Canopy disturbance and gap partitioning promote the persistence of a pioneer tree population in a near‐climax temperate forest of the Qinling Mountains,China

An unresolved question of temperate forests is how pioneer tree species persist in mature forests. In order to understand the responsible mechanisms, we investigated a near‐climax mixed temperate forest dominated by Betula albosinensis in the Qinling Mountains of China. Through establishing four 50 m × 50 m plots, we examined the canopy disturbance characteristics and its effects on tree recruitments. We further test the intra‐ and interspecific effects on the recruitment of B. albosinensis. The obtained data demonstrated canopy disturbance was frequent but most small‐sized. The canopy gaps are caused mainly by adult B. albosinensis by snapping. The regeneration of coexistent tree species shows a distinct preference for gap size. B. albosinensis were clumped at the juvenile stage and small scales. B. albosinensis juveniles were positively associated with B. utilis juveniles and negatively associated with the conspecific and B. utilis large trees. In addition, B. albosinensis juveniles showed negative associations with contemporary other tree species. Our results suggested that canopy disturbance caused by canopy trees and gap partitioning among the coexistent tree species are important for the persistence of the mixed forest. As a main gapmaker, B. albosinensis appear to develop a self‐perpetuating life‐history trait and allow them to persist.     

不同生境栓皮栎天然更新幼苗植冠构型分析

栓皮栎存在于秦岭南坡的多种林分中,生活在不同生境中的个体往往形成不同的树冠形态和构型特征。为了说明不同生境条件下栓皮栎幼苗的植冠构型变化,采用典型抽样法,对秦岭南坡3种生境中(林冠下、林隙、林缘)的栓皮栎天然更新幼苗的侧枝、叶片特征及其空间分布进行了调查分析,结果表明:3种不同生境中栓皮栎幼苗植冠形态发生了明显的可塑性变化,(1)林冠下的幼苗明显为开阔型树冠,林隙和林缘处的幼苗树冠相对紧密;(2)幼苗的1级侧枝密度与分枝角度在3种生境下均差异显著(P0.05);从Ⅰ到Ⅳ层,林冠下幼苗的分枝角度在冠层内变化幅度不到5°,而林缘处幼苗的分枝角度变化高达40°;发生5个以上1级侧枝的概率以林冠下最大,为0.6;(3)从林缘、林隙到林冠下,幼苗的叶长、叶宽、单叶面积、叶面积指数逐渐降低,数量叶密度和比叶面积则逐渐增大,与其它两种生境相比,林冠下幼苗的叶片逐渐向树冠上层集中,且以更高序的侧枝为主要着生枝条;(4)林隙中栓皮栎幼苗的树高、地径明显优于林缘和林冠下,缩短了苗木进入主林层的时间,林隙对栓皮栎种群更新有利。在今后栓皮栎林的经营中,可以通过适当间伐来增加林隙数量,为森林更新和结构的优化的提供有利条件。     

Cultivation of Non‐timber Forest Products Alters Understory Light Availability in a Humid Tropical Forest in Mexico1

The planting of non‐timber forest products (NTFPs) in the understory of tropical forests is promoted in many regions as a strategy to conserve forested lands and meet the economic needs of rural communities. While the forest canopy is left intact in most understory plantations, much of the midstory and understory vegetation is removed in order to increase light availability for cultivated species. We assessed the extent to which the removal of vegetation in understory plantations of Chamaedorea hooperiana Hodel (Arecaceae) alters understory light conditions. We also examined how any changes in light availability may be reflected by changes in the composition of canopy tree seedlings regenerating in understory plantations. We employed a blocked design consisting of four C. hooperiana plantation sites; each site was paired with an adjacent, unmanaged forest site. Hemispherical canopy photographs were taken and canopy tree seedlings were identified and measured within 12 3 × 2 m randomly placed plots in each site for a total of 96 plots (4 blocks × 2 sites × 12 plots). Plantation management did not affect canopy openness or direct light availability but understory plantations had a higher frequency of plots with greater total and diffuse light availability than unmanaged forest. Comparisons of canopy tree seedling composition between understory plantations and unmanaged forest sites were less conclusive but suggest that management practices have the potential to increase the proportion of shade‐intolerant species of tree seedlings establishing in plantations. Given the importance of advanced regeneration in gap‐phase forest dynamics, these changes may have implications for future patterns of succession in the areas of forest where NTFPs are cultivated.     

Chance versus determinism in canopy gap regeneration in coastal scarp forest in South Africa

Question: Is tree regeneration in canopy gaps characterized by chance or predictable establishment. Location: Coastal scarp forests, Umzimvubu district, Eastern Cape Province, South Africa. Methods: Estimation of richness of gap‐filling species across canopy gaps of different size. Data are compared with regeneration under the canopy. Probability of self‐replacement of gap forming species is calculated. Results: Forest area under natural gap phase was 7.8%, caused mostly by windthrow (54%). The abundance and average size of gaps (87.8 m²) suggests that species diversity may be maintained by gap dynamics. However, only four of 53 gap‐filler species displayed gap size specialization and these were pioneer species. An additional 13 species were more common in larger gaps but there was no gradient in composition of gap‐filler species across gap size (p= 0.61). Probabilities of self‐replacement in a gap were low (< 0.3) and common canopy species were equally abundant in gaps and the understorey. Species composition in gaps showed no pattern of variation, i.e. was unpredictable, which suggests absence of a successional sequence within tree‐fall gaps. There was also only a slight increase in species richness in gaps at intermediate levels of disturbance. Conclusions: Coastal scarp forest appears not to comprise tightly co‐evolved, niche‐differentiated tree species. Unpredictable species composition in gaps may be a chance effect of recruitment limitation of species from the species pool. Chance establishment slows competitive exclusion and may maintain tree diversity in these forests. These data suggest that current levels (≤ 3 gaps per ha) of selective tree harvesting may not cause a reduction in species richness in this forest.     

Gap-phase dynamics and coexistence of a long-lived pioneer and shade-tolerant tree species in the canopy of an old-growth coastal temperate rain forest of Chiloé Island, Chile

Aim A major question with regard to the ecology of temperate rain forests in south‐central Chile is how pioneer and shade‐tolerant tree species coexist in old‐growth forests. We explored the correspondence between tree regeneration dynamics and life‐history traits to explain the coexistence of these two functional types in stands apparently representing a non‐equilibrium mixture. Location This study was conducted in northern Chiloé Island, Chile (41.6° S, 73.9° W) in a temperate coastal rain forest with no evidence of stand disruption by human impact. Methods We assessed stand structure by sampling all stems within two 50 × 20 m and four 5 × 100 m plots. A 600‐m long transect, with 20 uniformly spaced sampling points, was used to quantify seedling and sapling densities, obtain increment cores, and randomly select 10 tree‐fall gaps. We used tree‐ring analysis to assess establishment periods and to relate the influences of disturbances to the regeneration dynamics of the main canopy species. Results Canopy emergent tree species were the long‐lived pioneer Eucryphia cordifolia and the shade‐tolerant Aextoxicon punctatum. Shade‐tolerant species such as Laureliopsis philippiana and several species of Myrtaceae occupied the main canopy. The stem diameter distribution for E. cordifolia was distinctly unimodal, while for A. punctatum it was multi‐modal, with all age classes represented. Myrtaceae accounted for most of the small trees. Most tree seedlings and saplings occurred beneath canopy gaps. Based on tree‐ring counts, the largest individuals of A. punctatum and E. cordifolia had minimum ages estimated to be > 350 years and > 286 years, respectively. Shade‐tolerant Myrtaceae species and L. philippiana had shorter life spans (< 200 years). Most growth releases, regardless of tree species, were moderate and have occurred continuously since 1750. Main conclusions We suggest that this coastal forest has remained largely free of stand‐disrupting disturbances for at least 450 years, without substantial changes in canopy composition. Release patterns are consistent with this hypothesis and suggest that the disturbance regime is dominated by individual tree‐fall gaps, with sporadic multiple tree falls. Long life spans, maximum height and differences in shade tolerance provide a basis for understanding the long‐term coexistence of pioneer and shade‐tolerant tree species in this coastal, old‐growth rain forest, despite the rarity of major disturbances.     

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.     

Short‐term Dynamics and the Effects of Biotic and Abiotic Factors on Plant Physiognomic Groups in a Hurricane‐impacted Lower Montane Tropical Forest

We assessed the short‐term effects of biotic (density, plant size) and abiotic factors (light), on the dynamics of physiognomically different plant groups (palms, tree ferns, lianas, and trees) in a hurricane‐impacted tropical wet montane forest, John Crow Mountains, Jamaica. All plants ≥2 cm (dbh) found within 45, 25 × 25 m permanent sample plots (2.8125 ha), established according to a randomized block design along an elevation gradient, were tagged and measured (dbh) in 2006 and re‐assessed in 2012 after Hurricane Dean (2007). Hemispheric light was measured in 2007 and 2008. Tree and liana size class distributions changed due to high mortality in the smallest size classes and their densities declined; however, palm and tree fern density remained unchanged. The dynamics of trees were only related to tree fern and liana dynamics (e.g., tree mortality was negatively related to liana recruitment etc.). Although pre‐ and posthurricane light was related to palm density and the density of the other plant groups, respectively, there were no significant changes in light. Tree survivorship increased with increasing dbh while posthurricane light and overall density influenced the growth and survivorship of tree species. Species importance value did not change, suggesting that direct regeneration may be the model of forest recovery following this small‐scale disturbance. Over the short term, tree species showed life history trade‐offs that aid species coexistence after this moderate/low disturbance event. Our study highlights that hurricanes with low impacts can have differential short‐ and possibly long‐term effects on different plant groups.     

Small gap dynamics in the southern boreal forest of eastern Canada: Do canopy gaps influence stand development?

Question: To what extent do small‐scale disturbances in the forest canopy, created by natural disturbance agents, affect stand development? Doubts exist as to whether small canopy openings have any real effect on the understory tree recruitment, especially in boreal forests. Location: Conifer and mixed stands in the Gaspesian region in eastern Québec. The main natural disturbance agents are recurring outbreaks of Choristoneura fumiferana (eastern spruce budworm) and winds. Methods: Linear transects in 27 sites were used to describe the gap (< 0.1 ha) regime parameters, including gap fraction, gap size and change in disturbance severity through time. Three stand types were distinguished, based on a gradient of abundance of tree host species for the eastern spruce budworm. The impact of gaps was evaluated on the basis of changes in the number, the period of recruitment, and the composition of tree saplings present within gap areas. Changes were measured along the gap size gradient, and according to the pattern of recent budworm epidemics. Results: The gap fraction is highly variable (18%‐64%) and is on average relatively high (42%). Gap sizes have a positively skewed distribution. In most cases the growth rate among gap filling saplings increased sufficiently to date disturbance events. The composition and the structure of understory trees were affected by gap formation. The number of shade‐intolerant tree species did increase during or following periods of particularly severe canopy disturbances. However, the establishment or survival of shade intolerant species was not restricted to larger gaps or more intensely disturbed periods. Conclusions: In sub‐boreal forests of Eastern Canada, small scale disturbances in the tree canopy influence stand regeneration dynamics, but not to the extent that parameters such as sapling composition and recruitment patterns depend on gap regime characteristics.     

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.     

Variation in the population structure between a natural and a human‐modified forest for a pioneer tropical tree species not restricted to large gaps

The distribution of tree species in tropical forests is generally related to the occurrence of disturbances and shifts in the local environmental conditions such as light, temperature, and biotic factors. Thus, the distribution of pioneer tree species is expected to vary according to the gap characteristics and with human disturbances. We asked whether there was variation in the distribution of a pioneer species under different environmental conditions generated by natural disturbances, and between two forests with contrasting levels of human disturbance. To answer this question, we studied the distribution patterns and population persistence of the pioneer tree species Croton floribundus in the size and age gap range of a primary Brazilian forest. Additionally, we compared the plant density of two size‐classes between a primary and an early successional human‐disturbed forest. Croton floribundus was found to be widespread and equally distributed along the gap‐size gradient in the primary forest. Overall density did not vary with gap size or age (F‐ratio = 0.062, P = 0.941), and while juveniles were found to have a higher density in the early successional forest (P = 0.021), tree density was found to be similar between forests (P = 0.058). Our results indicate that the population structure of a pioneer tree species with long life span and a broad gap‐size niche preference varied between natural and human‐disturbed forests, but not with the level of natural disturbance. We believe this can be explained by the extreme environmental changes that occur after human disturbance. The ecological processes that affect the distribution of pioneer species in natural and human‐modified forests may be similar, but our results suggest they act differently under the contrasting environmental conditions generated by natural and human disturbances.     

Gap formation, microsite variation and the conifer seedling occurrence in a subalpine old-growth forest, central Japan

To evaluate the effects of canopy gaps and forest floor microsites (soil, fallen logs, root-mounds, buttresses and stumps) on regeneration of subalpine forests, the gap regeneration and seedling occurrence of conifers (Abies mariesii, Abies veitchii, Picea jezoensis var. hondoensis and Tsuga diversifolia) were studied in two stands of a subalpine old-growth forest, central Japan. The percentage of gap area to total surveyed area was 11.2–11.3% in the stands. Gap regeneration was not common for P. jezoensis var. hondoensis and T. diversifolia. In contrast, gap regeneration by advanced regeneration was relatively common for Abies. Seedling occurrence of P. jezoensis var. hondoensis and T. diversifolia was restricted on elevated surfaces such as stumps and root-mounds, while Abies seedlings could occur on soil as well as on elevated surfaces. Rotten stumps were the most favorable microsites for conifer seedling occurrence, which covered small area in the forest floor. Although canopy gaps were not always favorable for seedling occurrence, all conifer seedlings were larger under canopy gaps than under closed canopy. Canopy gaps and forest floor microsites clearly affected seedling occurrence and growth of conifers. This suggests that regeneration of conifers is related to the difference of growth advantage under canopy gaps and favorable microsites for seedling occurrence.     

The limited role of canopy gaps in the successional dynamics of a mature mixed Quercus forest remnant

Question: Are canopy gap dynamics responsible for driving the structural and compositional changes that have occurred over a 26‐year period in a mature Quercus forest remnant? Location: Dobbs Natural Area, an unlogged 3.6‐ha forest preserve in west‐central Indiana, USA. Methods: We analyzed mapped permanent plot data for a site that illustrates a trend common in Quercus‐dominated forests in eastern North America, where recruitment of new stems is dominated by mesophytic, shade‐tolerant species such as Acer saccharum, rather than Quercus. We developed a GIS database from stand census measurements taken in 1974 and 2000, employing it to conduct tree‐by‐tree comparisons that allow direct determination of ingrowth, mortality and survivorship, and to relate the spatial patterns of subcanopy dynamics to canopy gap occurrence. Results: The re‐census shows modest changes in canopy composition, but much greater turnover in the subcanopy. Nearly half of all individuals originally present died; much of this mortality resulted from a major decline in subcanopy Ulmus americana. While overall density remained fairly constant, the subcanopy experienced substantial ingrowth of shade‐tolerant Acer saccharum, Fagus grandifolia, and Tilia americana. Canopy gaps, although forming at rates in the upper range of regional averages, did not significantly benefit subcanopy populations of Quercus spp. or most other taxa with limited shade tolerance. Conclusions: Canopy gaps play a minor role in driving the recent demographic trends of this stand. The spatial and temporal scales of light availability in gaps do not support regeneration of most shade‐intolerant species. Compositional change parallels a historical shift in light regimes.     

Tree population dynamics,stand structure and species composition in the montane virgin forest of Vallibäcken,northern Sweden

The population dynamics of Betula pubescens and Picea abies in a boreal forest near Kvikkjokk, northern Sweden, are governed by a process of storm gap regeneration similar to the gap regeneration described for boreo-nemoral forests. Cumulative age distribution curves, interpreted as static survivorship curves, lead to a simple theory of differential survival based on properties of the species, i.e. shade tolerance and relative growth rate. The theory is sustained by diameter and height distributions and by the spatial distributions of logs and of trees in different life-phases. Species of the field and ground layers respond differentially to gap formation and the ensuing successional stages. Browsing by moose (Alces alces) may prevent tree species, mainly Sorbus aucuparia, Betula pubescens and Pinus sylvestris, from developing into a tree layer. The regeneration ability for tree species growing in a stand at 460 m a.s.l. is limited compared with the regeneration at 330 m a.s.l., and typical storm gap formation involving more than one tree seems to occur rarely if at all, while overthrown trees with exposed rootplates are uncommon. Spruce at 460 m a.s.l. shows also a lower growth rate and a lower height/diameter ratio compared to the lower situated stands.     

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.     

Treefall Gaps and Regeneration Composition in the Laurel Forest of Anaga (Tenerife): a Matter of Size?

The size of treefall gaps is an important determinant of regeneration composition in tropical and temperate forests. Preliminary studies in the laurel forest of Tenerife have shown that small gaps (<100 m²) were the most numerous. However, due to this small size, no significant differences were found between regeneration in gaps and regeneration below the canopy. Because infrequent large gaps (>100 m²) are present in the laurel forest, we analyzed the regeneration in these large uncommon gaps, considering their potentially important role in the dynamics of the system. Our main hypothesis is that large gaps are important disturbance to ensure the regeneration and stablishment of shade intolerant species. Only five gaps larger than 100 m² (ranging from 125–268 m²) were found in the study area. Data from a further 20 small gaps (<100 m²), analysed in a previous study, was also included. Control plots were examined close to the gaps in order to determine regeneration below the closed canopy. We did not find a significant difference between regeneration density in the gaps (<100 m²) and regeneration below the canopy in the control plots. Contrary to our expectations, regeneration was lower in the large gaps than under the canopy. The open canopy in the large gaps increases light intensity, and has a negative effect on the germination and growth of shade-tolerant tree species like Viburnum tinus (although non-statistically significant); however, the increase in light intensity is not sufficient to stimulate the germination of shade-intolerant tree species. The effects of treefall gaps in the dynamics of the laurel forest of Anaga should be not considered as significant in comparison to other factors such as human disturbances or infrequent disturbances (land slides or hurricanes).     

Gap formation and regeneration of tropical mangrove forests in Ranong,Thailand

Tropical mangrove forests are characterized by clear zonation along a tidal gradient, and it has been supposed that the zonation is primarily controlled by soil factors. However, effects of disturbance on mangrove forests are still not well understood and may play an important role on the vegetation patterns and forest dynamics in some forest formations. In this study, the pattern of disturbance regime and its effects on regeneration of tropical mangrove forests along a tidal gradient were investigated in Ranong, Thailand. We established one or two 0.5 ha plots in four vegetation zones, i.e. Sonneratia alba–Avicennia alba zone, Rhizophora apiculata zone, Ra – Bruguiera gymnorrhiza zone, Ceriops tagal–Xylocarpus spp. zone. Gap size (percentage gap area to total study area and individual gap size) was the largest in Sa–Aa zone which is located on the most seaward fringe, and it declined from seaward to inland. Canopy trees of S. alba and A. alba had stunted trunks and showed low tree density. On the contrary, canopy dominants in the other three inland zones, e.g. R. apiculata, B. gymnorrhiza, and Xylocarpus spp., had slender trunks and showed high tree density. Accordingly, differences in disturbance regime among the four zones were resulted from the forest structural features of each zone. Disturbance regime matched with regeneration strategies of canopy dominants. Seedlings and saplings of S. alba and A. alba, which need sunny condition for their growth, were abundant in gaps than in understorey. By contrast, R. apiculata, B. gymnorrhiza, and Xylocarpus spp., which can tolerate less light than S. alba and A. alba, had greater seedling and sapling density under closed canopy than gaps. Many large gaps may enhance the abundance of S. alba and A. alba in Sa–Aa zone, and a few small gaps may prevent the light demanding species to establish and grow in the other inland zones. Correspondence of disturbance regime and regeneration strategies (e.g. light requirement) of canopy dominants may contribute to the maintenance of the present species composition in each of the vegetation zones.     

Gap characteristics and regeneration in Bwindi Impenetrable National Park, Uganda

Before Bwindi Impenetrable forest, Uganda, became a national park in 1991, there was a high level of human activity in much of the forest, especially cutting of large trees for timber by pitsawyers. This created extensive gaps in this tropical Afromontane rain forest. We quantified and compared tree regeneration in three sites that were logged at different intensities. Gap sizes in Bwindi, even under fairly natural conditions are very large (mean =4460.1 m²). Logging further enlarged the gap sizes and had a negative impact on tree regeneration. The study shows the strong role of logging disturbance in promoting an alternative successional pathway, where the large gaps created by logging are in a low‐canopy state dominated by a dense tangle of herbs, shrubs, and herbaceous or semi‐woody climbers. We recommend periodic monitoring of gap size and tree regeneration in the gaps to ascertain the trend of recovery from past logging disturbance.     

Estimating Canopy Structure in an Amazon Forest from Laser Range Finder and IKONOS Satellite Observations1

Canopy structural data can be used for biomass estimation and studies of carbon cycling, disturbance, energy balance, and hydrological processes in tropical forest ecosystems. Scarce information on canopy dimensions reflects the difficulties associated with measuring crown height, width, depth, and area in tall, humid tropical forests. New field and spaceborne observations provide an opportunity to acquire these measurements, but the accuracy and reliability of the methods are unknown. We used a handheld laser range finder to estimate tree crown height, diameter, and depth in a lowland tropical forest in the eastern Amazon, Brazil, for a sampling of 300 trees stratified by diameter at breast height (DBH). We found significant relationships between DBH and both tree height and crown diameter derived from the laser measurements. We also quantified changes in crown shape between tree height classes, finding a significant but weak positive trend between crown depth and width. We then compared the field‐based measurements of crown diameter and area to estimates derived manually from panchromatic 0.8 m spatial resolution IKONOS satellite imagery. Median crown diameter derived from satellite observations was 78 percent greater than that derived from field‐based laser measurements. The statistical distribution of crown diameters from IKONOS was biased toward larger trees, probably due to merging of smaller tree crowns, underestimation of understory trees, and overestimation of individual crown dimensions. The median crown area derived from IKONOS was 65 percent higher than the value modeled from field‐based measurements. We conclude that manual interpretation of IKONOS satellite data did not accurately estimate distributions of tree crown dimensions in a tall tropical forest of eastern Amazonia. Other methods will be needed to more accurately estimate crown dimensions from high spatial resolution satellite imagery.     

Amazon forest carbon dynamics predicted by profiles of canopy leaf area and light environment

Tropical forest structural variation across heterogeneous landscapes may control above‐ground carbon dynamics. We tested the hypothesis that canopy structure (leaf area and light availability) – remotely estimated from LiDAR – control variation in above‐ground coarse wood production (biomass growth). Using a statistical model, these factors predicted biomass growth across tree size classes in forest near Manaus, Brazil. The same statistical model, with no parameterisation change but driven by different observed canopy structure, predicted the higher productivity of a site 500 km east. Gap fraction and a metric of vegetation vertical extent and evenness also predicted biomass gains and losses for one‐hectare plots. Despite significant site differences in canopy structure and carbon dynamics, the relation between biomass growth and light fell on a unifying curve. This supported our hypothesis, suggesting that knowledge of canopy structure can explain variation in biomass growth over tropical landscapes and improve understanding of ecosystem function.     

Assessing the above‐ground biomass of a complex tropical rainforest using a canopy crane

Abstract Current estimates of the total biomass in tropical rainforests vary considerably; this is due in large part to the different approaches that are used to calculate biomass. In this study we have used a canopy crane to measure the tree architectures in a 1 ha plot of complex mesophyll vine forest at Cape Tribulation, Australia. Methods were developed to measure and calculate the crown and stem biomass of six major species of tree and palm (Alstonia scholaris (Apocynaceae), Cleistanthus myrianthus (Euphorbiaceae), Endiandra microneura (Lauraceae), Myristica insipida (Myristicaceae), Acmena graveolens (Myrtaceae), Normanbya normanbyi (Arecaceae)) using the unique access provided by the crane. This has allowed the first non‐destructive biomass estimate to be carried out for a forest of this type. Allometric equations which relate tree biomass to the measured variable ‘diameter at breast height’ were developed for the six species, and a general equation was also developed for trees on the plot. The general equation was similar in form to equations developed for tropical rainforests in Brazil and New Guinea. The species equations were applied at the level of families, the generalized equation was applied to the remaining species which allowed the biomass of a total of 680 trees to be calculated. This has provided a current estimate of 270 t ha⁻¹ above‐ground biomass at the Australian Canopy Crane site; a value comparable to lowland rainforests in Panama and French Guiana. Using the same tree database seven alternative allometric equations (literature equations for tropical rainforests) were used to calculate the site biomass, the range was large (252–446 t ha⁻¹) with only three equations providing estimates within 34 t ha⁻¹ (12.5%) of the site value. Our use of multiple species‐specific allometric equations has provided a site estimate only slightly larger (1%) than that obtained using allometric equations developed specifically for tropical wet rainforests. We have demonstrated that it is possible to non‐destructively measure the biomass in a complex forest using an on‐site canopy crane. In conjunction the development of crown maps and a detailed tree architecture database allows changes in forest structure to be followed quantitatively.