Spatial distribution of regeneration in West-Carpathian uneven-aged silver fir forests

The objective of our study was to examine whether distribution of regeneration in uneven-aged fir (Abies alba Mill.) forests is related to the spatial pattern of trees. In 12 sample plots of size 0.45–1.00 ha (in total 8.65 ha, with stand basal areas ranging from 27.6 m² ha^–1 to 39.5 m² ha^–1), all live and dead trees above 5 cm in d_1.3 were mapped and their diameters measured. In eight plots, all live and dead fir saplings were mapped. In three plots, the number of live fir saplings and seedlings was registered on small systematically distributed circular plots. The values of an analytically developed index of stand influence were compared in patches occupied and unoccupied by live or dead fir regeneration. Contrary to preliminary assumptions, only in a few cases did saplings and trees 5–15 cm in d_1.3 appear more often in gaps and looser stand patches. Rather, in many plots, the opposite tendency was observed. The seedling density showed a weak but positive correlation with the index of influence. If the spatial pattern of regeneration reflects the spatially varying mortality of juvenile trees, then no evidence was found that stand competition was the most important factor inducing this mortality. On the contrary, on the basis of the results obtained, we can presume that the survival rate of juvenile firs was higher in patches with a relatively higher local basal area. Thus, it was hypothesised that, first, dispersion of regeneration in uneven-aged fir forests is controlled by easy-to-change edaphic factors such as humus form and acidity of the upper soil horizons, and second, that these soil features are linked with the spatial pattern of trees.     

Spatial and volume patterns of an unmanaged submontane mixed forest in Central Europe: 160 years of spontaneous dynamics

The long-term development of stand characteristics and tree spatial patterns (TSP), their mutual relation, and linkage to site and tree species were studied in the Boubín primeval forest (protected since 1858). Surveys were carried out in 1851, 1961/1964, 1972, 1984/1989, 1996 and 2010 on one to six research plots sized 0.58-1.00 ha. To see how results from these surveys could be generalized, results were also compared with whole-area data sets (46 ha) from 1972 to 2010.The proportion of Abies alba continually decreased, from 20% in 1851 to 2-3% in 2010. This decrease started no later than at the beginning of the 20th century. In contrast, the proportion of Fagus sylvatica slightly increased. The sum of dead and living wood volume was stable during the 159 year period, with deviations of only up to 5%. From 1961/64 to 2010, the number of living trees continually decreased, but the mean-tree volume and volume of dead wood increased. The distribution of dead wood always differed from the distribution of living trees.A random TSP was always most common, which seems to be typical for this type of forest. This was true even when A. alba was gradually forced out from the stand structure. Nevertheless, the pronounced decrease of A. alba found during the 1961/64 survey compared with 1851 was reflected in a tendency towards clustering, as gaps from A. alba dieback gradually closed. In these gaps, competitive pressure was lower, resulting in more clustered distributions. Regular distribution was recorded only rarely. F. sylvatica maintained a random or clustered TSP over long periods, depending on site conditions. On water-affected plots, it had a consistently higher tendency towards clustering. The only step change in TSP occurred due to Hurricane Emma in 2008. Otherwise, the most marked TSP changes over time were found for Picea abies, in which there was a gradual trend from clustered to random, connected with a decreasing number of individuals.The main reason for the decline of A. alba and the decreased number of living trees from 1961/64-2010 was the on-going effect of a high stock of red deer at the turn of the 19th and 20th centuries. The decreasing trend in the number of living trees showed no marked change, even 60 years after the game pressure was strongly reduced. However, the vertical structure became increasingly homogenized due to the long-term absence of severe abiotic and/or biotic disturbances.     

Short term effects of alternative silvicultural treatments on stand attributes in irregular balsam fir-black spruce stands

The eastern Canadian boreal forest exhibits a specific disturbance regime where forest fires are less frequent than in the western part. This particularity may explain the abundance of irregular stands with distinct ecological features. To ensure sustainable forest management, these characteristics require the implementation of an adapted silviculture regime. In this context, two selection cutting methods were developed and compared with more conventional techniques, initially designed for cutting more regular stands of the boreal forest (cutting leaving small merchantable stems, careful logging preserving advance regeneration). The comparison focused on the capacity of treatments to maintain the primary attributes of irregular boreal forests, including complex vertical structure, abundant tree cover, species composition, and an abundance of dead wood. Mortality and regeneration processes were also compared.     

Microsite patterns of conifer seedling establishment and growth in a mixed stand in the southern Alps

In many forests of the Alps, permanent forest cover and, therefore, its continuous renewal is the main silvicultural goal. Regenerating these forests must be based on a sound understanding of the ecology of the tree species in question. The regeneration of silver fir (Abies alba Mill.) and Norway spruce (Picea abies (L.) Karst.) in the upper montane zone of the southern central Alps has so far received little attention. The aim of this study was, therefore, to characterize the ecological niche for the establishment and growth of silver fir and Norway spruce seedlings in this zone. The study was conducted as a case study on a plot of 1.5 ha in the upper montane zone of southern Switzerland, at an altitude of 1380 m. The basal area of the mixed Norway spruce–silver fir stand was 43.7 m² ha⁻¹. Seedlings were censused on 375 plots of 1.0 m² area. Seedling density was 1.30 ± 4.25 m⁻² (mean ± standard deviation) for Norway spruce and 0.87 ± 1.50 m⁻² for silver fir. Logistic regression models were used to test the effect of microsite characteristics on seedling occurrence, and general linear models for effects on seedling height growth and biomass increment. Most seedlings received less than 10% light (photosynthetic photon flux density) as compared to values in the open. Silver fir occurrence was positively related to microsites at the edge of canopy trees, but unrelated to ground cover type, light and micro-relief. Norway spruce occurrence was only, and positively, related to the presence of mosses. Height growth and biomass increment of seedlings of both species were only loosely correlated with microsite conditions. Seedlings without canopy cover grew faster than those under canopy cover, probably as a result of light and moisture limitations under the canopy of adult trees. Diffuse radiation was positively correlated with average annual biomass increment of silver fir, but not of Norway spruce seedlings. In general, the results suggest that silver fir seedlings have less specific microsite requirements than Norway spruce seedlings in terms of ground cover. They are also more shade tolerant, and therefore, grow faster than Norway spruce in low-light environments of the upper montane zone of the southern Alps. While the microsite concept can be helpful in designing silvicultural operations, it has limitations when only patch characteristics that are easy to assess are used, and others neglected. Finally, our study suggests that counting the number of green shoots is a promising method to quickly and non-destructively estimate the biomass of a great number of small seedlings.     

Gap dynamics in a near-natural spruce forest at Mt. Brocken,Germany

We analyzed indices of gap dynamics, i.e. gap formation and gap closure rates as well as turnover rates, in a montane conifer forest in the temperate zone and asked: (1) Are the characteristics of gap dynamics comparable with those of other temperate forests? (2) Do gap dynamics exhibit temporal trends, and (3) if so, are they related to recent climate change? We also addressed methodological issues concerning the differences observed at differing temporal and spatial scales of the used datasets. Thus, we asked further how the indices of gap dynamics are influenced (4) by differences in the length of observation periods and (5) by differences in gap size categories included in the study?     

Optimizing the joint production of timber and bilberries

Berries and mushrooms are increasingly appreciated products of Finnish forests. Therefore, there is a need to integrate them in silvicultural planning. Bilberry (Vaccinium myrtillus L.) is an economically important wild berry that is widely collected for household consumption and sale in North Karelia, Finland. In this study, bilberry yield models developed recently were included in a stand growth simulator and the joint production of timber and bilberry was optimized by maximizing soil expectation value (SEV) with 3% discounting rate, assuming that 75% of the bilberry yield is harvested. The effect of bilberry production on the optimal stand management increased with increasing bilberry price. With high bilberry prices (4–8 € kg⁻¹) it was optimal to manage the mixed stand of Scots pine, Norway spruce and birch, and the pure stand of Norway spruce so as to promote bilberry production. In the Scots pine stand, where bilberry yields are higher, bilberry production affected optimal stand management already with a price of 2 € kg⁻¹. Compared to timber production, joint production led to longer rotation lengths, higher thinning intensities, more frequent thinnings, and higher share of Scots pine in the mixed stand. The contribution of bilberries to the total SEV increased with increasing bilberry price and discounting rate. In the mixed stand and pine stand the SEV of bilberry production, calculated with 3% discounting rate, exceeded the SEV of timber production when bilberry price was 4 € kg⁻¹.With 4% discounting rate this happened already with bilberry price of 2 € kg⁻¹. It was concluded that forest management which promotes bilberry yields is the most profitable in pine stands where the potential bilberry yields are high.     

Regeneration in experimental gaps of subalpine <Emphasis Type="Italic">Picea abies</Emphasis> forest in the Slovenian Alps

In 1998, we analysed regeneration success in four 20 year old medium-sized (600–800 m²) and four small experimental gaps (100–200 m²) in subalpine Norway spruce forest in Triglav National Park, Slovenia. We assessed the influence of site (depressions, slopes), gap size, and position within gap (centre, gap south, and north edge) on regeneration success and interactions between regeneration density, height, height increment, direct and diffuse light, ground vegetation, soil depth, thickness of humus horizons, presence of woody debris, and microrelief. Seedling density was higher in depressions, where more seedlings developed in central positions of gaps in both gap sizes, and in microsites with deeper soils, less competition from ground vegetation and convex microsites. On slopes, a higher seedling density was found in small gaps, and at the northern edge of gaps regardless of gap size. Here, seedling density was negatively influenced by diffuse radiation and positively by soil depth. The results suggest that regeneration should be initiated from medium to large gaps in depressions and small gaps on slopes. Development of established seedlings was enhanced by higher radiation levels on both sites, therefore favourable extension of the gaps would be lateral to northwards.     

Factors affecting the probability of windthrow at stand level as a result of Gudrun winter storm in southern Sweden

A storm in January 2005 in southern Sweden (Gudrun) caused large damage to forests, archaeological sites, buildings, power supply, telecommunications, and infrastructure. In this work, we analyzed the factors affecting the probability of windthrow at stand level (i.e. tree- and stand characteristics) in southern Sweden. We selected a sub-sample (1721 plots) of existing plots from the Swedish National Forest Inventory (inventoried in 2003 and 2004). To assess whether these plots had suffered damage or not, aerial photographs were taken in summer 2005. In order to analyze the effects of measured characteristics on damage, field data for each sub-sample plot were merged with the corresponding result of the aerial photo interpretation.We found that the plots damaged were typically well stocked mature stands, dominated by Norway spruce (Picea abies (L.) Karst.), and often also recently thinned. Probability of damage increased with increase in stand height. In mixed stands, decrease of proportion of Norway spruce decreased the probability of damage. An admixture of deciduous trees, leafless in this winter storm, showed larger effect in this sense than that of Scots pine.     

Climate and historical stand dynamics in the tropical pine forests of northern Thailand

Forest recruitment is the outcome of local- and regional-scale factors such as disturbances and climate. The relative importance of local- and regional-scale factors will determine the spatial scale at which temporal pulses of recruitment occur. In seasonal tropical forests, where the annual dry-season is a critical bottleneck to seedling survival, multi-year periods of relatively cool, wet dry seasons may be required for successful tree recruitment. Consequently, when such conditions are present, region-wide synchronisation of recruitment may occur. To examine the case for regional synchronisation of forest dynamics in the seasonal tropical pine forests of northern Thailand, we investigated forest age structures at three spatial scales: stand, site and region. We compared forest age structures with instrumental climatic records beginning in 1902. We found significant statistical evidence of synchronous recruitment at the stand- and site-scales, but not at the regional-scale. While correlations between recruitment and climate were not statistically significant, recruitment success was often linked to favourable climatic conditions. For example, recruitment at all sites was associated with multi-year periods of cool-wet dry seasons. The lack of significant correlations between recruitment and climate appears to reflect complex interactions among local disturbance history, regional climate variability and pine recruitment.     

The effects of burning and dead-wood creation on the diversity of pioneer wood-inhabiting fungi in managed boreal spruce forests

The loss of natural forest habitats due to forestry is the main reason for the decline in boreal forest biodiversity of the Nordic countries of Europe. Ecological rehabilitation may provide means to recover and sustain biodiversity. We analyzed the effects of controlled burning and dead-wood creation (DWC) on the diversity of pioneer wood-inhabiting fungi in managed Norway spruce (Picea abies) forests in southern Finland. Altogether 18 stands were first subjected to a partial cutting with ordinary logging residues in form of cut stumps and treetops left on site. The subsequent rehabilitation treatments consisted of a controlled burning applied in half of the stands and three levels of dead-wood creation (5, 30 and 60 m³ ha⁻¹). The DWC involved creation of logs; felling of whole trees to mimic downed logs formed by natural disturbance processes. Each treatment was replicated three times. Inside each stand, substrates were sampled in two different biotopes; one on mineral soil and one on mineral soil with a thin peat layer. We surveyed the fungal flora on the logs (n = 364) and the ordinary residue stumps (n = 1767) and tops (n = 845) five years after the treatments.When comparing different stands, controlled burning had a significant effect on species composition; certain species were significantly more frequent on substrates in burned stands than in unburned stands, indicating that these species were favored by controlled burning. By contrast, we found no significant effects of DWC levels or biotope on species composition or richness. When comparing different substrates, 99% of the logs hosted at least one species and the occurrence probability of certain species was significantly higher on logs than on ordinary residue stumps and tops. Yet, volume-based rarefaction analyses showed that residues were more species dense than the logs, indicating that ordinary logging residues constitute important resources for many pioneer species.We conclude that controlled burning combined with DWC have strong effects on biodiversity; it modifies the composition of the pioneer wood-inhabiting fungal species found in managed forests and may thereby also influence the further succession and diversity of the secondary fungal flora.     

Forest stand dynamics and sudden oak death: Mortality in mixed-evergreen forests dominated by coast live oak

Sudden oak death (SOD), caused by the recently discovered non-native invasive pathogen, Phytophthora ramorum, has already killed tens of thousands of native coast live oak and tanoak trees in California. Little is known of potential short and long term impacts of this novel plant–pathogen interaction on forest structure and composition. Coast live oak (Quercus agrifolia) and bay laurel (Umbellularia californica) form mixed-evergreen forests along the northern California coast. This study measured tree mortality over a gradient of disease in three time periods. Direct measurements of current mortality were taken during 2004, representing a point-in-time estimate of present and ongoing mortality. Past stand conditions, c. 1994, were estimated using a stand reconstruction technique. Future stand conditions, c. 2014, were calculated by assuming that, given a lack of host resistance, live trees showing signs of the disease in 2004 would die. Results indicate that coast live oaks died at a rate of 4.4–5.5% year⁻¹ between 1994 and 2004 in highly impacted sites, compared with a background rate of 0.49% year⁻¹, a ten-fold increase in mortality. From 2004 to 2014, mortality rates in the same sites were 0.8–2.6% year⁻¹. Over the entire period, in highly impacted sites, a 59–70% loss of coast live oak basal area was predicted, and coast live oak decreased from 60% to 40% of total stand basal area, while bay laurel increased from 22% to 37%. Future stand structures will likely have greater proportions of bay laurel relative to coast live oak.     

Evidence of mixed-severity fires in the foothills of the Rocky Mountains of west-central Alberta, Canada

This study presents new evidence of historic low-to-moderate-severity fires, intermixed with high-severity fires, in the foothills of the Rocky Mountains of west-central Alberta, Canada. High-severity fires that burned 120-300 years ago initiated even-aged cohorts of fast-growing lodgepole pine at each of the six study sites. Evidence of subsequent, low-to-moderate-severity fires included single and double fire scars on thin-barked lodgepole pine that were as small as 3.6 cm in diameter at the time of the fire, but survived. These low-to-moderate-severity fires resulted in structurally complex stands with a broad range of tree diameters and multiple cohorts of lodgepole pine, white and black spruce, and subalpine fir. At the site level, fire return intervals were variable, ranging from 29 to 167 years, but most were <80 years. Of the 9 years in which we documented low-to-moderate-severity fires, only the fires in 1889 and 1915 scarred trees at more than one site, indicating that these fires were small and had local effects. The new knowledge of historical, low-to-moderate-severity fires provided by this study has important implications for ecologically sustainable forest management. Although we recognize that further research needs to determine the extent of low-to-moderate-severity fires at the landscape scale, our results clearly indicate that a mixed-severity fires occurred at least locally. A broader range of silvicultural systems than is currently practiced would be consistent with historic forest dynamics.     

Tree rings indicate different drought resistance of a native (Abies alba Mill.) and a nonnative (Picea abies (L.) Karst.) species co-occurring at a dry site in Southern Italy

Climate changes induced by the anthropogenic alteration of the atmospheric radiative balance are expected to change the productivity and composition of forest ecosystems. In Europe, the Mediterranean is considered one of the most vulnerable regions according to climatic forecasts and simulations. However, although modifications in the inter-specific competition are envisaged, we still lack a clear understanding of the ability of the Mediterranean vegetation to adapt to climate changes. We investigated how two co-occurring tree species commonly used in afforestation programmes, the native Abies alba Mill. and the nonnative Picea abies L. Karst., adapt to climate change by assessing their growth performance and physiological responses in relation to past climate variability. Growth was addressed by analysing tree-ring width and carbon and oxygen stable isotopes. Statistical relationships between isotopic value and monthly climate data suggest that the two species underwent ecophysiological adaptation to Mediterranean climatic constraints. These adaptations are also expressed in the ring-width data. Based on the carbon isotope ratio reflecting the stomatal response to drought, we found that the precipitation in the first period of the growing season, i.e. early spring, is a major factor influencing the annual growth of A. alba, which although native, proved to be sensitive to drought. P. abies, on the other hand, showed a higher tolerance to summer drought stress. These findings should help define criteria for sustainability and effective forest conservation in the Mediterranean region.     

The potential of transforming simple structured pine plantations into mixed Mediterranean forests through natural regeneration along a rainfall gradient

The aim was to study the potential for using natural regeneration as a basis for transformation of simply structured conifer plantations into mixed Mediterranean forests. We studied the variation along a rainfall gradient, in the natural regeneration of tree species in the understory of planted 40- to 50-year-old Aleppo pine (Pinus halepensis) forests. The study was conducted within the Mediterranean zone of Israel, which extends from the semiarid northern Negev desert (rainfall ca. 300 mm yr⁻¹) in the south to the humid Upper Galilee in the north (ca 900 mm yr⁻¹). Cover and height, density, and species composition of regenerating trees were measured on south- and north-facing slopes in forest sites of comparable silvicultural history (site preparation methodology, planting density and thinning regime) distributed along the rainfall gradient. Altogether, 12 species of regenerating native broadleaved trees were found in the understory of the various forest sites. Surface cover, density and species richness increased linearly along the entire rainfall gradient, on both north- and south-facing slopes, ranging from zero in the driest forest sites up to 85% cover, 7980 trees ha⁻¹ and 4.5 species per 200 m², respectively, in the most humid ones. Species composition of regenerating trees was also related to rainfall amount, through changes in the relative importance of species along the rainfall gradient. The effect of topographic aspect on tree regeneration was inconsistent, i.e., the interaction Rainfall × Aspect was significant. Nevertheless, the general trend showed better regeneration on north-facing slopes. Most of the regenerating trees in the understory were small, i.e., less than 100 cm in height, with no clear effect of rainfall amount and topographic aspect on the relative abundance of height classes. Regeneration by Aleppo pine was highly variable among and within the different forest sites and ranged from 0 to 1565 trees ha⁻¹, with no clear relationships with rainfall amount and topographic aspect. In light of our results we propose that the future structure of forests should vary with respect to annual rainfall amount within possible silvicultural scenarios.     

Regeneration growth in different light environments of mixed species, multiaged, mountainous forests of Romania

Growth of regenerating trees in different light environments was studied for the mountainous, mixed-species forests in the Carpathian Mountains of Romania. The primary species in these mixtures were silver fir (Abies alba Mill.), European beech (Fagus sylvatica L.) and Norway spruce (Picea abies (L.) Karst). Seedlings/saplings of these species were selected and measured in different stands from two different geographical locations. Regenerating trees were measured for height and diameter growth during the summer of 2002. For each seedling/sapling, percentage of above canopy light (PACL) and stand basal area (BA) were used to assess available and occupied growing space respectively. Regeneration growth was compared against these two variables and regression relationships were developed. Using these models, we predicted the dynamics of regeneration as both growth and species composition. Our results showed that in low-light environments (PACL<20–35%; BA>30 m²/ha), shade tolerant fir and beech clearly outcompeted the spruce. Therefore, in dense stands, spruce could be eliminated by the shade tolerant species. For intermediate levels of cover (PACL=35–70%; BA=15–35 m²/ha) the spruce grew at comparable rates as the beech and fir. All three species showed similar growth rates in open conditions (PACL>80–90%; BA<15–20 m²/ha) with the spruce having a tendency to outgrow the others. However, in terms of establishment, such conditions favor spruce and inhibit fir and beech.     

Logging residue removal after thinning in Nordic boreal forests: Long-term impact on tree growth

The aim of this study was to determine the effect of whole-tree harvesting (WTH) on the growth of Scots pine (Pinus sylvestris L.) and Norway spruce (Picea abies (L.) Karst.) as compared to conventional stem harvesting (CH) over 10 and 20 years. Compensatory (WTH + CoF) and normal nitrogen-based (CH + F or WTH + F) fertilisation were also studied. A series of 22 field experiments were established during 1977-1987, representing a range of site types and climatic conditions in Finland, Norway and Sweden. The treatments were performed at the time of establishment and were repeated after 10-13 years at 11 experimental sites. Seven experiments were followed for 25 years.Volume increment was on average significantly lower after WTH than after CH in both 10-year periods in the spruce stands. In the pine stands thinned only once, the WTH induced growth reduction was significant during the second 10-year period, indicating a long-term response.Volume increment of pine stands was 4 and 8% and that of spruce stands 5 and 13% lower on the WTH plots than on CH during the first and the second 10-year period, respectively. For the second 10-year period the relative volume increment of the whole-tree harvested plots tended to be negatively correlated with the amount of logging residue. Accordingly, the relative volume increment decreased more, the more logging residue was harvested, stressing the importance of developing methods for leaving the nutrient-rich needles on site.If nutrient (N, P, K) losses with the removed logging residues were compensated with fertiliser (WTH + CoF), the volume increment was equal to that in the CH plots. Nitrogen (150-180 kg ha⁻¹) or N + P fertilisation increased tree growth in all experiments except in one very productive spruce stand. Pine stands fertilised only once had a normal positive growth response during the first 10-year period, on average 13 m³ ha⁻¹, followed by a negative response of 5 m³ ha⁻¹ during the second 10-year period. The fertilisation effect of WTH + F and WTH + CoF on basal area increment was both smaller and shorter than with CH + F.     

Long-term browsing impact around diversionary feeding stations for moose in Southern Norway

Diversionary winter feeding of browsing ungulates is an increasingly common management practice although evidence for its efficacy to reduce excessive browsing remains ambiguous. Moreover, comparative estimates on changes in browsing pressure (proportion of available shoots browsed) following long-term winter feeding are currently lacking. We quantified spatiotemporal changes in browsing pressure of moose (Alces alces L.) on commercial and non-commercial tree species around 30 feeding stations after 15–20 years of winter feeding. The results were compared with browsing pressure indices recorded at the same feeding stations 10 years previously. We expected leader stem and lateral twig browsing to have increased over time at a fine spatial scale (≤200 m from feeding station) and to have increased in spatial extent away from feeding stations. Furthermore, we tested whether moose browsing patterns conformed to central-place foraging theory. Despite 2–3-fold higher faecal pellet group numbers in the vicinity of feeding stations, leader stem browsing increased only on the commercially valuable Norway spruce (Picea abies (L.) Karst), a species normally avoided by moose. Lateral twig browsing largely decreased within 200 m of feeding stations while at a broader scale (≤1 km from feeding stations), leader stem browsing was high on most tree species (ca. 60% of available stems browsed) as expected, Peak browsing on lateral twigs of Scots pine (Pinus sylvestris L.) occurred further from feeding stations after 15–20 years of winter feeding than 10 years earlier. Browsing pressure on active feeding stations (n = 18) was comparable to that on inactive feeding stations (n = 12), suggesting the occurrence of rebrowsing. The ability of central-place foraging theory to explain fine-scale browsing patterns around feeding stations decreased as diversionary feeding continued over time. Long-term diversionary winter feeding of moose in unproductive boreal forests has serious implications for the intensity of fine-scale browsing pressure, which may lead to resource depletion close to feeding stations followed by high browsing pressure at distances further away from feeding stations. Our study emphasizes the importance of considering the longevity of a winter feeding programme at its inception.     

Explaining conifer dominance in Afrotemperate forests: Shade tolerance favours Podocarpus latifolius over angiosperm species

We contrasted the regeneration success of angiosperm canopy species with that of Podocarpus latifolius to test whether this conifer was sufficiently shade tolerant to be favoured on infrequently disturbed and well-shaded sites. Seedling and sapling population structures were measured in several habitats representing a light gradient in warm temperate forest in the Drakenberg mountains of South Africa. The angiosperm-dominated forest was well-shaded (∼5.5% PAR) and lacked ground vegetation. Seedlings of angiosperm canopy species were abundant but the absence of saplings indicated regeneration failure. In contrast, P. latifolius was represented by all size classes beneath the intact canopy indicating continuous regeneration in angiosperm-dominated forest. Angiosperm regeneration was similarly poor in forest gaps, which were dominated by grasses, ferns and vines. P. latifolius entered the advanced regeneration in gaps by establishing in shade before gap formation. The floor of the Podocarpus-dominated forest was less shaded (∼7.5% PAR) than angiosperm-dominated forest and dominated by grass, which suppressed most conifer regeneration. Gaps in Podocarpus forest were more than double the size of gaps in angiosperm forest and dominated by vines and understorey shrubs. There was no clear winner of the angiosperm–conifer contest in Podocarpus forest gaps. Consequently, gaps maintain limited angiosperm diversity in Podocarpus forest. Angiosperm canopy species regenerated continuously in the high light (∼11.5% PAR) thicket environment. Few seedlings of P. latifolius were recorded in the thicket environment. Our findings are consistent with the hypothesis that shaded and infrequently disturbed forest sites favour shade-tolerant conifers over relatively light-demanding angiosperm species. The outcome of the competitive interaction between angiosperm and conifer at the regeneration phase depends on the relative shade-tolerance of associated species.     

Photosynthetic characteristics of Fagus sylvatica and Quercus robur established for stand conversion from Picea abies

Efforts in Europe to convert Norway spruce (Picea abies) plantations to broadleaf or mixed broadleaf-conifer forests could be bolstered by an increased understanding of how artificial regeneration acclimates and functions under a range of Norway spruce stand conditions. We studied foliage characteristics and leaf-level photosynthesis on 7-year-old European beech (Fagus sylvatica) and pedunculate oak (Quercus robur) regeneration established in open patches and shelterwoods of a partially harvested Norway spruce plantation in southwestern Sweden. Both species exhibited morphological plasticity at the leaf level by developing leaf blades in patches with an average mass per unit area (LMA) 54% greater than of those in shelterwoods, and at the plant level by maintaining a leaf area ratio (LAR) in shelterwoods that was 78% greater than in patches. However, we observed interspecific differences in photosynthetic capacity relative to spruce canopy openness. Photosynthetic capacity (A₁₆₀₀, net photosynthesis at a photosynthetic photon flux density of 1600 μmol photons m⁻² s⁻¹) of beech in respect to the canopy gradient was best related to leaf mass, and declined substantially with increasing canopy openness primarily because leaf nitrogen (N) in this species decreased about 0.9 mg g⁻¹ with each 10% rise in canopy openness. In contrast, A₁₆₀₀ of oak showed a weak response to mass-based N, and furthermore the percentage of N remained constant in oak leaf tissues across the canopy gradient. Therefore, oak photosynthetic capacity along the canopy gradient was best related to leaf area, and increased as the spruce canopy thinned primarily because LMA rose 8.6 g m⁻² for each 10% increase in canopy openness. These findings support the premise that spruce stand structure regulates photosynthetic capacity of beech through processes that determine N status of this species; leaf N (mass basis) was greatest under relatively closed spruce canopies where leaves apparently acclimate by enhancing light harvesting mechanisms. Spruce stand structure regulates photosynthetic capacity of oak through processes that control LMA; LMA was greatest under open spruce canopies of high light availability where leaves apparently acclimate by enhancing CO₂ fixation mechanisms.     

Effects of non-native spruce plantations on small mammal communities in subarctic birch forests

Voles and shrews are key species in northern forest ecosystems. Thus, it is important to quantify to what extent new forestry practices such as planting of non-native tree species impact these small mammals. In northern Norway stands of coastal subarctic birch forests have increasingly been converted to non-native spruce stands during the last century. This leads to changes in the forest floor vegetation and soil conditions that can be expected to negatively impact the community of ground-dwelling small mammals. In this 10-year trapping study we contrasted seasonal small mammal population abundances in spruce plantations with four birch forest varieties. Six different small mammal species were trapped (in descending order of abundance; common shrew Sorex araneus, red vole Myodes rutilus, field vole Microtus agrestis, grey-sided vole M. rufocanus, pygmy shrew S. minutus and water shrew Neomys fodiens). None of the voles appeared to exhibit temporal dynamics resembling population cycles. The three most numerous species were clearly less abundant in the spruce plantations compared to the other forest types. Autumn abundances were most impacted by spruce plantations, indicating that growth rates in the reproductive season were more influenced than winter declines. Species associated with productive forest habitats (i.e. field vole and common shrew) were most impacted by tree species conversion. Still young spruce plantations inter-mixed with birch trees and the ecotone habitat, sustained small mammal abundances comparable to the native birch forests. This implies that managing spruce plantations to maintain a mix of different tree species and high spatial heterogeneity (i.e. more ecotones), will reduce the negative impacts on the small mammal community. On the contrary, if young spruce plantations, as they age become spruce monocultures covering larger parts of the landscapes than they do presently, the negative effects on small mammal communities may be larger than observed in the present study.