Productivity of temperate broad-leaved forest stands differing in tree species diversity

? Understanding the effects of tree species diversity on biomass and production of forests is fundamental for carbon sequestration strategies, particularly in the perspective of the current climate change. However, the diversity-productivity relationship in old-growth forests is not well understood.

? We quantified biomass and above-ground production in nine forest stands with increasing tree species diversity from monocultures of beech to stands consisting of up to five deciduous tree species (Fagus sylvatica, Fraxinus excelsior, Tilia spp., Carpinus betulus, Acer spp.) to examine (a) if mixed stands are more productive than monospecific stands, (b) how tree species differ in the productivity of stem wood, leaves and fruits, and (c) if beech productivity increases with tree diversity due to lower intraspecific competition and complementary resource use.

? Total above-ground biomass and wood production decreased with increasing tree species diversity. In Fagus and Fraxinus, the basal area-related wood productivity exceeded those of the co-occurring tree species, while Tilia had the highest leaf productivity. Fagus trees showed no elevated production per basal area in the mixed stands.

? We found no evidence of complementary resource use associated with biomass production. We conclude that above-ground productivity of old-growth temperate deciduous forests depend more on tree species-specific traits than on tree diversity itself.

     

Soil seed banks in four 22-year-old plantations in South China: Implications for restoration

To better understand the potentials of the soil seed banks in facilitating succession towards a more natural forest of native tree species, we quantified the size and composition of the soil seed banks in established plantations in South China. The seed banks were from four typical 22-year-old plantations, i.e., legume, mixed-conifer, mixed-native, and Eucalyptus overstory species. Species diversity in the seed banks was low, and the vegetation species differed from those found in the seed bank in each plantation. A total of 1211 seedlings belonging to eight species emerged in a seedling germination assay, among which Cyrtococcum patens was most abundant. All species detected were shrubs and herbs, and no viable indigenous tree seeds were found in soil samples. Size and species composition of the seed banks might be related to the overstory species compositions of the established plantations. The seed bank density in soils was highest in the mixed-conifer plantation followed by Eucalyptus, mixed-native, and legume plantations. Species richness among the seed banks of plantations was ranked as follows: Eucalyptus > mixed-conifer > mixed-native = legume. The results indicated that the soil seed banks of the current plantations are ineffective in regenerating the former communities after human disturbances. Particularly, the absence of indigenous tree species seeds in the seed banks would limit regeneration and probably contribute to arrested succession at the pioneer community stage. It would appear from these data that the soil seed banks under the current plantations should not be considered as a useful tool leading the succession to more natural stages. Introduction of target indigenous species by artificial seeding or seedling planting should be considered to accelerate forest regeneration.     

Through droughts and hurricanes: Tree mortality,forest structure,and biomass production in a coastal swamp targeted for restoration in the Mississippi River Deltaic Plain

Coastal swamps are among the rapidly vanishing wetland habitats in Louisiana due to accelerated sea-level rise and hydrological alterations that alter the natural flooding regime. In particular, the swamp forests of Lake Maurepas, Louisiana, have degraded considerably, and research regarding their condition might suggest approaches for their restoration. We measured forest structure, species composition, tree mortality, annual aboveground net primary production (ANPP) of woody species, and aboveground biomass allocation to leaf litter and wood, and soil strength at forty study plots within the Lake Maurepas basin over 5 years to evaluate the current condition of this coastal forested wetland. Local measures of salinity and regional measures of flooding were used to predict ANPP and aboveground biomass allocation. The 5-year study period included an intense drought as well as years characterized by hurricane-induced flooding. The forty study plots could be divided into four distinct habitat clusters based on standing biomass, structural variables, and salinity. The majority of the plots were co-dominated by Taxodium distichum and Nyssa aquatica. Acer rubrum var. drummondii and Fraxinus pennsylvanica were common mid-story species throughout the western and southern parts of the study area, while Salix nigra, Morella cerifera, and Triadica sebiferum were more important at the more degraded plots in the eastern part of the basin. Annual mean soil salinity reached unprecedented level (2–5 psu) during the drought and cumulative tree mortality reached up to 85% in areas characterized by frequent saltwater intrusions. The ANPP was higher during the drought period in 2000–2001 than during subsequent years, and was dominated by T. distichum. At most sampling plots, litter production exceeded wood production annually. A negative correlation between aboveground biomass allocation to litter and flooding indicated that biomass allocation shifted from litter toward wood during wet years. Overall, the majority of the plots sampled produced less than 400 g m⁻² yr⁻¹ of aboveground biomass annually due to the interacting negative effects of saltwater intrusion and prolonged flooding with nutrient-poor water. Reintroduction of Mississippi River water to the Maurepas system has the potential to benefit these swamps greatly by restoring a greater flow of nutrients, sediments, and fresh water through the wetlands. The historically slow (i.e., multi-decadal) process of swamp deterioration was greatly sped by low salinity (i.e., 2–5 psu) saltwater intrusions during a drought in 1999–2000. The majority of the coastal swamps in the Pontchartrain Basin are deteriorating, and most of this swamp area will be lost to open water in the foreseeable future if no restoration action is taken.     

Gap partitioning among temperate tree species across a regional soil gradient in windstorm-disturbed forests

Canopy closure and soil characteristics are commonly used to explain regeneration distribution at local and regional scales, although very few studies take both factors into account. The combination of environmental variables defined at broad and local scales is necessary to provide regeneration distribution models with a small resolution (tree scale) that are valid on a large spatial scale (regional scale). Our aim was to quantify how gap partitioning among tree species at the seedling stage varies across large soil and stand type gradients. Regeneration inventories performed 5 years after gap creation were used to analyse the combined effects of soil type, stand type, and position within canopy gaps on the regeneration development of eight western European broadleaved species: Acer campestre, Acer pseudoplatanus, Betulapendula, Carpinusbetulus, Fagussylvatica, Fraxinusexcelsior, Quercus sp., and Salixcaprea. A clear pattern of gap partitioning among the eight species was observed. All species had higher density at the gap edge except birch and willow showing the highest presence in gap centres. For all species, the probability of presence of tall seedlings (height > 0.5 m) increased from gap edge to gap centre. Small seedlings presented the opposite trend except birch and willow. Soil pH influenced probability of presence for each species, but did not affect the pattern of gap partitioning among species. Both local (location within the gap) and regional (soil pH and stand type) scale factors affect recruitment distribution and are thus necessary to predict seedling distribution. The models developed may be used to determine the optimal gap size in order to obtain a given species composition according to soil and stand type conditions.     

Twentieth-century decline of large-diameter trees in Yosemite National Park,California, USA

Studies of forest change in western North America often focus on increased densities of small-diameter trees rather than on changes in the large tree component. Large trees generally have lower rates of mortality than small trees and are more resilient to climate change, but these assumptions have rarely been examined in long-term studies. We combined data from 655 historical (1932–1936) and 210 modern (1988–1999) vegetation plots to examine changes in density of large-diameter trees in Yosemite National Park (3027 km²). We tested the assumption of stability for large-diameter trees, as both individual species and communities of large-diameter trees. Between the 1930s and 1990s, large-diameter tree density in Yosemite declined 24%. Although the decrease was apparent in all forest types, declines were greatest in subalpine and upper montane forests (57.0% of park area), and least in lower montane forests (15.3% of park area). Large-diameter tree densities of 11 species declined while only 3 species increased. Four general patterns emerged: (1) Pinus albicaulis, Quercus chrysolepis, and Quercus kelloggii had increases in density of large-diameter trees occur throughout their ranges; (2) Pinus jeffreyi, Pinus lambertiana, and Pinus ponderosa, had disproportionately larger decreases in large-diameter tree densities in lower-elevation portions of their ranges; (3) Abies concolor and Pinus contorta, had approximately uniform decreases in large-diameter trees throughout their elevational ranges; and (4) Abies magnifica, Calocedrus decurrens, Juniperus occidentalis, Pinus monticola, Pseudotsuga menziesii, and Tsuga mertensiana displayed little or no change in large-diameter tree densities. In Pinus ponderosa–Calocedrus decurrens forests, modern large-diameter tree densities were equivalent whether or not plots had burned since 1936. However, in unburned plots, the large-diameter trees were predominantly A. concolor, C. decurrens, and Q. chrysolepis, whereas P. ponderosa dominated the large-diameter component of burned plots. Densities of large-diameter P. ponderosa were 8.1 trees ha⁻¹ in plots that had experienced fire, but only 0.5 trees ha⁻¹ in plots that remained unburned.     

Experimental test of the impacts of feral hogs on forest dynamics and processes in the southeastern US

The foraging activities of nonindigenous feral hogs (Sus scrofa) create widespread, conspicuous soil disturbances. Hogs may impact forest regeneration dynamics through both direct effects, such as consumption of seeds, or indirectly via changes in disturbance frequency or intensity. Because they incorporate litter and live plant material into the soil, hogs may also influence ground cover and soil nutrient concentrations. We investigated the impacts of exotic feral hogs in a mixed pine-hardwood forest in the Big Thicket National Preserve (Texas, USA) where they are abundant. We established sixteen 10 m × 10 m plots and fenced eight of them to exclude feral hogs for 7 years. Excluding hogs increased the diversity of woody plants in the understory. Large seeded (>250 mg) species known to be preferred forage of feral hogs all responded positively to hog exclusion, thus consumption of Carya (hickory nuts), Quercus (acorns), and Nyssa seeds (tupelo) by hogs may be causing this pattern. The only exotic woody species, Sapium sebiferum (Chinese tallow tree), was more than twice as abundant with hogs present, perhaps as a response to increased disturbance. Hogs increased the amount of bare soil by decreasing the amounts of plant cover and surface litter. Plots with hogs present had lower soil C:N, possibly due to accelerated rates of nitrogen mineralization. These results demonstrate that hogs may influence future overstory composition and reduce tree diversity in this forest. Management of hogs may be desirable in this and other forests where large-seeded species are an important component of the ecosystem. Further, by accelerating litter breakdown and elevating nitrogen in the soil, hogs have the potential to impact local vegetation composition via nitrogen inputs as well.     

Herb-layer diversity in deciduous forests: Raised by tree richness or beaten by beech?

Where natural production capacity permits, modern silvicultural management in Central Europe frequently aims at the development of mixed broadleaved stands, instead of pure European beech (Fagus sylvatica) stands. It is crucial to study the effects of these tree-layer diversity variations on herb-layer vegetation, since herb-layer vegetation contributes significantly to ecosystem functioning in forests. In Hainich National Park (Thuringia, Germany), we conducted observational research in deciduous stands to investigate whether herb-layer diversity was related to canopy-layer diversity, and to ascertain possible causal mechanisms. We found that herb-layer vegetation of deciduous forest stands rich in canopy species appeared to be more diverse than herb-layer vegetation of beech-dominated stands. We surmise that herbaceous understorey diversity was indirectly influenced by canopy tree species through the medium of the altered environmental factors soil pH and litter layer thickness. Apparently, lower beech proportion had a more profound effect than the number of secondary tree species. There were no correlations between herb-layer diversity and light transmissibility of the canopy layer, indicating that the light factor was not crucial for herb-layer diversity. At least for the Hainich research sites, our results indicated that small-scale light and soil heterogeneity is insignificant for herb-layer diversity. We found several herb-layer species whose occurrence was particularly correlated with tree-layer diversity and environmental factors. Remarkably, all species positively correlated with soil pH were important for the phytosociological classification of the research sites. Beech-dominated research sites showed high tree-layer volumes, whereas research sites with high tree-layer diversity tended to feature lower tree-layer volumes. These findings could be the result of differing former silvicultural systems and varying soil clay contents affecting tree species composition. In contrast, herb-layer biomass was positively correlated with tree-layer diversity. Herb-layer productivity might be promoted in more diverse research sites by increased nutrient supply and base saturation. It is also possible that greater beech proportion interfered with herb-layer productivity. However, herb-layer biomass was also positively correlated with herb-layer diversity. Hence, our study hints that positive diversity-functioning relationships might occur in the herb-layer of the deciduous forest under investigation.     

Intra and inter-annual variation in seed rain in a secondary dry tropical forest excluded from chronic disturbance

Domestic livestock grazing and firewood extraction are chronic and pervasive disturbances in dry secondary forest that reduce diversity and modify composition. Exclosure of such disturbances may allow the reestablishment of original diversity. To investigate potential vegetation recovery following 30 years of perturbation, seed rain was evaluated monthly during 3 years in 96 0.78 m² seed traps located in eight exclosures established in 2006 in a species-poor secondary dry forest in central Mexico. Twelve additional seed traps were located outside the exclosures in 2010 for comparison purposes. Baseline diversity of seed sources (trees, ?5 cm DBH) was recorded to compare standing tree composition with seed rain. Also, baseline diversity of potential primary (birds) and secondary (ants) dispersal vectors were registered to infer current plant-animal interactions. Total seed rain over 3 years within the exclosures comprised 180,375 seeds from 61 woody species, an average of 1.12 seeds/m²/month. Half of these species (31) and 95% of the seeds falling within the exclosures belonged to trees typical of perturbed sites. Sixty percent of all recorded seeds belonged to an early-successional, bird-dispersed shrub (Hamelia patens Jacq., Rubiaceae). Seed rain at one non-excluded plot was dominated by the wind-dispersed early-successional tree Heliocarpus sp. (Tiliaceae). Richness and seed density increased significantly in the second year of exclusion; however, by the third year, richness remained constant while seed density decreased. Seed rain was richer and more abundant during the dry season and was dominated by wind-dispersed species. Phenology, successional status and the dispersal modes of woody species explained most variance in richness and seed density of seed rain in early-successional dry forest. Seeds of early-successional species were dispersed in equal measure by abiotic and biotic vectors, whereas biotic vectors dispersed a higher density of late-successional seeds. The amount and timing of fall of abiotically dispersed seeds of the three most abundant species in the standing tree community may prolong their current domination of the landscape. However, even when richness in the seed rain within exclosures remained constant through time, composition was gradually shifting towards animal-dispersed species which may start a positive feedback producing an increase in animal visitation and a further increase of richness.     

Prescribed burning and understory composition in a temperate deciduous forest,Ohio, USA

Since the advent of widespread suppression in the mid-20th century, fire has been relatively rare in deciduous forests of the eastern United States. However, widespread prescribed burning has recently been proposed as a management tool to favor oak (Quercus spp.) regeneration. To examine the potential effects of fire introduction on the understory community, we experimentally burned small plots and simulated aspects of fire at a forested site in southeastern Ohio. Treatments included two burn intensities, litter removal, increased soil pH, and a control. Treatments were arranged in a randomized block design in two landscape positions (dry upland and moist lowland) and two canopy conditions (gap, no gap). Post-fire vegetation was identified to species, and stems were counted 1, 3, and 14 months after burning. Community composition was more strongly affected by fire in upland plots than in lowlands, but was not affected by canopy openness. Both cool and hot burns reduced post-fire seedling emergence of Acer rubrum, a common overstory tree. Hot burns facilitated germination of Vitis spp., Rhus glabra, and Phytolacca americana, species common in disturbed habitats, and increased graminoid abundance. Cool burns and litter removal facilitated germination of Erechtites hieracifolia and Liriodendron tulipifera suggesting that litter removal is the mechanism by which fire favors colonization. These results suggest that fire applied frequently in the Central Hardwoods Region would cause compositional shifts to graminoids and disturbance-adapted forbs by increasing germination from the seed bank. Fire did not favor species with dormant underground buds, as studies in other ecosystems would suggest. Vegetational responses were noticeably weaker in the second year after burning, indicating that a single fire has only a short-term effect.     

Understory vegetation in old-growth and second-growth Tsuga canadensis forests in western Massachusetts

We compared the understory communities (herbs, shrubs, and tree seedlings and saplings) of old-growth and second-growth eastern hemlock forests (Tsuga canadensis) in western Massachusetts, USA. Second-growth hemlock forests originated following clear-cut logging in the late 1800s and were 108–136 years old at the time of sampling. Old-growth hemlock forests contained total ground cover of herbaceous and shrub species that was approximately 4 times greater than in second-growth forests (4.02 ± 0.41%/m² versus 1.06 ± 0.47%/m²) and supported greater overall species richness and diversity. In addition, seedling and sapling densities were greater in old-growth stands compared to second-growth stands and the composition of these layers was positively correlated with overstory species composition (Mantel tests, r > 0.26, P < 0.05) highlighting the strong positive neighborhood effects in these systems. Ordination of study site understory species composition identified a strong gradient in community composition from second-growth to old-growth stands. Vector overlays of environmental and forest structural variables indicated that these gradients were related to differences in overstory tree density, nitrogen availability, and coarse woody debris characteristics among hemlock stands. These relationships suggest that differences in resource availability (e.g., light, moisture, and nutrients) and microhabitat heterogeneity between old-growth and second-growth stands were likely driving these compositional patterns. Interestingly, several common forest understory plants, including Aralia nudicaulis, Dryopteris intermedia, and Viburnum alnifolium, were significant indicator species for old-growth hemlock stands, highlighting the lasting legacy of past land use on the reestablishment and growth of these common species within second-growth areas. The return of old-growth understory conditions to these second-growth areas will largely be dependent on disturbance and self-thinning mediated changes in overstory structure, resource availability, and microhabitat heterogeneity.     

Disruption of <Emphasis Type="Italic">Juniperus thurifera</Emphasis> woodland structure in its northwestern geographical range: potential drivers and limiting factors

Enhancement of Juniperus thurifera recruitment and colonisation by oak and pine species has been related at the local level to changes in livestock pressure. We used forest inventory data from Castilla y León Autonomous Region (Central Spain), an area comprising 34% of the world range of J. thurifera, to assess whether this process is occurring at a larger scale. We compared tree composition and density in a set of 659 permanent plots over a 10-year period. Logistic models and redundancy analysis were used to assess the effect on this process of parameters such as livestock pressure, propagule availability and climatic conditions. Between 1992 and 2002, juniper woodlands became denser (1.31% juniper stem year⁻¹) and tree diversity increased due to rapid colonisation by oaks and pines (2.21% occupied plots year⁻¹). In addition, the presence of juniper increased in other types of forests at a moderate rate (0.6% y⁻¹). Thus, we observed both a disruption of the borders between current forest types and a generalised increase in α-diversity of tree species. The seed source was the main factor explaining colonisation rate, suggesting that the pace of colonisation is critically constrained by the spatial configuration of the landscape and the local propagule availability of the colonising species. If the current colonisation trends continue, monospecific juniper woodlands will become very scarce by the end of the twenty-first century.     

The impact of atmospheric deposition and climate on forest growth in European monitoring plots: An individual tree growth model

In the climate change discussion, the possibility of carbon sequestration of forests plays an important role. Therefore, research on the effects of environmental changes on net primary productivity is interesting. In this study we investigated the influence of changing temperature, precipitation and deposition of sulphur and nitrogen compounds on forest growth. The database consisted of 654 plots of the European intensive monitoring program (Level II plots) with 5-year growth data for the period 1994–1999. Among these 654 plots only 382 plots in 18 European countries met the requirements necessary to be used in our analysis. Our analysis was done for common beech (Fagus sylvatica), oak (Quercus petraea and Q. robur), Scots pine (Pinus sylvestris) and Norway spruce (Picea abies). We developed an individual tree growth model with measured basal area increment of each individual tree as responding growth factor and tree size (diameter at breast height), tree competition (basal area of larger trees and stand density index), site factors (soil C/N ratio, temperature), and environmental factors (temperature change compared to long-term average, nitrogen and sulphur deposition) as influencing parameters. Using a mixed model approach, all models for the tree species show a high goodness of fit with Pseudo-R² between 0.33 and 0.44. Diameter at breast height and basal area of larger trees were highly influential variables in all models. Increasing temperature shows a positive effect on growth for all species except Norway spruce. Nitrogen deposition shows a positive impact on growth for all four species. This influence was significant with p < 0.05 for all species except common beech. For beech the effect was nearly significant (p = 0.077). An increase of 1 kg N ha⁻¹ yr⁻¹ corresponds to an increase in basal area increment between 1.20% and 1.49% depending on species. Considering an average total carbon uptake for European forests near 1730 kg per hectare and year, this implies an estimated sequestration of approximately 21–26 kg carbon per kg nitrogen deposition.     

Stand characteristics and downed woody debris accumulations associated with a mountain pine beetle (Dendroctonus ponderosae Hopkins) outbreak in Colorado

Lodgepole pine (Pinus contorta Dougl. ex Loud.)-dominated ecosystems in north-central Colorado are undergoing rapid and drastic changes associated with overstory tree mortality from a current mountain pine beetle (Dendroctonus ponderosae Hopkins) outbreak. To characterize stand characteristics and downed woody debris loads during the first 7 years of the outbreak, 221 plots (0.02 ha) were randomly established in infested and uninfested stands distributed across the Arapaho National Forest, Colorado. Mountain pine beetle initially attacked stands with higher lodgepole pine basal area, and lower density and basal area of Engelmann spruce (Picea engelmannii [Parry]), and subalpine fir (Abies lasiocarpa (Hook.) Nutt. var. lasiocarpa) compared to uninfested plots. Mountain pine beetle-affected stands had reduced total and lodgepole pine stocking and quadratic mean diameter. The density and basal area of live overstory lodgepole declined by 62% and 71% in infested plots, respectively. The mean diameter of live lodgepole pine was 53% lower than pre-outbreak in infested plots. Downed woody debris loads did not differ between uninfested plots and plots currently infested at the time of sampling to 3 or 4–7 years after initial infestation, but the projected downed coarse wood accumulations when 80% of the mountain pine beetle-killed trees fall indicated a fourfold increase. Depth of the litter layer and maximum height of grass and herbaceous vegetation were greater 4–7 years after initial infestation compared to uninfested plots, though understory plant percent cover was not different. Seedling and sapling density of all species combined was higher in uninfested plots but there was no difference between infested and uninfested plots for lodgepole pine alone. For trees ≥2.5 cm in diameter at breast height, the density of live lodgepole pine trees in mountain pine beetle-affected stands was higher than Engelmann spruce, subalpine fir, and aspen, (Populus tremuloides Michx.), in diameter classes comprised of trees from 2.5 cm to 30 cm in diameter, suggesting that lodgepole pine will remain as a dominant overstory tree after the bark beetle outbreak.     

Mixed litter decomposition in a managed Missouri Ozark forest ecosystem

We monitored the decomposition of mixed leaf litter (Quercus spp., Carya spp., and Pinusechinata) in a Missouri Ozark forest eight years after experimental harvest. Leaf litter mass losses and changes in carbon chemistry (extractive, acid soluble, and acid insoluble fractions) were measured over 32 months in field incubations to determine the effects of litter composition and stand manipulation on decomposition and nitrogen (N) concentration in the remaining litter. The decay (k) rate over this period ranged between 0.39 (±0.010) and 0.51 (±0.002) year⁻¹ for oak, oak–hickory, and oak–pine litter. There were significant main effects of stand manipulation (p = 0.03) and litter type (p < 0.01) on decay. Mass losses of oak and oak–hickory litter were 7% (p = 0.02) and 4% (p = 0.04) higher on harvested stands than controls, respectively. Mass loss of oak–hickory litter was 3% faster than oak–pine (p = 0.03) and 6% faster than oak (p = 0.02) litter on control stands, whereas the oak–hickory litter mass loss was 5% higher than oak litter on harvested stands (p = 0.01). The decay (k) rate had a linear relationship with initial leaf litter nitrogen content and lignin-to-N ratio. The nitrogen concentration in remaining litter had a nonlinear relationship to cumulative mass loss suggesting an exogenous source of N. In summary, this study demonstrated significant effects of timber harvest and litter mixtures on decomposition and N dynamics in a managed Missouri Ozark forest.     

Changes in size of soil seed bank in Robinia pseudoacacia L. (Leguminosae), an exotic tall tree species in Japan: Impacts of stand growth and apicultural utilization

This study examined the factors that influence the size of a soil seed bank in Robinia pseudoacacia stands. We proposed two hypotheses: the amount of buried seeds of R. pseudoacacia is proportional to stand biomass rather than stand age, and apicultural utilization influences the size of the soil seed bank. R. pseudoacacia generally produces seeds with various degrees of physical dormancy. In addition, this tree is short-lived, and fecundity declines after 30–40 years in relation to tree vigor. However, R. pseudoacacia is intensively used in apiculture as an important honey source, and supply of honeybees by beekeepers could influence seed crops. We investigated 25 plantations in Hokkaido (Central, Hidaka, and Oshima) and in Nagano, Japan, where many naturally regenerated R. pseudoacacia stands occur. We found great variation in the size of the soil seed bank among stands, ranging from 13,757 to 6.4 seeds m⁻² per plot. A generalized linear mixed model revealed that both basal area (BA_Rp) and apicultural utilization best explained the size of the soil seed bank. Both the positive effect of BA_Rp and small contribution of stand age to the model implied that the soil seed bank is transitory, and will not persist for very long time. The large contribution of apicultural utilization to the size of the soil seed bank implied pollinator limitation under natural conditions. R. pseudoacacia often has large flower crops, and native pollinators can not keep up with the demand for pollination. Thus, the supply of honeybees by beekeepers should improve the pollination success of R. pseudoacacia. From these results, we conclude that we must consider stand history with respect to apicultural utilization if the land is harvested or a stand declines.     

Belowground competition in a broad-leaved temperate mixed forest: pattern analysis and experiments in a four-species stand

We investigated fine root biomass and distribution patterns in a species-rich temperate Carpinus–Quercus–Fagus–Tilia forest and searched for experimental evidence of symmetry or asymmetry in belowground competition. We conducted extensive root coring and applied the recently introduced in situ-root growth chamber technique for quantifying fine root growth under experimentally altered intra- and interspecific root neighbourhoods in the intact stand. In 75% of all soil cores, fine roots of more than two tree species were present indicating a broad overlap of the root systems of neighbouring trees. Quercus trees had more than ten times less fine root biomass in relation to aboveground biomass or productivity (stem growth) and a much higher leaf area index/root area index ratio than Carpinus, Fagus and Tilia trees. The root growth chamber experiments indicated a high belowground competitive ability of Fagus in interspecific interactions, but a low one of Quercus. We conclude that (1) interspecific root competition is ubiquitous in this mixed stand, (2) root competition between trees can be clearly asymmetric, and (3) tree species may be ranked according to their belowground competitive ability. Fagus was found to be the most successful species in belowground competition which matches with its superiority in aboveground competition in this forest community.     

Epiphyte diversity and biomass loads of canopy emergent trees in Chilean temperate rain forests: A neglected functional component

We document for the first time the epiphytic composition and biomass of canopy emergent trees from temperate, old-growth coastal rainforests of Chile (42°30′S). Through tree-climbing techniques, we accessed the crown of two large (c. 1 m trunk diameter, 25–30 m tall) individuals of Eucryphia cordifolia (Cunoniaceae) and one large Aextoxicon punctatum (Aextoxicaceae) to sample all epiphytes from the base to the treetop. Epiphytes, with the exception of the hemi-epiphytic tree Raukaua laetevirens (Araliaceae), were removed, weighed and subsamples dried to estimate total dry mass. We recorded 22 species of vascular epiphytes, and 22 genera of cryptogams, with at least 30 species of bryophytes, liverworts and lichens. The dominant vascular epiphytes were Fascicularia bicolor (Bromeliaceae), Raukaua laetevirens, Sarmienta repens (Gesneriaceae), and filmy ferns (Hymenophyllaceae). Epiphyte loads per tree ranged between 134 and 144 kg dry mass, with 60–70% water. The hemi-epiphytic tree R. laetevirens added between 1 and 2.6 t of dry mass to each host tree. A main component of epiphyte biomass, making 70% of the weight, was detritus and roots, while leaves, stems, and fronds made up the remaining 30%. Emergent trees hold a high proportion of the regional diversity of epiphytes: 33% of all flowering epiphytes, and 50% of all filmy ferns described for Chilean temperate forests. Dry epiphyte biomass associated only with the emergent E. cordifolia trees in coastal forests was estimated in 10 t/ha. Epiphyte biomass may store up to 300 l of water in each emergent tree, and add 40–150% of photosynthetic biomass to the tree crowns. Based on this evidence, epiphytes may play key but generally neglected roles in ecosystem carbon uptake, water storage, and nutrient cycling. Moreover, emergent trees represent nuclei of biodiversity and ecosystem functions distributed throughout mature forests. Forest management should recognize large trees as significant management units for the preservation of biodiversity and ecological functions.     

Positive association between understory species richness and a dominant shrub species (Corylus avellana) in a boreonemoral spruce forest

Old growth stands of boreonemoral spruce (Picea abies) forests frequently have a shrub layer dominated by hazel (Corylus avellana) – a species which is generally excluded in intensively managed forests due to clearcutting activities. We sampled understory species composition, richness and biomass, as well as environmental variables beneath these two species and also within forest ‘gaps’ in order to determine the effect of overstory species on understory vegetation. Species richness and biomass of herbaceous plants was significantly greater under Corylus compared with plots under Picea and in forest gaps. Indicator species analysis found that many species were significantly associated with Corylus. We found 45% of the total species found under woody plants occurred exclusively under Corylus. Light availability in spring and summer was higher in gaps than under forest cover but no difference was found between plots under Corylus and Picea. Hence, reductions in light availability cannot explain the differences in species composition. However, Ellenberg indicator values showed that more light demanding species were found under Corylus compared to Picea, but most light demanding species were found in gaps. The litter layer under Picea was three times thicker than under Corylus and this may be an important mechanism determining differences in understory composition and richness between the woody species. The presence of Corylus is an important factor enhancing local diversity and small-scale species variation within coniferous stands. Hence, management should maintain areas of Corylus shrubs to maintain understory species diversity in boreal forests.     

Native woody species regeneration in exotic tree plantations at Munessa-Shashemene Forest,southern Ethiopia

Regeneration of native woody species was studied in the plantations and the adjacent natural forest at Munessa-Shashemene Forest Project Area, Ethiopia. The aim of the study was to test the hypothesis that tree plantations foster regeneration of native woody species. A total of 60 plots, having 10 × 10 m area each, were studied in monoculture plantations of 4 exotic species (Cupressus lusitanica, Eucalyptus globulus, E. saligna, Pinus patula) and an adjacent natural forest. Ages of the plantations ranged between 9 and 28 years. Soil seed bank analysis was also undertaken from soil samples collected in each of the 60 plots to examine the similarity between the soil seed flora and aboveground vegetation. A total of 56 naturally regenerated woody species were recorded beneath all plantation stands with densities ranging between 2300 and 18650 individuals / ha in different stands. There was a significant difference among plantation stands with regard to understorey density (standard deviation: 4836 ± 1341). Vegetation diversity was assessed through analyses of floristic composition, species richness and abundance. Generally, seedling populations were the most abundant components of the regeneration in most of the plantation stands, forming 68 % of the total regeneration count in all stands. A total of 77 plant species represented by 44 herbs, 13 woody species, 8 grasses and 12 unidentified species were recorded in the soil seed bank from all stands. Similarity between the soil seed bank and aboveground flora was very low implying that the role of soil seed banks is negligible rather dispersal plays an important role in the process of regeneration. These results support the concept that forest plantations can foster the regeneration of native woody species, thereby increasing biological diversity, provided that there are seed sources in the vicinity of the plantations.