor. In contrast, tree falls accounted for more than two‐thirds of the area disturbed, but the canopy gaps associated with felled trees were much smaller than for log decks, roads and skids. Canopy openings decreased in size with distance from each felled tree crown. At 0.5 years postharvest, the area initially affected by the felling of each tree was approximately 100 m in radius for CL and 50 m for RIL. Initial decreases in gap fraction during the first 1.5 years of regrowth diminished in subsequent years. Throughout the 3.5‐year period of forest recovery, tree‐fall gap fractions remained higher in CL than in RIL treatments, but canopy gap closure rates were higher in CL than in RIL areas. During the observed recovery period, the canopy gap area affected by harvesting decreased in radius around each felled tree from 100 to 40 m in CL, and from 50 to 10 m in RIL. The results suggest that the full spatial and temporal dynamics of canopy gap fraction must be understood and monitored to predict the effects of selective logging on regional energy balance and climate regimes, biogeochemical processes including carbon cycling, and plant and faunal population dynamics. This paper also shows that remote sensing of log decks alone will not provide an accurate assessment of total forest area impacted by selective logging, nor will it be closely correlated to damage levels and canopy gap closure rates.     

格氏栲天然林林窗植物群落功能性状的变异

林窗是森林更新演替的重要环节, 揭示林窗环境下功能性状变异来源及其相对贡献, 有助于阐明植物对林窗环境的响应。该研究以中亚热带格氏栲(Castanopsis kawakamii)天然林为对象, 设置9个不同大小的林窗样地, 运用方差分解探讨林窗、物种和个体对叶性状变异的相对贡献, 采用线性回归分析不同大小林窗下群落性状变化及种间和种内性状变异的重要性。研究发现: (1)格氏栲天然林林窗植物比叶面积、叶干物质含量、叶厚和叶绿素含量由种间性状变异主导, 叶氮含量由种内性状变异主导, 叶磷含量受林窗大小影响最大。(2)群落叶磷含量与林窗大小具有显著正相关关系, 土壤温度和水解氮含量对群落叶磷含量具有显著正效应, 土壤有效磷含量具有显著负效应。(3)沿林冠开放度的群落叶磷含量变化主要由种内性状变异引起, 优势种扮演着重要角色。结果表明, 格氏栲天然林林窗环境下植物功能性状仍以种间性状变异为主(平均41%), 但沿林窗环境梯度的群落性状变化主要源自种内性状变异, 通过植物表型可塑性响应环境改变, 优势种作用明显。     

Tree replacement in small canopy gaps of a Tsuga canadensis forest in the Southern Appalachians,Tennessee

Summary Tree species replacement was studied in 95 canopy gaps created by the fall of single trees in an undisturbed, old-growth forest in Great Smoky Mountains National Park, Tennessee. When large trees (dbh>70 cm) of the very shade tolerant species, Tsuga canadensis, die and fall, they are usually replaced by less tolerant species such as Betula alleghaniensis, Liriodendron tulipifera, and Magnolia fraseri. Species diversity of the replacement trees, measured by the index, 1/p_i ², was 5.77 compared to a diversity of 1.66 for the fallen trees.     

Whole-tree water transport scales with sapwood capacitance in tropical forest canopy trees

The present study examines the manner in which several whole‐tree water transport properties scale with species‐specific variation in sapwood water storage capacity. The hypothesis that constraints on relationships between sapwood capacitance and other water relations characteristics lead to predictable scaling relationships between intrinsic capacitance and whole‐tree behaviour was investigated. Samples of sapwood from four tropical forest canopy tree species selected to represent a range of wood density, tree size and architecture, and taxonomic diversity were used to generate moisture release curves in thermocouple psychrometer chambers, from which species‐specific values of sapwood capacitance were calculated. Sapwood capacitance was then used to scale several whole‐tree water transport properties determined from measurements of upper branch and basal sap flow, branch water potential, and axial and radial movement of deuterated water (D₂O) injected into the base of the trunk as a tracer. Sapwood capacitance ranged from 83 to 416 kg m^?3 MPa^?1 among the four species studied and was strongly correlated with minimum branch water potential, soil‐to‐branch hydraulic conductance, daily utilization of stored water, and axial and radial movement of D₂O. The species‐independent scaling of several whole‐tree water transport properties with sapwood capacitance indicated that substantial convergence in plant function at multiple levels of biological organization was revealed by a simple variable related to a biophysical property of water transport tissue.     

Stem water storage and diurnal patterns of water use in tropical forest canopy trees

Stem water storage capacity and diurnal patterns of water use were studied in five canopy trees of a seasonal tropical forest in Panama. Sap flow was measured simultaneously at the top and at the base of each tree using constant energy input thermal probes inserted in the sapwood. The daily stem storage capacity was calculated by comparing the diurnal patterns of basal and crown sap flow. The amount of water withdrawn from storage and subsequently replaced daily ranged from 4 kg d^–1 in a 0·20-m-diameter individual of Cecropia longipes to 54 kg d^–1 in a 1·02-m-diameter individual of Anacardium excelsum, representing 9–15% of the total daily water loss, respectively. Ficus insipida, Luehea seemannii and Spondias mombin had intermediate diurnal water storage capacities. Trees with greater storage capacity maintained maximum rates of transpiration for a substantially longer fraction of the day than trees with smaller water storage capacity. All five trees conformed to a common linear relationship between diurnal storage capacity and basal sapwood area, suggesting that this relationship was species-independent and size-specific for trees at the study site. According to this relationship there was an increment of 10 kg of diurnal water storage capacity for every 0·1 m² increase in basal sapwood area. The diurnal withdrawal of water from, and refill of, internal stores was a dynamic process, tightly coupled to fluctuations in environmental conditions. The variations in basal and crown sap flow were more synchronized after 1100 h when internal reserves were mostly depleted. Stem water storage may partially compensate for increases in axial hydraulic resistance with tree size and thus play an important role in regulating the water status of leaves exposed to the large diurnal variations in evaporative demand that occur in the upper canopy of seasonal lowland tropical forests.