Estimating leaf area index in different types of mature forest stands in Switzerland: a comparison of methods

Leaf area index (LAI) was estimated at 15 sites in the Swiss Long-Term Forest Ecosystem Research Programme (LWF) in 2004–2005 using two indirect techniques: the LAI-2000 plant canopy analyzer (Licor Inc.) and digital hemispherical photography, applying several exposure settings. Hemispherical photographs of the canopy were analysed using Hemisfer, a software package that offers several new features, which were tested here: (1) automatic thresholding taking the gamma value of the picture into account; (2) implementation of several equations to solve the gap-fraction inversion model from which LAI estimates are derived; (3) correction for ground slope effects, and (4) correction for clumped canopies. In seven broadleaved stands in our sample set, LAI was also estimated semi-directly from litterfall. The various equations used to solve the gap-fraction inversion model generated significantly different estimates for the LAI-2000 measurements. In contrast, the same equations applied in Hemisfer did not produce significantly different estimates. The best relationship between the LAI-2000 and the Hemisfer estimates was obtained when the hemispherical photographs were overexposed by one to two stops compared with the exposure setting derived from the reading of a spotmeter in a canopy gap. There was no clear general relationship between the litterfall and the LAI-2000 or the hemispherical photographs estimates. This was probably due to the heterogeneity of the canopy, or to biased litterfall collection at sites on steep slopes or sites subject to strong winds. This study introduces new arguments into the comparison of the advantages and drawbacks of the LAI-2000 and hemispherical photography in terms of applicability and accuracy.     

Calculation method for sunshine duration in canopy gaps and its application in analyzing gap light regimes

Understory light is essential to the growth and survival of plants, yet the light varies temporally and spatially within forest gaps. Measurement of understory light levels using light sensors is both time and labor intensive. Sunshine duration (SD) has a strong correlation with solar radiation and has been the variable most widely used for estimating solar radiation. The power of SD-based methods for estimating solar radiation lies in its ability to quickly estimate light levels. Although several calculation methods for SD within canopy gaps are available, all the methods oversimplify canopy gaps by classifying them as cylindrical or ellipsoid and thus have a relatively low level of accuracy. In this study we developed a calculation method for SD at any given point within natural canopy gaps and we used SD to estimate solar radiation within 12 canopy gaps on Mt. Taibai in China based on the Angström-Prescott model. We then evaluated the SD-based solar radiation by the total and direct solar radiation derived from a gap light index based on hemispherical photographs (HP). Finally, we analyzed the spatial-temporal characteristics of light levels within these 12 gaps by using the solar radiation derived from SD at hundreds of simulated points in each gap. Our results showed that (1) SD-based solar radiation was not statistically different from HP-based direct solar radiation and had a strong linear correlation with HP-based total solar radiation; (2) growing-season daily mean solar radiation within the 12 gaps varied from 0.08 to 13.28 MJ m⁻² day⁻¹ with an average of 4.13 MJ m⁻² day⁻¹; (3) solar radiation had a positive correlation with the ratio of the square root of the canopy gap area to the mean canopy height. This relationship was significant but solar radiation had no correlation with the canopy gap area; (4) among most gaps solar radiation was greatest in May while potential SD was longest in June 2008. From these results we can conclude that the SD-based method for estimating solar radiation developed in this study can quickly and accurately estimate light levels at any specified point within canopy gaps. SD-based solar radiation appears to be a good choice for studies on the spatial-temporal characteristics of gap light levels.     

Effect of settings of digital fisheye photography to estimate relative illuminance within forest under low light conditions

In order to evaluate the effects of settings of digital fisheye photography on estimates of relative illuminance (RI) within a forest under low light conditions, we assessed the effectiveness of some estimators, i.e., canopy openness (CO), sky factor (SF), and diffuse transmittance (DIF). The estimates were calculated from digital hemispherical photographs taken with two different exposure settings, i.e., the reference exposure setting and the auto-exposure setting. We also presented a new estimator (%Exposure), which does not require image analysis, and compared it with other estimators calculated from a digital hemispherical photograph using image analysis. The CO, SF, and DIF calculated from digital hemispherical photographs taken with auto-exposure setting were not significantly correlated with RI. However, those calculated from digital hemispherical photographs taken with reference exposure settings were significantly correlated with RI. On the other hand, %Exposure showed a slightly higher correlation with RI compared with the other estimators.     

Instantaneous and potential radiation effect on underplanted European beech below Norway spruce canopy

In even-aged, 120-year-old Norway spruce stands with underplanted beech (in 1995) four permanent research plots were established, each 100 × 100 m in size. Twelve subplots were selected on each plot along a light gradient from complete canopy closure to open light conditions. On each plot, photon flux density (PFD) was measured continuously, and during same time interval (August 2008, 2009), color digital hemispherical photographs were taken. Cumulative PFD values for measuring points were compared with potential PFD radiation obtained from hemispherical photograph analysis for different angle of hemisphere and different parameters from hemispherical image analysis (gap fraction, total openness and direct, indirect and total amount of radiation). Cumulative and average daily values for the plots were compared; 120° hemispherical photograph angle, gap fraction and total openness were the variables that explained the largest proportion of variance in light transmittance. Determination coefficients between direct and total light component were highest for the total potential radiation and lowest for the diffuse light component. Comparison between potential and instantaneous light measurements for radial, height increment and SLA of young beech showed that instantaneous radiation measurements explained height increment best; 120° of hemisphere proved to be the best explaining angle.     

Correlations between canopy gaps and species diversity in broad-leaved and Korean pine mixed forests

Regeneration of tree species associated with canopy gaps in broad-leaved Korean pine forests was investigated. Species diversity in gaps and under closed canopy was compared, the relationship between biodiversity and gap structure was analyzed. Results indicate that there were significant differences between tree species diversity in gaps and that under canopy (p<0.01). In terms of Shannon-Wiener index, evenness index, and abundance index, the biodiversity in gap community were higher than those under forest canopy in regeneration layer. In terms of Simpson’s dominance index, the dominance of certain species in the regeneration layer increased from gaps to closed canopy (p<0.01). In contrast, trends of biodiversity changes of succession layer in gaps and under closed canopy were opposite. Tree species diversity of different layers reacted directly to the change of gap size class. For example, Shannon-Wiener index and abundance index is higher and Simpson’s dominance index is the lowest in succession layer of medium-size gap (100–250 m²) in the broad-leaved Korean pine forest of Changbai Mountains. Shannon-Wiener index reached the highest in a size of ≥250 m² and <100 m², reached the lowest in a size of 200–250 m² in the regeneration layer. Simpson’s dominance index reached its maximum when the gap size was between 200 and 250 m². Generally, species of different layers reacted differently to the changes of gap size classes. The gap size class with more seedlings did not correspond to size class containing more medium-size trees. Tree species diversity indices in the two layers behaved reciprocally during the development process of forest gaps. __________ Translated from Chinese Journal of Applied Ecology, 2005, 16(12): 2,236–2,240 [译自: 应用生态学报, 2005, 16(12): 2,236–2,240]     

Can a natural experiment be used as a tool to design partial cutting regimes? The decline of <Emphasis Type="Italic">Austrocedrus chilensis</Emphasis> forests,an example

The forests of Austrocedrus chilensis in Argentina suffer mortality from what is locally known as ‘mal del ciprés’. This disturbance usually appears as aggregations of dead trees of varying sizes. The mortality process seems unpredictable, and regeneration establishment is variable under these conditions. The highly variable mortality rates results in a natural experiment that allows the correlation of overstory mortality with regeneration rate. This correlation should then make it possible to estimate the regeneration success which would follow after different levels of partial cutting, allowing the purposeful direction of the regeneration process. We collected data from eight Austrocedrus chilensis-dominated stands and reconstructed overstory mortality over time and correlated it to the regeneration recruitment. Mortality showed two patterns: increasing ‘continuous’ mortality over time, and ‘pulses’ of important basal area loss in short periods. Overstory mortality had an effect on the recruitment and composition of the regeneration by allowing different species establishment depending on the degree of mortality. By quantifying mortality basal area and recruitment over time, we calculated estimates that can provide partial cutting guidance. Partial cuttings would then allow the purposeful direction of the regeneration process. Stand reconstruction studies provided a useful tool to help designing regeneration systems using a natural experiment. Some specific management implications for symptomatic A. chilensis forests are given.     

The influence of canopy patch mosaics on understory plant community composition in boreal mixedwood forest

We assessed the composition of understory vascular plant communities in relation to the mosaic of canopy patch types, and their associated structure and environment, within unmanaged, mature boreal mixedwood forests in western Canada. Within a 30 km² area, we sampled patches of four different canopy types: conifer-dominated, broadleaf-dominated, mixed conifer-broadleaf, and canopy gaps (total n = 98). There were significant differences in understory composition among the four patch types (based on multi-response permutation procedure (MRPP)) and these were mainly due to differences in relative abundances of understory species. The understory communities of conifer patches were characterized by low abundances of shade intolerant species while shade-tolerant and evergreen species were indicators (based on an indicator species analysis (ISA)). Understory communities under gap and broadleaf patches were characterized by higher abundances of grasses and shade intolerant species. Gap, broadleaf, and mixed patches had higher abundances of certain shrub species than did conifer patches. The patch types also differed in terms of their environmental conditions. Conifer patches had drier, cooler soils and the lowest understory light. Broadleaf patches had the warmest soils while understory light during the leaf-off period was similar to that of canopy gaps. Gap patches had the lowest litter cover and PO₄⁻ availability and the highest light. Seven environmental variables (soil moisture, soil temperature, total light during the leaf-off period, cover of coarse and fine downed woody material, and availability of NH₄⁺ and Ca²⁺) were significantly related to understory species composition (in a constrained ordination by means of a distance-based redundancy analysis (db-RDA); 16.5% of variation in understory community data explained). Even within a single patch type, there was substantial environmental variation that was related to understory species composition. Our study suggests that the mosaic of canopy patches within mixedwood forests supports coexistence of both early and late successional understory plant species in mixedwood stands. Maintaining the mixture of canopy patch types within mixedwood stands will be important for conserving the natural patterns of understory plant composition in boreal mixedwood forests.     

Differential overstory leaf flushing contributes to the formation of a patchy understory

Understory individuals were found to form patches in a 100-year-old deciduous broad-leaved forest. The closed forest canopy was uniform, and so the light conditions at various locations across the forest floor differed little after the leaf flush of the overstory. To explain the distribution pattern in the understory, a hypothesis was proposed: in spring, the forest floor is divided into patches according to the timing of leaf flush of the overstory individuals, and the light conditions are more favorable for understory plants under the crowns of trees with later-flushing leaves. In the plot, three groups of early, intermediate, and late, were recognized in the overstory concerning the timing of leaf flush. As for the start of leaf flush, a difference of 31.6 days was recognized among tree species, and for the end of leaf flush, a difference of 40.3 days. In the spring of 1998, the relative photosynthetic-photon-flux density under an intensively studiedCastanea crenata tree (late-flushing species) usually showed higher values than that under a similarly studiedAcer mono tree (early-flushing species). Analysis of the spatial-distribution pattern using Morisita’s1δ index revealed that the understory community had an aggregated distribution. In the overstory, the late- and the intermediate-flushing-species groups showed aggregated distributions, while the early-flushing-species group showed random distribution. Spatial correlation between the understory and the overstory was analyzed by using Morisita’sRδ index. The distribution of whole understory community spatially co-occurred with that of the late-flushing-species group of the overstory. In contrast, the understory community was less developed below the members of the early-flushing-species group of the overstory. We consider that the data presented here support our hypothesis, and we suggest that the growth and survival of understory individuals were promoted in the places receiving light for long periods in spring.     

Oak (<Emphasis Type="Italic">Quercus robur</Emphasis> L.) regeneration as a response to natural dynamics of stands in European hemiboreal zone

The oak (Quercus robur L.) regeneration intensity was assessed in the core area of the Białowieża National Park (BNP) in Poland with respect to the selected ecological factors. The emphasis was placed on the response of oak regeneration to disturbances, including the large-scale dieback of spruce stands. Defining their effect could help predicting the role of oak in naturally developing lowland forest ecosystems in the European hemiboreal zone. The results of the study challenge the opinion that the ‘lime-oak-hornbeam forest’ is a ‘climax’ community, confirming a very poor regeneration represented by only two saplings taller than 0.5 m per hectare. By contrast, in spruce-dominated communities, from 49 to 848 taller saplings per hectare were found. The occurrence of saplings was associated with discontinuous canopy of late seral stage of stands, as well as with large gaps. Most of the best quality grown-up oak saplings developed in the immediate neighbourhood of spruce logs. The results of the research indicate that ‘lime-oak-hornbeam forest’ (Tilio-Carpinetum) should be rather perceived as a transient community, evolved from relict, culturally modified, oak woodlands. Because spruce had become a dominating species only after abandoning in mid-1800s the historical regime of anthropogenic disturbances (involving frequent forest fires), the observed phenomena related to the disintegration of spruce stands had probably no precedent over the last 500 years. To confirm whether the massive decline of spruce stands will finally result in the successful establishment of the new canopy oak generation, both in Białowieża and other forests of hemiboreal zone, further research is needed.     

Stand structure and the regeneration ofChamaecyparis pisifera (Sieb. et Zucc.) Endl. on sites with different soil development in an old-growth coniferous forest, central Japan

Stand structure and the regeneration ofChamaecyparis pisifera on sites with and without well-developed soil in an old-growth coniferous forest, Akasawa Forest Reserve, central Japan, were investigated, along with their relationships to other important coniferous species. Of stems ofC. pisifera≥5 cm diameter at breast height, stems with intermediate size were absent in stands on the developed-soil site, while those with smaller size increased in stands on the undeveloped-soil site, which supported a higher density ofC. pisifera. In the stands without recent tree-fall of canopy stems on the developed-soil site, canopy stems ofC. pisifera were distributed uniformly and the understory stems, patchily. On the other hand, both canopy and understory stems ofC. pisifera in stands on the undeveloped-soil site were distributed patchily. In the understory,C. pisifera occurred as clonal patches formed by vegetative reproduction on various types of microsites including bare rocks. However, the clonal patches did not occur in a stand on the developed-soil site with dense saplings of an evergreen conifer,Thujopsis dolabrata, which can expand vegetatively with a well developed soil layer. Gap formation may induceC. pisifera to develop tree-form stems from shrubby clonal patches. On developed-soil sites,C. pisifera stands that survive a long disturbance-free period on this site need catastrophic disturbance for their regeneration, which will eliminateT. dolabrata in the understory and create bare soil for colonization ofC. pisifera from seeds. On undeveloped-soil sites,C. pisifera replaces itself continuously by effective vegetative reproduction. Ground instability and the wet condition of this site may promote the replacement.     

Light distribution in scattered-trees open woodlands in Western Spain

We have studied the percentage of radiation transmitted through the tree canopy to the underlying pasture and crops in dehesas of Southwest Spain by means of fish-eye photographs taken at different distances from the tree. Thirty-six trees were studied covering all the diametric classes (0.1–14 m canopy width) of two stands, with mean density of 19 mature trees ha⁻¹. Intercepted light decreased with distance following an logistic curve, indicating a rapid increase in the light availability with distance from the tree. For mature trees, radiation was constant beyond 20 m. Applying a multivariable regression light equation, distance, stem diameter and canopy width explained more than 88% of the light variability for each orientation studied. A simple model was built up from light equations, tree growth curves and allometric relationships. From this model, we have estimated that radiation available for crops and pasture decreased up to 21% due to the presence of trees in a standard dehesa with 24 mature trees ha⁻¹ and 13% of canopy cover. In addition, we have generated different radiation maps of virtual dehesas differing in tree age, density and arrangement, which could be useful to determine optimal tree planting schemes and consequent pruning and thinning regimes.     

Canopy light transmittance in Douglas-fir--western hemlock stands

We measured vertical and horizontal variation in canopy transmittance of photosynthetically active radiation in five Pseudotsuga menziesii (Mirb.) Franco-Tsuga heterophylla (Raf.) Sarg. (Douglas-fir-western hemlock) stands in the central Cascades of southern Washington to determine how stand structure and age affect the forest light environment. The shape of the mean transmittance profile was related to stand height, but height of mean maximum transmittance was progressively lower than maximum tree height in older stands. The vertical rate of attenuation declined with stand age in both the overstory and understory. A classification of vertical light zones based on the mean and variance of transmittance showed a progressive widening of the bright (low variance and high mean) and transition (high variance and rapid vertical change) zones in older stands, whereas the dim zone (low variance and mean) narrowed. The zone of maximum canopy surface area in height profiles, estimated by inversion of transmittance profiles, changed from relatively high in the canopy in most young stands ("top-heavy") to lower in the canopy in older stands ("bottom-heavy"). In the understory, all stands had similar mean transmittances, but the spatial scale of variation increased with stand age and increasing crown size. The angular distribution of openness was similar in all stands, though the older stands were less open at all angles than the younger stands. Understory openness was generally unrelated to transmittance in the canopy above. Whole-canopy leaf area indices, estimated using three methods of inverting light measurements, showed little correspondence across methods. The observed patterns in light environment are consistent with structural changes occurring during stand development, particularly the diversification of crowns, the creation of openings of various sizes and the elaboration of the outer canopy surface. The ensemble of measurements has potential use in distinguishing forests of differing ages that have similar stature.     

Variation in canopy structure, light and soil nutrition across elevation of a Sri Lankan tropical rain forest

Mixed dipterocarp forests are perhaps the single most important rain forest type in the wet tropics. Only a few studies have purposefully examined differences in resource availability across mixed dipterocarp forest landscapes by simply measuring the abiotic variables of light, soil nutrition and soil water availability in relation to forest structure. We sought to directly measure the environment of canopy gaps across elevation and geology—from lowland mixed dipterocarp forest (100 m amsl) to lower montane dipterocarp forest (1200 m amsl) in southwest Sri Lanka. Middle elevation gap sites (300–900 m amsl) were subdivided into valley, mid-slope and ridge topographic positions. Eighteen natural disturbances all of which were canopy openings caused by tree fall, were randomly selected within primary rain forest that ranged across 100–1200 m elevation. Plots were placed in gap centers and in adjacent understories and measurements taken of forest structure (basal area, canopy height, canopy cover index, CCI), shade (light sensors—photosynthetically active radiation [PAR], canopy hemispherical photographs—global site factor [GSF]) and soil nutrition (pH, exchangeable Al, K, Mg and Ca; Total N; and plant available P). Soil moisture was measured at bi-weekly intervals for five years across middle elevation sites only (300–900 m amsl). Stand basal area, mean canopy height, and canopy cover index all declined with increase in elevation. Understory PAR and GSF decreased with increases in canopy height, basal area and CCI. Size of canopy opening decreased with increase in elevation, but PAR and GSF increased. Valley sites had significantly greater levels of mean percent soil water content as compared to mid-slope and ridge sites of middle elevation sites. However, at the onset of the southwest monsoons in May all sites were similar. Differences were most pronounced during the dry season (December–April). No differences in soil moisture content could be found between gap and understory microsites. K and Ca in gap centers and adjacent forest understories increased with increase in elevation and change in associated geology. pH increased and Al decreased with elevation and associated geology but only for forest understory conditions. Results demonstrate strong differentiation in soil and light resources with elevation that appears related to size of tree-fall disturbance, stature of the forest, topographic position and underlying geology and soil-weathering environment. This suggests that forest management and conservation practices need to develop and tailor techniques and treatments (silviculture) to the forest that emulate and/or account for change in elevation, geology and topographic position. Further studies are needed to identify which are the primary underlying mechanisms (e.g. temperature, wind, soil nutrients, soil moisture availability) defining change in forest structure across elevation.     

Patterns in spatial extent of gap influence on understory plant communities

Gap formation in forests can have impacts on forest ecosystems beyond the physical boundary of the canopy opening. The extent of gap influence may affect responses of many components of forest ecosystems to gap formation on stand and landscape scales. In this study, spatial extent of gap influence on understory plant communities was investigated in and around 0.1 and 0.4 ha harvested canopy gaps in four young Douglas-fir (Psuedotsuga menziesii) dominated stands in western Oregon. In larger gaps, the influence of gap creation on understory plant communities in surrounding forests was minimal. The area showing evidence of gap influence extended a maximum of 2 m beyond the edge of the canopy opening, suggesting that the area affected by gap creation did not differ greatly from the area of physical canopy removal. In smaller gaps, influence of the gap did not extend to the edge of the canopy opening. In fact, the area in which understory vegetation was influenced by gap creation was smaller than the physical canopy opening. Gap influence appears to be limited to areas where ruderal or competitor species are able to replace stress-tolerator species, likely due to elimination or reduction of these species by physical disturbance or competition. The limited gap influence extent exhibited here indicates that gap creation may not have a significant effect on understory plant communities beyond the physical canopy opening. This suggests a limited effectiveness of gaps, especially smaller gaps, as a tool for management of understory plant diversity, and perhaps biodiversity in general, on a larger scale.     

Comparison of stand dynamics after dieback caused by pine wilt disease among pine forests with different management regimes in western Japan

In pine forests damaged by pine wilt disease, in western Japan, the effect of protection regimes of pine trees on the stand dynamics were examined in the following four stands: (1) lightly damaged stand (age 30–40 years) with no procedure in operation for protecting pine trees; (2) severely damaged stand (age 30–40 years) with no procedure in place for protecting pine trees; (3) severely damaged stand (age 50 years) with a selective cutting of infected trees; (4) severely damaged stand (age 30–40 years) with a selective cutting of infected trees. All the stands had been abandoned before the pine wilt disease damage. The understory structure of the severely damaged stand with no protection procedure was similar to that of the lightly damaged stand. Frequent invasion by tree species and acceleration in the growth of understory trees occurred after the dieback in the selective cutting stand. These results suggest that a deficiency in the canopy layer caused by the dieback resulted in low disturbance intensity in the early stages after the dieback, but the selective cutting increased the intensity by the reduction in the understory as well as the canopy layer. The intensity of the disturbance in the selective cutting stands was larger in the younger stand because it had a higher density of selectively cut pine trees. The different stand structure of pine forests occurred after the dieback because the intensity of the disturbance varied as a result of the selective cutting operation and the stand age.     

Assessing the relationships between stand development and understory vegetation using a 420-year chronosequence

The large-scale conversion of old forests to tree plantations has made it increasingly important to understand how understory vegetation responds to such landscape changes. For instance, in some forest types a reduction in understory richness and cover is thought to result from the development of canopy closure in plantations, although there is a paucity of empirical data demonstrating this relationship. We used a 420-year forest chronosequence as a case study to assess the relationship between stand age, tree canopy cover and understory vascular plant richness and composition in the Siskiyou Mountains of Oregon. The chronosequence consisted of six young managed (age 7–44) and nine older unmanaged (age 90–427) stands. All stands were similar in underlying geology, slope, elevation, and aspect. We found a non-linear relationship between stand age and richness, in which richness was highest in the youngest stands, reached a low in mid-aged stands (∼55 years), then increased in the oldest stands. We also found that percent tree canopy cover was correlated with total understory cover, richness, diversity, and species composition. In general, young stands were characterized by high shrub and graminoid cover and old stands were characterized by an abundant herb layer. Our work suggests that a major component of our study landscape is currently entering the forest stage (canopy closure) characterized by low levels of vascular plant species richness and cover. We use our results to discuss the potential effects of future forest management on understory plants.     

Forest gap dynamics and tree regeneration

When one or a few canopy trees die (or are injured) in a forest, small openings, which are called ‘gaps,’ are formed in the forest canopy and are then filled with other trees. This sort of forest dynamics is termed gap dynamics; a large number of papers and data on gap dynamics have been accumulated since the 1970’s, and gap dynamics has been described in many forest types. In this review, I introduce the basic concepts of gap dynamics and summarize major issues on gap dynamics relating to tree regeneration, with many references. Although enormous studies on gap dynamics of natural forests have been conducted, applications of gap dynamics to forest practice are limited. However, accumulated knowledge on gap dynamics should be useful for sustainable forest ecosystem management, as much of the literature suggests. Recipient of the Japanese Forestry Society Award 2000.     

Species diversity of broad-leaved trees in Cryptomeria japonica plantations in relation to the distance from adjacent broad-leaved forests

The species diversity of broad-leaved trees in relation to the distance from a broad-leaved forest was investigated in Cryptomeria japonica plantations with crown snow damage near the Japan Sea in central Japan. The number and diversity of species and stem density decreased with distance from the broad-leaved forest in maturing gap stands (51–58 years old; >10 years after crown snow damage), but not in recent gap stands (34–42 years old; <10 years after crown snow damage), and increased with improved light conditions in the interior of recent gap stands, but not in that of maturing gap stands. The stem densities of tall and small tree species and woody lianas were greater in the interior of recent gap stands than in maturing gap stands. Woody lianas, which are characteristically shade intolerant, had a high stem density in the interior of recent gap stands. In contrast, the stem density of shrubs was greater in maturing gap stands than in recent gap stands, irrespective of distance. Shade-tolerant shrubs had a high stem density in maturing gap stands. Wind-dispersed and frugivore-dispersed species were concentrated at the edges of maturing gap stands, but some frugivore-dispersed species, which may persist in soil seed banks, occurred in the interior of recent gap stands. Gravity-dispersed species tended to occur both at the plantation edge and in the interior. The differences in the occurrence patterns of broad-leaved trees in the two types of stands reflected the difference in the effect of both the distance from the adjacent broad-leaved forest and the light conditions related to canopy gaps, with the time since gap formation in a region where crown snow damage often occurs.     

Cutting and resprouting of Detarium microcarpum and herbaceous forage availability in a semiarid environment in Burkina Faso

The tree-shrub savanna ‘Forêt Classée de Nazinon’ (Burkina Faso) is submitted to a management of grazing and rotational cutting of Detarium microcarpum. This species resprouts after cutting. In order to investigate whether this silvopastoral land use system is sustainable, aboveground herbaceous biomass was measured on subplots under uncut trees (‘uncut’), next to the stubs of cut trees (‘cut’) and on subplots not influenced by the (former) crowns of trees (‘open grassland’) in four lots. These lots were cut one, three, six and seven years before the study. Vegetation composition of the lots and the composition of the diet of cattle were also determined. Comparisons were made between treatments and lots. Herbaceous biomass was lower in the open grassland subplots than in uncut or cut subplots. We speculate that soil enrichment and more efficient precipitation input in (former) tree crown zones could have resulted in this pattern. Cutting and subsequent resprouting of trees did not lead to significant differences in herbaceous biomass between cut and uncut subplots. The most simple explanation for this is that the trees could extend their roots beyond the location of their neighbouring trees. Biomass and coverage of perennial grasses, mainly Andropogon ascinodis and Andropogon gayanus, did not change in lots cut one, three or six years before the study, but decreased dramatically in lots that were cut seven years before the study. When foraging, cattle spent more than 90% of their time feeding on these species. This indicates that, as a consequence of tree cutting, forage availability may be reduced to the point where local herdsmen are forced to take their cattle to another region. This revised version was published online in August 2006 with corrections to the Cover Date.