Impact of deer on temperate forest vegetation and woody debris as protection of forest regeneration against browsing

A 3-year field experiment with paired exclosure (fenced areas, excluding deer) and control plots (unfenced areas, free access to deer), with two treatments with and without woody debris, was carried out at two sites in a temperate forest in eastern France. The aim of the experiment was to assess the effect of browsing by roe deer (Capreolus capreolus) and red deer (Cervus elaphus) on the diversity and richness of plant species and to test the effectiveness of using woody debris to protect seedlings and saplings from deer browsing. In presence of deer, both plant species richness and diversity were reduced the first year of the study, but this negative impact of deer then disappeared after 3 years. Deer browsing mostly affected species composition of plant communities. We observed a decrease in the abundance of preferred species such as Carpinus betulus, Rubus fructicosus, Rubus idaeus, Anemone nemorosa and Epilobium angustifolium, and palatable species such as Acer spp., Carex spp., Festuca spp. and Mycelis muralis, whereas unpalatable species such as Lamium spp., or species particularly resilient to browsing such as grasses (Brachypodium spp. and Luzula spp.) increased in abundance. The use of woody debris as protection against browsing by deer did not limit damage to seedlings and saplings of the main commercially valuable species, Abies alba and Quercus spp. Instead of limiting deer impact, use of woody debris seemed to increase the negative effect of deer browsing on regeneration in control plots relatively to those without protection.     

Interactions among forestry regeneration treatments,plant vigour and browsing damage by deer.

In a large field experiment we studied the influence of regenerationmethods on the extent to which roe deer (Capreoluscapreolus L.) browse on spruce (Picea abies L.Karst) seedlings. Our objective was to evaluate if treatments that are good atstimulating regeneration may increase browsing damage by deer. Data werecollected between 1993 and 1995, and in the winters of 1993–94 and1994–95 the frequency of browsed seedlings was 13.2% and 13.4%,respectively. The frequency of browsing damage varied considerably betweensitesand clearcuts. Browsing damage was more frequent when the regeneration methodsincluded insecticide treatments. Among insecticide-free treatments, browsingwasheavier on scarified plots than on herbicide-treated or control plots (in whichseedlings were planted on untreated ground and given no post-plantingtreatment). Containerised seedlings were browsed more than bare-rooted ones.Neither the age of clearcut when planting, nor removal of slash, had any effecton browsing. It was demonstrated that regeneration methods that increased plantvigour (as assessed by leader growth and needle colour index) led to morebrowsing damage. Thus, our results support the plant vigour hypothesis, whichstates that many herbivores prefer to feed on vigorous plants because they aremore nutritious. However, the difference in browsing damage betweencontainerised and bare-rooted seedlings could not be explained solely by theplant vigour hypothesis.     

Regeneration dynamics of dominant tree species along an altitudinal gradient in moist temperate valley slopes of the Garhwal Himalaya

The present study was undertaken in moist temperate forest of Mandal-Chopta area in the Garhwal region of Uttarakhand, India. The aim of the present study was to understand the regeneration dynamics of the dominant tree species along an altitudinal gradient in naturally regenerating, restricted access forest. The overall regeneration status was fairly high in the study area. Most of the native canopy and undercanopy dominants had frequent reproduction and expanding populations, which suggests the stability of forest structure/composition and further expansion of dominant species. The overall regeneration of trees in the forest had a greater contribution of middle and understorey species. Because of infrequent reproduction and declining populations of some of the dominant native species viz., Abies pindrow, Alnus nepalensis and Betula alnoides, structural/compositional changes in the future are expected in respective forests dominated by them. Abies pindrow and Taxus baccata need immediate attention by forest managers for their survival in the area. Seedlings were found to be more prone to competition from herb and shrubs than saplings.     

Forest structure and woody plant species composition along a fire chronosequence in mixed pine–oak forest in the Sierra Madre Oriental, Northeast Mexico

Although wildfires are occurring frequently in the pine–oak forests in the Sierra Madre Oriental (northeastern Mexico), data on post-fire succession and forest structure are still rare. Our objectives were to (1) assess the changes in woody plant species composition after fire and to (2) to relate successional patterns to environmental variability. Based on their fire history 23 plots were selected in the Parque Ecológico Chipinque (PECH). Changes in forest structure across the chronosequence of burned stands were deduced from density, height and diameter measurements of trees and shrubs (>5 cm in diameter) in all plots of 1000 m². Differences in woody plant species composition among the plots were evaluated using Shannon evenness measure and the Whittaker's measure and by Hierarchical cluster analysis and Detrended Correspondence Analysis. Hierarchical cluster analysis showed a high similarity among all recently burned plots, independed of the aspect. Multivariate analysis showed that local environmental factors, including time since fire, continue to structure species composition. Oak species (mainly Quercus rysophylla) resprouted successfully after fire and dominated young post-fire stands. Pine species (Pinus pseudotrobus and Pinus teocote) only appeared 18 years after fire and were the dominating species in mature stands (62 years after fire). In contrast, woody plant species composition in older stands tended also to be influenced by factors such as aspect and by the potential solar radiation (PSR) during the growing season. The results demonstrate that in the PECH, natural regeneration is sufficient and woody plant species composition will be similar to pre-fire conditions after 60 years of succession. We conclude that the park managers should consider incorporating natural disturbance regimes into their management practices.     

Effects of 13-year fencing on browsing by sika deer on seedlings on Mt. Ohdaigahara, central Japan

We examined the effects of fencing on deer browsing on seedlings 13 years after the building the experimental fence on Mt. Ohdaigahara, central Japan, where the sika deer (Cervus nippon) population is high. There was no difference in the species number of seedlings between 1991 and 2004. The density of seedlings in the fenced plot in 2004 was three times higher than in 1991. Although the unfenced plots had no seedlings higher than 20 cm, there were seedlings up to 120 cm in the fenced plot. These results suggest that 13-year fencing promoted regeneration of seedlings in this area.     

Analysis on the daily courses of water potential of nine woody species from Cerrado vegetation during wet season

IntroductionAS the concept of water pot6ntial (% was introduced by Saltyer (1967), there were some argumentson it among some physiologist. Some physiologistsopposed it beCause they believed Tvalue might notbe the beSt parameter for describing physiologicalprocesses. However, this concept has been accept6dand used by most plant physiologists for many years(Kramer 1988, Schulze et al. 1988, Passioura 1988,Boyer 1989).ACcording tO our experience (Prado et al. 1994,1995), we believed-t.hat th…     

Seasonal variations of leaf traits and drought adaptation strategies of four common woody species in South Texas,USA

Understanding physiological responses and drought adaptation strategies of woody plant leaf traits in sub-humid to semi-arid regions is of vital importance to understand the interplay between ecological processes and plant resource-allocation strategies of different tree species.Seasonal variations of leaf morphological traits,stoichiometric traits and their relationships of two drought tolerant woody species,live oak(Quercus virginiana)and honey mesquite(Prosopis glandulosa)and two less drought tolerant species,sugarberry(Celtis laevigata)and white ash(Fraxinus americana)were analyzed in a sub-humid to semi-arid area of south Texas,USA.Our findings demonstrate that for the two drought tolerant species,the leguminous P.glandulosa had the highest specific leaf area,leaf N,P,and lowest leaf area and dry mass,indicating that P.glandulosa adapts to an arid habitat by decreasing leaf area,thus reducing water loss,reflecting a resource acquisition strategy.While the evergreen species Q.virginiana exhibited higher leaf dry mass,leaf dry matter content,C content,C:N,C:P and N:P ratios,adapts to an arid habitat through increased leaf thickness and thus reduced water loss,reflecting a resource conservation strategy in south Texas.For the two less drought tolerant deciduous species,the variations of leaf traits in C.laevigata and F.americana varied between Q.virginiana and P.glandulosa,reflecting a trade-off between rapid plant growth and nutrient maintenance in a semi-arid environment.     

Dispersal potentials determine responses of woody plant species richness to environmental factors in fragmented Mediterranean landscapes

While maximizing plant species richness continues to be central in the design, conservation and reforestation action plans, plant life histories are receiving increasing attention in assessments for the conservation of biodiversity in fragmented landscapes. We investigated the determinants of woody plant species (trees, shrubs and climbers) richness in the forest patches of the Guadalquivir river valley, a Mediterranean agricultural landscape with ∼1% forest cover. We analyzed three species richness variables, total, and those corresponding to species with short-distance (ballistic, barochorous, myrmecochorous and short-distance anemochorous) and long-distance (anemochorous, endozochorous, exozoochorous, hydrochorous and dyszoochorous) dispersal systems, which significantly characterize earlier and late successional stages, respectively. We selected eleven predictor variables related to habitat structure (patch area, shape, distances to the nearest patch and reserve, and general isolation), physical environment (temperature, precipitation, elevation, and lithological heterogeneity), and anthropogenic influences (disturbance and proportion of old-growth forest). We used ordinary-least-squares multiple regression (OLS) and the Akaike's information criterion (corrected for spatial autocorrelation) and derived indices to generate parsimonious models including multiple predictors. These analyses indicated that plant species richness increase primarily along with increasing patch area and decreasing disturbance, but also detected secondary effects of other factors when dispersal was considered. While the number of species with potential long-distance dispersal tended to increase in more isolated patches of areas with greater precipitation and lithological heterogeneity (e.g. highlands at the valley edges), the number of species with short-distance dispersal increased towards drier and less lithologically complex zones with shorter between-patch distances (e.g. central lowlands). Beyond emphasizing the need to consider dispersal in fragmentation studies, our results show that woody plant species richness would be favoured by actions that increase patch area and reduce anthropogenic disturbances particularly in lowland forests.     

Short-term browsing by roe deer has little effect on survival and growth of sessile oak seedlings

We studied the effect of roe deer (Capreolus capreolus) browsing on the growth and mortality of sessile oak (Quercus petraea) seedlings. First, we measured the height and basal diameter, and counted the number of shoots of the 849 seedlings of an experimental plantation, 60 of which were individually protected by fencing. We then introduced two tame roe deer females every day for three weeks from late May to mid-June. Measurements and counts were repeated after roe deer removal, then once again the following winter. We found no significant effect of browsing on mortality, total height, or basal diameter of the seedlings. However, the browsed seedlings exhibited a lower number of shoots despite regrowth observed after roe deer removal. We propose to use the tolerance of oak to deer browsing to promote natural and inexpensive methods, to protect oak during regeneration, such as the spatial arrangement of plants or the use of slash or surrounding vegetation in favourable environments.     

Effects of deer and mice on seedling survival in a temperate mixed forest of Japan

To detect the magnitude of indirect positive effects of deer and mice on seedling survival of some woody species in a Japanese temperate forest, we analyzed the data from an earlier field experiment using a hierarchical Bayesian approach. The forest studied was inhabited by sika deer (Cervus nippon) and mice (Apodemus spp.), and the floor was covered with dwarf bamboo, Sasa nipponica, which negatively affected tree seedlings and was in turn negatively affected by deer and mice. The field experiment was designed as the combination of exclusion or removal of these factors: deer, mice and dwarf bamboo. A hierarchical Bayesian model was constructed and the parameters were estimated by the Markov chain Monte Carlo method. The model successfully showed the indirect positive effects of deer and mice, which improved the survival rate of five cohorts of the seedlings of three tree species by ameliorating the direct negative effect of dwarf bamboo. The cohorts studied were formed by the seedlings of Abies homolepis that emerged in 1997 and 2002, those of Fraxinus lanuginosa f. serrata that emerged in 1998 and 2002, and those of Fagus crenata that emerged in 1999. The positive indirect effect was especially large in F. crenata, which is known to be severely affected by dwarf bamboo. The total effect of deer was shown to be positive for all cohorts except the A. homolepis cohort that emerged in 1997, at which time the dwarf bamboo had not yet fully recovered from the browsing pressure of deer. The total effect of mice was shown to be positive for all of the cohorts. We conclude that these positive effects were due to the large negative effect of dwarf bamboo on the seedlings.     

Early-stage impacts of sika deer on structure and function of the soil microbial food webs in a temperate forest: A large-scale experiment

Modification of forest vegetation caused by an overabundance of mammalian herbivores has been reported in temperate and subarctic regions all over the world. However, the indirect effects of these herbivores on the structure and functions of soil decomposer systems are not fully understood, especially in temperate forests. We investigated the early effects of sika deer invasion on soil decomposer systems in a Japanese temperate forest using two large-scale experimental enclosures with low and high densities of deer (LD: 25 ha, 4 deer km⁻²; HD: 6.25 ha, 16 deer km⁻²) including control plots without deer (WD). Three years after deer introduction the understory cover of dwarf bamboo (Sasa nipponica) declined due to deer browsing in both enclosures. At the same time, measurements were made of the soil microbial community, soil nematode community, soil nitrogen (N) mineralization rate, and carbon (C) and N content in dwarf bamboo leaves. In LD, soil microbial biomass was lower from WD, probably due to the decrease of fresh aboveground litter from dwarf bamboo. Surprisingly, there were no cascading effects on total abundance of soil nematodes and soil total N mineralization potential which were unaffected by deer in the LD treatment, while soil NH₄⁺-N content was lower and soil nematode community structure was different (abundance of 4 families was higher and that of 3 families was lower, but the functional structure was not different) from WD. Specifically, the responses to deer introduction varied between microbes and nematodes, and the change of balance in the microbial food webs may have altered N mineralization processes. In contrast, in the HD treatment, all the variables measured were not significantly different from those of WD treatment. Intensive browsing by deer may have cancelled out the effects of the decrease in aboveground litter input on the soil decomposer systems through other pathways, such as a transitory increase in belowground litter input caused by induced changes in allocation patterns of bamboo. No changes in total N mineralization potential, leaf N, and composition of understory vegetation in both enclosures indicated that deer introduction did not facilitate nor retard N cycling regardless of deer density. This study showed that sika deer browsing can affect soil decomposer systems at an early stage of invasion even at low density, which contrasts with previous studies on the subject. Linking our findings of early-stage effects of deer on soil decomposer systems to longer-term dynamics of understory vegetation and tree regeneration will be needed to evaluate the adequacy of deer management practices with respect to the sustainability of soil nutrient supplies.     

Population structures and spatial patterns of two unpalatable Arisaema species(Araceae) with and without clonal reproduction in a riparian forest intensively grazed by Sika deer

General decline of understory cover can result from increased abundance of and foraging pressure by deer.But population size and degree of aggregation can increase for unpalatable understory plants that escape foraging pressure.Clonal reproduction can enable unpalatable plant species to increase their population sizes while trending toward spatially aggregated distributions.However,the details of the relationship between clonal reproduction in unpalatable plants and their dynamics under intensive deer herbivory are not clear.We compared the population structures and spatial patterns of two coexisting unpalatable plant species,Arisaema ovale(with clonal reproduction)and A.peninsulae(without clonal reproduction)in a riparian forest intensively grazed by Sika deer,and examined the null hypothesis that the extent of spatial aggregation and local population size would not differ between the clonal and non-clonal Arisaema species.In a 0.36-ha plot,A.ovale had a larger population size(1087 individuals)with a higher abundance ratio of small plants(p<0.01)than A.peninsulae(84 individuals).Analyses of spatial point processes showed that both populations were spatially aggregated(p<0.05).The spatial aggregation of A.peninsulae,however,became weaker than that of A.ovale,when we excluded one dense patch originating from irregular seed dispersion.These results,excluding the aggregated distribution observed in A.peninsulae,suggested a substantial contribution of clonal reproduction to the expansion of the local A.ovale population following intensive grazing by Sika deer.     

Initial vegetation recovery following a blowdown of a conifer plantation in monsoonal East Asia: Impacts of legacy retention, salvaging, site preparation, and weeding

All or a part of a sequence of forest practices (i.e., salvage logging, site preparation, planting crop trees, and weeding) has been implemented after natural disturbances for the rapid re-establishment of tree cover. Forest policies in Japan have recently changed from monocultural planting of coniferous crop trees to planting native broadleaved trees to restore forests and nurture local biodiversity following large windthrows. However, the effects of this new practice on preserving biodiversity, as well as the effects of legacy retention, have never been verified in Asia. Thus, the objective of our research was to compare the effects of legacy retention with plantation after salvaging on the initial stage of vegetation recovery in a blowdown area, specifically focusing on plant species diversity, the occurrence of alien species, and the composition of plant species. Following the analysis of our results, we finally describe appropriate practices to alter disturbed coniferous plantations to bring the species composition closer to that of the original natural mixed forests.A control (A, legacy retention) and three experimental treatment sites (B, salvage logged, site prepared, and Quercus crispula seedlings planted; C, same as B, but weeded once during the summer; and D, residual rows that emerged after establishing sites for planting) were prepared, and quadrats were set. Eleven indicators of the ground condition and the number of vascular plant species, including ferns, were quantified, and the number and abundance of residual and newly colonized plants of the main woody species were estimated.Our main findings were as follows: (1) in unsalvaged sites and residual rows, the diversity of plant species was poor, but a variety of plant species compositions were observed due to the heterogeneous conditions of the ground and ample residual plants; (2) in the planting site, many species appeared, but little variety of the species composition was observed due to the homogeneous condition of the ground and the destruction of residual plants; (3) a large number of alien species emerged in broad, unvegetated areas; (4) the impact of site preparation overwhelmed the impact of salvage logging on the initial recovery of plant species; and (5) to restore a natural mixed forest, a combination of legacy retention and plantation after salvaging would be the most appropriate.     

Comparison of fractal characteristics of species richness patterns among different plant taxonomic groups along an altitudinal gradient

This study was done using the non brown fractal model to quantify and compare the variations in the species richness of trees, shrubs, herbs and all plants along an altitudinal gradient and to characterize the dominating ecological processes that determine the variations. Two transects were sampled far away from any anthropogenic disturbances along the shady slopes of the Dongling mountains in Beijing, China. Both transects were continuous and 2 m wide, and every individual tree and shrub was recorded in each of them. Discrete quadrats of 1 m × 1 m were located along the transects A and B for estimation of the herb species richness along the altitudinal gradients. The level interval between the quadrats was 10 m and 25 m respectively. In this study, transects A and B were combined into one transect AB, and 40 m was selected as the optimal quadrat length along the altitudinal gradients for measuring the plant species richness patterns. Species richness in each quadrat was calculated using a program written in Matlab 6.0. Direct gradient analysis was used to describe the overall trends in the species richness of trees, shrubs, herbs and other plants with change in altitude, while the non-brown fractal model was used to detect more accurately their variations at various scales along the gradient. The model assumed that each class of ecological processes affecting the distribution of a variable could be represented by an independent spatial random function. Generally, ecological phenomena are determined not by a single ecological process but by multiple ones. These processes act on ecological patterns within their own spatial scales. In the non-brown fractal model, the spatial random functions are nested within a larger range of spatial scales. The relative contribution of the spatial random functions to the spatial variation of a variable is indicated by a weighting parameter that has to be greater than or equal to zero. In this paper, we reached the following results and conclusions. Firstly, the direct gradient method describes the general trends of trees, shrubs, herbs and all plants along the altitudinal gradient but is unable to provide further details on the altitudinal variations in the species richness. The non-brown fractal model brought out the altitudinal variations in the species richness of trees, shrubs and herbs at various scales and related them to the ecological processes. The sharp changes in the double-log variograms suggest that the non-brown fractal model is suitable for characterizing the altitudinal patterns in the species richness of trees, shrubs and herbs at various scales but is not appropriate for explaining the variations in the plant species richness, since no significant changes were found in the double-log variograms in this case. Secondly, for the trees, the double-log variogram was divided into two scale ranges (0–245 m and 245–570 m), with a fractal dimension of 1.83 and 1.10, respectively, implying that changes in the tree species richness were random at small scales (0–245 m) and almost linear at large scales (245–570 m) along the altitudinal gradients. This suggests that altitudinal variations in the tree species richness are dominated by short-range processes at small scales and by long-range processes at large scales. Thirdly, for shrubs and herbs, the double-log variograms exhibited three ranges (0–101 m, 125–298 m and 325–570 m), and the fractal dimensions were 1.78 and 1.97, 1.56 and 1.43, and 1.08 and 1.25, respectively. The results indicate that, as in the case of trees, species richness of shrubs and herbs are distributed randomly at small scales and change in a linear manner at large scales although variations in the herb species richness is less heterogeneous than shrub species richness at large scales. These results also indicate that species richness of shrubs and herbs change approximately like brown movement at middle scales. The results also suggest that altitudinal variations in the specie richness of shrubs and herbs are dominated by three ecological processes, short-range ecological processes at small scales, long-range ecological processes at large scales, and brown fractal processes at middle scales. Interestingly, comparisons of the variations in the species richness of shrubs and herbs reveal that shrubs and herbs present the same scale range in spatial variation in species richness but display different trends in species richness along the altitudinal gradient, i.e. the shrub species richness decreased with increasing elevation whereas the herb species richness peaked at the mid-high elevation. These patterns suggest that although the scales at which the main processes affect patterns in species richness are the same, the processes are completely different, or the processes are similar but the responses of the shrubs and herbs to the ecological processes are different. Finally, the plant species richness did not show any obvious pattern along the altitude gradient and maintained a constant fractal dimension across all scales, this is perhaps because the processes defining the patterns of plant species richness had similar weights and acted over closely related scales. __________ Translated from Acta Phytoecologica Sinica, 2005, 29(6): 901–909 [译自: 植物生态学报]     

竹类植物种质资源圃抗寒性竹种的选择

通过对江苏省林业科学研究院收集引种的竹类植物22属108个竹种(含变种或变型)生长情况和冻害程度调查,将该院竹类植物种质资源圃的竹种抗寒性划分成5个等级,提出了一批值得发展推广的抗寒性强和比较强的观赏、经济竹种;同时,对抗寒性一般及抗寒性弱和极弱的竹种的引种与栽培提出了较好的建议。     

Influences of climate,fire, grazing,and logging on woody species composition along an elevation gradient in the eastern Cascades,Washington

Across western North America, current ecosystem structure has been determined by historical interactions between climate, fire, livestock grazing, and logging. Climate change could substantially alter species abundance and composition, but the relative weight of the legacy of historical factors and projected future conditions in informing management objectives remains unresolved. We integrated land use histories with broad scale climatic factors to better understand how inland Pacific Northwest ecosystems may develop under projected climates. We measured vegetation structure and age distributions in five vegetation types (shrub steppe to subalpine forest) along an elevation gradient in the eastern Cascades of Washington. We quantitatively assessed compositional changes, and qualitatively summarized the environmental history (climate, fire and fire suppression, grazing, and logging) of each site. Little change was evident in woody species composition at the shrub steppe site. At the shrub steppe/forest ecotone, densities of drought-tolerant Artemisia tripartita and Pinus ponderosa increased. In the dry conifer, montane, and subalpine forest sites, increases in Pseudotsuga menziesii, Abies grandis, and Abies lasiocarpa, respectively, and decreases in Pinus ponderosa, Larix occidentalis, and Pinus contorta, respectively, have shifted species composition from fire and drought-tolerant species to shade-tolerant species. Fire suppression, grazing, and logging explain changes in species composition more clearly than climate variation does, although the relative influence of these factors varies with elevation. Furthermore, some of the observed changes in composition are opposite what we expect would be most suited to projected future climates. Natural resource managers need to recognize that the current state of an ecosystem reflects historical land uses, and that contemporary management actions can have long-term effects on ecosystem structure. Understanding the processes that generated an ecosystem's current structure will lead to more informed management decisions to effectively respond to projected climate changes.     

Effects of uprooting tree on herbaceous species diversity, woody species regeneration status and soil physical characteristics in a temperate mixed forest of Iran

We conducted a study to examine the pattern of development of herbaceous plant species, woody species regeneration and soil physical characteristics after tree uprooting in 20-ha areas of Experimental Forest Station of Tarbiat Modares University located in a temperate forest of Mazandaran province in the north of Iran. Soil bulk density, soil texture and moisture from pit and mound (PM) were measured in the laboratory. Results show that the soil bulk density was most in soil deeper layers at mound top, and the soil moisture content was most in soil deeper layers at Pit bottom. Our study supports that the micro-topography of PM (pit and mound) topography will create a mosaic of environmental conditions. This environmental heterogeneity could be responsible for the diversity of herbaceous plant species and regeneration of woody species. It is recommend that the fallen trees with PM structure should remain in the protected area without clearing as the best option for forest restoration. This information can be useful for forest management that attempts to emulate natural processes.     

Regeneration strategies in a temperate hardwood floodplain forest of the Upper Rhine: sexual versus vegetative reproduction of woody species

The regeneration mechanisms of woody species in the hardwood floodplain forest of the Upper Rhine are not well known, although they are of primary importance for future forest development. To gain a better understanding of the balance between sexual and asexual regeneration strategies and to assess the role of the seed bank in regeneration, the similarities in species composition and abundance of four fundamental compartments involved in regeneration (the seed rain, the seed bank, the recruits and the canopy) were compared in three hardwood forest stands with different flooding conditions. The results show that the floristic composition of the recruits is very similar to that of the canopy, whereas the composition of the seed bank is very dissimilar to the latter and comprises very few hardwood species. However, some species such as Fraxinus excelsior, Carpinus betulus and Acer pseudoplatanus which are very rare in the seed bank release a high number of diaspores, and seedlings of these species germinate abundantly in the field in the year following seed release. Moreover, the germinating seeds originate from the litter layer and not from the soil itself. This shows that most woody species regenerating by seed in the hardwood Rhine forest build transient seed banks and that the role of persistent seed banks for regeneration is very limited. Furthermore, it appears that many woody species have developed strategies favouring vegetative propagation for their regeneration, particularly understorey species, such as Cornus sanguinea and Prunus padus. As vegetatively grown individuals better withstand prolonged inundation in their early life stages than seedlings, species relying on vegetative regeneration strategies might be advantaged by regular and prolonged flooding of the Rhine forest over species regenerating only by seeds.     

Forage production of woody fodder species and herbaceous vegetation in a silvopastoral system in northern Greece

Introduction of woody plants in silvopastoral systems could be an appropriate land-use for the poor sandy soils of the Mediterranean semiarid zone. Forage production of four woody fodder species and herbaceous vegetation in relation to plant spacing and animal (sheep) grazing was studied in a silvopastoral system on such a site in Macedonia, northern Greece. The woody fodder species tested were Robinia pseudoacacia L., Gleditsia triacanthos L., Amorpha fruticosa L. and Morus alba L. They were planted at the spacings 1.5 × 1.5 m,2.5 × 2.5 m and 3.5 × 3.5 m and kept in a shrubby form by topping. They were grazed by sheep (stocking rate of 1.1 sheep/ha/year) in early July and late August of 1992, 1993 and 1994. Among the species Robinia yielded the highest forage production (394 kg/ha). The spacing 1.5 × 1.5 m had the highest (P ≥ 0.05) forage production per area unit (280 kg/ha) while the spacing 3.5 × 3.5 m had the highest forage production per plant (91.8 g/plant). No competition was observed between the woody and the herbaceous plants in the various spacings probably due to topping of the woody species every winter. Percent utilisation of the native herbage dropped as forage of the woody species increased. This revised version was published online in June 2006 with corrections to the Cover Date.     

Structure and regeneration dynamics of dominant tree species along altitudinal gradient in a dry valley slopes of the Bhutan Himalaya

The aim of our study was to characterize the structure and regeneration dynamics of dominant tree species along altitudinal gradient in dry valley slopes of the Bhutan Himalaya. In the typical dry valley slopes of west-central Bhutan, we analyzed forest stratification, tree crown projection, and population structure from lower dry valley bottom (1520 m a.s.l.) to upper humid ridge top (3370 m a.s.l.) across five major forest types: i.e. (1) dry pine forest with Pinus roxburghii as a dominant species; (2) mixed broad-leaved forest with Quercus lanata (evergreen), Quercus griffithii, (deciduous), and Rhododendron arboreum (evergreen) as major canopy species; (3) evergreen broad-leaved forest dominated by Quercus semecarpifolia, Quercus glauca and Quercus oxyodon; (4) cool conifer forest dominated by Abies densa and Tsuga dumosa; (5) cold conifer forest with A. densa and Juniperus recurva as dominants. In general, regeneration pattern of major dominant species shifted from inverse-J (lower altitudes), to sporadic (mid-altitudes), and to uni-modal type (upper altitudes) corresponding to three regeneration trends: (1) invasive at the lower, warm, dry forest under relatively strong human disturbances; (2) stable/balanced at the mid-altitude, in a relatively stable, mature moist evergreen broad-leaved forest with gap regeneration; and (3) poor/low regeneration at the upper, cool, humid conifer forest with a continuously cattle-grazed understory. Overall, regeneration patterns were balanced in all forest types, however care should be taken to prevent excessive exploitation of dry pine forest of lower valley bottom, and to control cattle grazing at higher altitudes. Our findings can be incorporated into management plans for sustainable management and conservation of mountain forests.