Composition and structure of natural mixed-oak stands in northern and central Portugal

Stand composition and structure of natural mixed-oak stands of common-oak (Quercus robur L.) and pyrenean-oak (Quercus pyrenaica Willd.) were studied. Diverse compositional and structural elements in early and late successional stand stages were analysed. The study was conducted in north and central Portugal where different natural mixed oak forests types are located. The following mixed-oak forest types involving common-oak and pyrenean-oak were studied: common-oak & other hardwoods; common-oak & cork-oak (Quercus suber L.); ash (Fraxinus angustifolia Vahl) & pyrenean-oak; and pyrenean-oak & madrone (Arbutus unedo L.). Measurements were made in early and late successional stand stages on the different mixed oak forest types. Different stand characteristics and indices were used to describe and compare stand structure and composition. The study showed changes in species diversity and stand structure. Most tree species in mature stands are present in early stages but with higher abundance. Shannon diversity index may change between 0.798 and 1.915. Significant differences on species diversity and abundance were found depending on the forest type and successional stage. Mature mixed-oak forests have high species diversity with an abundance of small to medium tree size species. Species distribution and diameter differentiation indices range from 0.30 to 0.70 and 0.52 to 0.82, respectively, revealing significant structural complexity. The average number of standing and downed dead trees was 265 and 83 trees ha⁻¹ for early and late stage, respectively, with 6.9 and 65.4 m³ ha⁻¹. Higher values of stand diversity index were 41 and 53 in more complex and developed forests. Later stand stages have complex structure, with a wider range of tree diameter distribution and higher degree of irregularity.     

Structure and diversity of small-mammal communities of lowland forests in the rural central European landscape

Small-mammal communities were monitored over 4 years in South Moravian rural lowland forests in order to study relationships with various forest habitat types. Early successional sites (plantations) and forest edges maintained communities with higher abundance and diversity. Lowest total abundance was observed in unmanaged lowland forest and highest abundance in plantations with irregular undergrowth management. Low diversity was typical of non-fruiting monocultures. In plantations, both diversity and abundance were affected by herb layer removal, with sites subject to regular cutting, displaying lower levels. Significant differences in diversity and species richness were only detected between early and late successional sites, with the former showing higher values. Two basic small-mammal community groups were determined; (1) those inhabiting sites with a thick herb undergrowth and an open tree canopy (plantation) and (2) those inhabiting stands with sparse or no herb layer and a closed canopy (high forest). Forest stands support long-term populations of dominant small forest-dwelling mammal species and, as such, serve as reservoirs, even in rural landscapes dominated by intensive arable farming. Managed early succession lowland forests represent important refuges for a number of small open-habitat mammals negatively affected by intensive farming.     

Calibration and assessment of seasonal changes in leaf area index of a tropical dry forest in different stages of succession

A simple measure of the amount of foliage present in a forest is leaf area index (LAI; the amount of foliage per unit ground surface area), which can be determined by optical estimation (gap fraction method) with an instrument such as the Li-Cor LAI-2000 Plant Canopy Analyzer. However, optical instruments such as the LAI-2000 cannot directly differentiate between foliage and woody components of the canopy. Studies investigating LAI and its calibration (extracting foliar LAI from optical estimates) in tropical forests are rare. We calibrated optical estimates of LAI from the LAI-2000 with leaf litter data for a tropical dry forest. We also developed a robust method for determining LAI from leaf litter data in a tropical dry forest environment. We found that, depending on the successional stage of the canopy and the season, the LAI-2000 may underestimate LAI by 17% to over 40%. In the dry season, the instrument overestimated LAI by the contribution of the woody area index. Examination of the seasonal variation in LAI for three successional stages in a tropical dry forest indicated differences in timing of leaf fall according to successional stage and functional group (i.e., lianas and trees). We conclude that when calculating LAI from optical estimates, it is necessary to account for the differences between values obtained from optical and semi-direct techniques. In addition, to calculate LAI from litter collected in traps, specific leaf area must be calculated for each species rather than from a mean value for multiple species.     

Plant diversity and soil properties in pristine and managed stands from Bosnian mixed forests

We compared pristine and managed beech–fir mixed forestsin Bosnia to assess the impact of management on plant speciesdiversity, vegetation structure and soil–vegetation interaction.Traditionally, management of Bosnian mixed forests has beenby the opening of small gaps, similar to those occurring naturally.Species composition revealed three successional vegetation groups,two mainly in managed forests (regenerating and mature) andone mainly in pristine old growth forests. Managed sites showeda tendency to be more diverse in plant species than pristinesites, and were also more heterogeneous. Diversity was clearlyexplained by vegetation groups; mature managed stands showingthe highest plant diversity and regenerating stands the lowest,both in terms of species richness and abundance distributions.Plant diversity decreased with canopy cover and increased withcarbon concentration in the A_h horizon, being accurately modelledby both factors. Although topographical conditions were homogeneousamong sites, vegetation was very sensitive to small changesin environmental variables at the local scale. Vegetation reflecteda combination of past situations, represented by C in the A_hhorizon, and current conditions, represented by canopy coverand the C : N ratio in the LF organic layers, suggesting bothsensitivity to present-day environmental conditions as wellas the imprint of past events.     

Resilience of stand structure and tree species diversity in subtropical forest degraded by clear logging

Subtropical forests in the Ryukyu Islands have been degraded by silvicultural practices, and thus their structural attributes are being shifted to other states dominated by a few tree species. This study clarified the mechanisms of the change, and examined the effect of clear logging on the resilience of a subtropical forest. Sprouting regeneration and typhoon disturbance were introduced into an individual-based model, SEIB-DGVM, for describing stand development and succession. The regeneration dynamics from young secondary to old-growth stands were reproduced fairly well with the model. Sprouting recruitment produced high stem density at the beginning of stand development, which caused a self-thinning trajectory following the −3/2 power law. In the late development stage after 70 years, tree species diversity fluctuated because of the regenerative response of sprouting species and the facilitatory effect of typhoon disturbance on the coexistence of subordinate species. The death of canopy trees because of typhoon disturbances reduced the dominance of Castanopsis sieboldii, and depressed its dominance in the understory. Consequently, the understory species could establish by virtue of fallen canopy trees, and tree species diversity increased at the stand level. Clear logging experiments in the model revealed that species diversity deteriorated, especially in the stand dominated by sprouting species. Resilience of subtropical forests was determined by initial species composition before clear logging. Our simulation results suggest that repeated logging drives subtropical forests with high species diversity to a stand monopolized by C. sieboldii.     

Diversity and succession of adventive and indigenous vascular understorey plants in Pinus radiata plantation forests in New Zealand

The vegetation of Pinus radiata plantation forests in New Zealand was studied to examine how the indigenous flora has responded to this novel habitat. A chronosequence of stands about 5, 16 and 27 years was assessed in each of four different biogeographic regions to test the effects of several stand and site factors on the succession of vascular understorey plant communities. A total of 202 indigenous and 70 adventive vascular plant species were found across all study areas, with considerable geographic variation among forests in species composition, species richness (range 48–135 species), and the percentage of indigenous species (50–86%). Both richness and cover of adventive species decreased significantly over time, whereas richness and cover of indigenous species was highest in the oldest stands, and overall species richness was lowest at mid-rotation. The guild composition changed from dominance of grasses and forbs in young stands to dominance of ferns and understorey trees in mature stands. These temporal changes were accompanied by a decrease in light-demanding pioneer species and an increase in shade tolerant, later seral species adapted to a forest environment. Measurements of the degree of canopy closure in stands with low or high stocking and modelling of temporal changes of canopy closure indicated that these understorey plant dynamics are influenced by changes in light availability as stands age. Despite the successional changes within forests, geographic variation more strongly influenced understorey communities because stands within a forest area were grouped together in DCA and TWINSPAN analyses, along rainfall and temperature gradients. Although the canopy species of such intensively managed plantation forests is an alien element in the New Zealand flora, the sheltered forest environment of older stands allows the establishment of a mostly indigenous forest understorey community with considerable similarities to indigenous forests located nearby.     

Gap dynamics in the Western Carpathian mixed beech old-growth forests affected by spruce bark beetle outbreak

Understanding how species-specific disturbances affect the dynamics of mixed forests is becoming increasingly important due to rapidly changing disturbance regimes. This study estimated the effect of Norway spruce (Picea abies (L.) Karst.) mortality on the disturbance processes in two mixed beech stands of the Western Carpathians that were affected by a bark beetle outbreak. We evaluated the size distribution, fraction of canopy and expanded gaps, the characteristics of gapmakers and the contribution of different species to gap size. The measured canopy gap fraction was <5%, and most canopy gaps were small (<100 m²). Spruce was the most abundant gapmaker, and its share among gapmakers was 3–6 times higher than its share in the canopy. We found that the increase in spruce mortality due to the outbreak resulted in a fine-scale mortality pattern. However, spruce gapmakers did not contribute much to gap area size, as shown by a weak correlation between the number of spruce gapmakers and the area of expanded gaps. Diameter distribution of living versus recently dead trees showed that beech mortality occurred disproportionately in large size classes. However, dead spruce trees were equally frequent in all diameter classes, which means beetles did not exclusively attack larger trees in these stands during the outbreak. We conclude that spruce mortality may have influenced successional processes by giving a competitive advantage to two other species that were not affected by the outbreak, provided that a high deer browsing intensity does not hinder the regeneration of new seedlings.

     

Effect of thinning on the development of compression wood in stems of Corsican pine

This study considered the effects of thinning on the development of compression wood in stems of 35-year-old stand of Corsican pine (Pinus nigra L.). Part of the stand had been thinned at 5-yearly intervals and part left unthinned. Twenty trees each from the thinned and unthinned stands were randomly selected and felled. Measurements were made on tree height, stem diameter, stem slenderness and canopy depth. Wood samples were removed from the central part of the main log and cross-sectional measurements made on ring width, basic density and compression wood content. Cross-sectional area of compression wood was found to be three time higher in stems from the unthinned trees in comparison with those from the thinned trees. No significant differences in mean radial ring width or basic density were found between treatments. Correlations indicated that, with increasing in stem diameter, compression wood content increased in the unthinned trees, while a decline in compression was observed in the thinned trees. Tree height was also positively correlated with compression wood content in unthinned trees, while no equivalent relationship was observed in thinned trees. Observations from this study, while not conclusive, suggest that phototropic stimulus may be producing stem inclinations in the unthinned stand as trees compete for space in the canopy, whereas crown competition has been largely eliminated in the thinned stand; and that this is responsible for compression wood levels recorded in this study.     

Impacts of changing climate on the productivity of Norway spruce dominant stands with a mixture of Scots pine and birch in relation to water availability in southern and northern Finland

A process-based ecosystem model was used to assess the impacts of changing climate on net photosynthesis and total stem wood growth in relation to water availability in two unmanaged Norway spruce (Picea abies) dominant stands with a mixture of Scots pine (Pinus sylvestris) and birch (Betula sp.). The mixed stands were grown over a 100-year rotation (2000-99) in southern and northern Finland with initial species shares of 50, 25 and 25% for Norway spruce, Scots pine and birch, respectively. In addition, pure Norway spruce, Scots pine and birch stands were used as a comparison to identify whether species' response is different in mixed and pure stands. Soil type and moisture conditions (moderate drought) were expected to be the same at the beginning of the simulations irrespective of site location. Regardless of tree species, both annual net canopy photosynthesis (P(nc)) and total stem wood growth (V(s)) were, on average, lower on the southern site under the changing climate compared with the current climate (difference increasing toward the end of the rotation); the opposite was the case for the northern site. Regarding the stand water budget, evapotranspiration (E(T)) was higher under the changing climate regardless of site location. Transpiration and evaporation from the canopy affected water depletion the most. Norway spruce and birch accounted for most of the water depletion in mixed stands on both sites regardless of climatic condition. The annual soil water deficit (W(d)) was higher on the southern site under the changing climate. On the northern site, the situation was the opposite. According to our results, the growth of pure Norway spruce stands in southern Finland could be even lower than the growth of Norway spruce in mixed stands under the changing climate. The opposite was found for pure Scots pine and birch stands due to lower water depletion. This indicates that in the future the management should be properly adapted to climate change in order to sustain the productivity of mixed stands dominated by Norway spruce.     

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.

     

Temperate-like stand dynamics in relict Mediterranean-fir (Abies pinsapo, Boiss.) forests from southern Spain

Introduction and statement of the research questions

Gap dynamics have been widely studied in forests of Abies spp. from temperate and boreal regions. The local microclimate and competition for light have been identified as the main factors controlling changes in species composition and canopy structure, however little is known on dynamics of such forests in Mediterranean.

Experimental design and aims

We studied forest structure and dynamics of Abies pinsapo stands in southern Spain, in contrasting habitats and successional status. In addition past regeneration patterns and their relationship to canopy structure, disturbances and forest-use history were investigated.

Results

Stands structure attributes were within the range described for temperate conifer biomes. The age structure revealed two main cohorts comprised of a few > 100 year-old trees and abundant younger trees established in a single recruitment event after the stands were protected in the 1950s. Initial growth-rate analyses indicated that A. pinsapo regenerated mainly in small canopy gaps, while only 15% recruited from the forest understorey. For the last ten years, basal area increment was lower than 10 cm² y^?1 in 91% of studied trees and growth rate differences between trees narrowed.

Conclusion

Stand dynamics in A. pinsapo forests maintain general features of temperate fir forests. Tree establishment over time and current stand structure fit to known changes in forest use. Widespread growth decline trends might be linked to stand stagnation and global warming.     

The changing Himalayan landscape: pine-oak forest dynamics and the supply of ecosystem services

This review summarizes the current state of knowledge on pine and oak forest dynamics in the mid-montane central Himalayan forest and the ecosystem services associated with these vegetation types. Forest ecosystems play a crucial role in the livelihood of the central Himalayas as well as the adjacent plains, providing a number of tangible and intangible ecosystem services, at each stage of succession. The successional sequence starts from warm temperate grasslands, followed by early successional pine forests, mid-successional pine-oak mixed forests and eventually culminating in a late successional oak community. This successional sequence is considerably influenced by disturbances like fire, grazing, and lopping, which maintain the vegetation types in their current form and can act as potential drivers of change. Fire and grazing in grasslands and pine forests inhibit the successional process by hindering the establishment of pioneer and late successional species, respectively. Potential land-cover changes with forest succession can lead to changes in ecosystem services supply. We found that the number of ecosystem services associated with these vegetation types increase from early to late successional community. Current management approaches fail to include the dynamic nature of vegetation, which is essential for maintenance of ecosystem service supply. In conclusion, the trade-offs between ES of global (biodiversity and carbon) and local importance (fuel wood and fodder) have to be examined carefully in order to have effective conservation and management plans for the region.     

Breeding bird community composition in different successional vegetation in the montane coniferous forests zone of Taiwan

To examine the relationship between forest succession following fire and the composition of bird communities, we investigated the vegetation structure, bird population density, foraging behavior and guild structure in bamboo grasslands (11 years since the last fire), pine savanna (41 years), pine woodland (58 years), old-growth hemlock forest (never burned), and old-growth spruce forest (never burned) in the Tatachia area of central Taiwan. Canopy height, total foliage cover, tree density, total basal area of tree, total basal area of snags, foliage height diversity, and tree species richness all increased with successional age. However, shrub cover peaked in intermediate successional stages. The vertical profile of foliage cover was more diverse in later successional forests, which had more breeding bird species and ecological guilds. All the breeding bird species recorded in early and intermediate stages were also found distributed in the late successional forests. Because Taiwan has high precipitation and humidity, and most forest fires in Taiwan are caused by human activities, forest fires and large areas of early successional vegetation were probably rare in the mountain areas of Taiwan prior to the arrival of humans. Therefore, bird species have not had enough time to adapt to areas with early or intermediate successional vegetation. Moreover, late successional forests host all the major plant species found in the early and intermediate stages and have higher foliage height diversity index, which was positively correlated with the bird species richness and bird species diversity index in this study. As a result, all breeding bird species and guilds in the area can be found in late successional forests. Efforts for conserving avian diversity in Taiwan should focus on protecting the remaining native old-growth forests.     

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.     

Effects of artificial afforestation and successional stage on a lowland forest bird community in southern China

To examine the relationship between forest succession after severe logging forestry practices and the composition of avian communities, we investigated how forest bird composition and guild structure change as a function of structural properties along a successional gradient, including a climax mature forest (>400 years), a rehabilitated mixed forest (50-70 years), and a disturbed Masson pine forest (70 years) of the Dinghushan Nature Reserve, Guangdong Province, China. Of a total of 51 resident species recorded, mixed forests hosted the highest numbers of individuals and species, reflecting the high species richness of both forest and non-forest species. For forest-dependent species, however, mature stands had the highest observed and estimated species richness. Of 6 habitat-use guilds identified, vertical-profile generalists and understory-birds formed the two dominant guilds, accounting for 54.0% and 38.7% of all individuals respectively. The results of canonical correspondence analysis (CCA) clearly showed that most forest-dependent species were associated with high proportions of native canopy cover and the mean density of dead trees and large trees, which are characteristic of old-growth mature forests (horizontal heterogeneity) at stand level. Accordingly, conservation efforts should focus on the specialized requirements of the most habitat-restricted species in the future, especially for understory insectivores (Babblers) and large-tree users in mature subtropical monsoon forests of southern China. Moreover, since regenerating mixed forests are very similar to mature forests in both vegetation structure and bird community composition, we recommend that logging cycles (>50 years) be increased to a minimum of 50 years in southern China, so that a balance between economic and ecological interest can be reestablished.     

Promoting old-growth characteristics and long-term wood production in Douglas-fir forests

Trade-offs among wood production, wood quality and ecological characteristics in the management of harvested forest stands are explored through model simulation of various silvicultural regimes. Long-term production of merchantable wood, production of various types of high-quality wood, and the level of certain quantitative ecological indicators are projected for coniferous forests of Pacific Northwestern USA. The set of ecological indicators used is based on the species composition and physical structure of old, unlogged forest stands. Simulations are performed with an ecological model of forest stand dynamics that tracks the fate of live and dead trees. Short rotations (<50 years) produce the least amount of high-quality wood over the multi-century simulation period. They also fail to generate ecological attributes resembling those of old forest stands. Production of high-quality wood is moderate to high under all rotations of 80 years or more; however, most ecological indicators require longer rotations unless alternatives to clearcutting are applied. Alternatives examined include retention of 15% cover of live tree canopy at each harvest in combination with artificial thinning between harvests. Thinning from below can expedite the development of large live and dead trees, and canopy height diversity without greatly diminishing wood quantity or quality. Proportional thinning retains understory stems, thereby expediting the recruitment of shade-tolerant trees. A possible drawback to thinning, particularly proportional thinning, is the diminished production of clean-bole wood at rotations of 150 and 260 years. It is concluded that most wood quantity, wood quality and ecological objectives can be met with long rotations (ca. 260 years). Certain objectives can be met with shorter rotations (80–150 years) when treatments of thinning and canopy tree retention are applied.     

Biomass allocation strategies and productivity of tropical trees related to successional status

The productivity and biomass allocation strategies of two early successional (ES) and two late successional (LS) tropical tree species were compared and related to their successional status. Apart from distinct differences in clean bole length, crown depth, maximum crown width and leaf area index (LAI), the ES species showed higher allocation to the shoot, particularly to the bole, whereas the LS species had higher allocation to the root. The ES species with shallow root system had more root biomass within the upper 20 cm of the soil profile while the LS species with deeper roots had a higher proportion of root biomass distributed below the 20 cm depth. The productivity of the shoot of ES species was significantly higher than that of LS species. However, root productivity for ES species was higher only up to 4 years of age; the differences were not significant between 5 and 7 years. The implication of these results for agroforestry and mixed plantation forestry is emphasized.     

A stand density index for complex mixed species forests in the northeastern United States

Quantifying stand density is important for accurate prediction of net biomass and carbon accumulation, for estimating growth and mortality risks of trees, stands, and regions, and for the management of forests for multiple goods and services. Building on previous work relating maximum stand density to wood specific gravity, we develop a stand density equation for the mixed species forests of the northeastern United States using data from the US Forest Service, Forest Inventory and Analysis program. We used quantile regression, in concert with a quantile selection and evaluation procedure, to ensure conformity between our density measure and previously developed guidance for well-studied stand types. The resulting strictly additive relative density measure appears to provide reasonable prediction of maximum density even for plantations of exotic conifers in the region. The results suggest that maximum stand densities after accounting for wood specific gravity may be lower in northeastern North America than in the south or west.     

The successional status of tropical rainforest tree species is associated with differences in leaf carbon isotope discrimination and functional traits

We characterised the among species variability in leaf gas exchange and morphological traits under controlled conditions of seedlings of 22 tropical rainforest canopy species to understand the origin of the variability in leaf carbon isotope discrimination (Δ) among species with different growth and dynamic characteristics (successional gradient). Our results first suggest that these species pursue a consistent strategy in terms of Δ throughout their ontogeny (juveniles grown here versus canopy adult trees from the natural forest). Second, leaf Δ was negatively correlated with WUE and N, and positively correlated with g_s, but among species differences in Δ were mainly explained by differences in WUE. Finally, species belonging to different successional groups display distinct leaf functional and morphological traits. We confirmed that fast growing early successional species maximise carbon assimilation with high stomatal conductance. In contrast, fast and slow growing late successional species are both characterised by low carbon assimilation values, but by distinct stomatal conductance and leaf morphological features. Along the successional gradient, these differences result in much lower Δ for the intermediate species (i.e. fast growing late successional) as compared to the two other groups.