Janusz Szewczyk

Dynamics of seedling banks in beech forest: results of a 10-year study on germination, growth and survival

The dynamics of a seedling bank was studied in four plots located in the Krynica Experimental Forest, southern Poland. Each sample plot was divided into eight sectors, in which germinants and seedlings were mapped, measured, and labelled; their survival was checked twice a year between 1989 and 1996. Maps of relative light intensities on the forest floor in sample plots were produced on the basis of measurements conducted in June and August 1991. During the entire period of study, European beech (Fagus sylvatica L.) made up over 96% of germinants and seedlings. Numbers of beech germinants varied strongly (from 0 to 1178) among plots and years. Each year of abundant germination was followed by one or two years of scarce germination, and there was an increasing trend in the number of germinants in three plots between 1989 and 1996. Seedling emergence was almost independent of relative light intensity and first-season survival was positively, but weakly correlated with it. Long-term survival of seedlings was low and strongly related to relative illumination. Results of analyses of spatial patterns in seedlings suggested that some microsites were more conducive than others to germination and early survival of seedlings. Survival among taller germinants was greater than among shorter ones. A true seedling bank was observed only in two plots, where relative light intensities were 9 and 15% of full sunlight. In a plot with relative light intensity of 4%, the seedling bank was unstable and in the least illuminated plot (<3% of full sunlight) no seedling bank was formed. Results indicate that, even in a strongly shade-tolerant species like beech, formation of a seedling bank requires the occurrence of canopy openings in the close proximity.

Tree mortality and effects of release from competition in an old-growth Fagus-Abies-Picea stand

. In a montane mixed Fagus-Abies-Picea forest in Babia Gora National Park (southern Poland), the dynamics of an old-growth stand were studied by combining an 8-yr annual census of trees in a 1-ha permanent sample plot with radial increments of Abies and Picea growing in the central part of the plot. The mortality among the canopy trees was relatively high (10% in 8 yr), but the basal area increment of surviving trees slightly exceeded the losses caused by tree death. DBH increment was positively correlated with initial diameter in Abies and Picea, but not in Fagus. For individual trees smaller than the median height, basal area increment was positively related to the basal area of old snags and the basal area of recently deceased trees in their neighbourhood, but negatively related to the basal area of live trees. Dendrochronological analysis of the past growth patterns revealed numerous periods of release and suppression, which were usually not synchronized among the trees within a 0.3 ha plot. The almost normal distribution of canopy tree DBH and the small number of young individuals in the plot indicated that stand dynamics were synchronized over a relatively large area and, hence, were consistent with the developmental phase concept. On the other hand, the lack of synchronization among periods of growth acceleration in individual mature Abies and Picea trees conforms more closely to the gap-dynamics paradigm.

Spatial and temporal variability of natural regeneration in a temperate old-growth forest

Abstract• The natural regeneration of shade-tolerant tree species is characterized by large spatial heterogeneity and temporal variability. The mechanisms producing those patterns are still poorly understood and the knowledge of long-term fluctuations in regeneration processes is very limited.• We used data from long-term study plots in an old-growth stand dominated by European beech and silver fir to address three questions: (1) Is a tendency towards clumping in seedlings associated with a particular spatial scale? (2) Are the spatial patterns in seedlings constant over time? (3) Is the distribution of seedlings related to light intensity?• Over the study period the seedling density varied from 2.5 to 6 ind./m2. Stronger fluctuations occurred at a fine scale. Seedlings were clumped at distances ranging from a few centimeters to a few meters. The distribution of places with high seedling densities was relatively constant over time, especially in beech. Positive but weak rank correlations were found between light intensity and densities of beech and fir seedlings.• Despite large fluctuations in seedlings densities, caused by mast-seeding, a permanent seedling bank was maintained. Very pronounced clumping of tree seedlings found in this study was only partially explained by differences in light intensity.Résumé• La régénération naturelle des essences tolérantes à l’ombre est caractérisée par une grande hétérogénéité spatiale et une variabilité temporelle. Les mécanismes de production de ces modèles sont encore mal compris et la connaissance des fluctuations à long terme des processus de régénération est très limitée.• Nous avons utilisé des données provenant d’études à long terme de parcelles, dans une vieille forêt ancienne dominée par le hêtre et le sapin pectiné, pour répondre à trois questions : (1) Y a-t-il une tendance à des bouquets de semis associée à une échelle spatiale particulière? (2) Les modèles spatiaux des semis sont-ils constants dans le temps? (3) La distribution des semis est-elle liée à l’intensité de la lumière?• Au cours de la période d’étude, la densité des semis a varié de 2.5 à 6 ind./m2. De fortes fluctuations se sont produites à une échelle fine. Les plants étaient distribués en bouquets à des distances allant de quelques centimètres à quelques mètres. La distribution des emplacements avec des densités de semis élevées a été relativement constante au fil du temps, surtout avec le hêtre. Des corrélations positives mais faibles, ont été trouvées entre l’intensité de la lumière et les densités de semis de hêtre et de sapin.• En dépit de fortes fluctuations des densités de semis, causées par la chute des graines, un stock permanent de semis a été maintenu. Des bouquets très marqués de semis, retrouvés dans cette étude n’ont été que partiellement expliqués par des différences dans l’intensité lumineuse.

Regeneration processes on coarse woody debris in mixed forests: do tree germinants and seedlings have species-specific responses when grown on coarse woody debris?

ummary Tree regeneration on coarse woody debris (CWD) is considered to be one the most ecologically valuable aspects of CWD in forest systems. However, most studies have focused solely on uncovering the differences in establishment and growth on CWD (regarded as a homogeneous substrate) in comparison with the forest floor. Our study concentrates on the underlying mechanisms of germinant and seedling colonization patterns and demographic responses relative to various properties of CWD. We analysed the effects of CWD properties (decay class, form, species and diameter) on: (i) germinant and seedling annual counts; (ii) annual germinant and seedling survival; and (iii) seedling growth and height. Our study comprised three species (beech, fir and spruce) over 7-year span in two old-growth stands in the Western Carpathians, and employed generalized linear models and mixed models to test for differences between species. CWD properties affected regeneration at the germination stage. There were some demographic differences between species relative to CWD properties. Decay class had the most pronounced effect on beech, not only on its establishment but also survival and growth. Beech and spruce established and survived in higher densities on beech CWD, while their height growth was enhanced on conifer-derived CWD, particularly on spruce CWD. Stumps enhanced establishment of all species and survival of conifer seedlings. Synthesis. Our study shows that CWD properties do influence seedling establishment, growth, height distribution and survival. Moreover, there may be trade-offs between seedling growth and survival among tree species growing on different species of CWD. This highlights the need to include CWD heterogeneity as a factor that can affect the role of CWD in regeneration in forest ecosystems.

High Reproductive Effort and Low Recruitment Rates of European Beech: Is There a Limit for the Superior Competitor?

ABSTRACT European beech is a superior competitor among the trees of Central Europe, often growing in pure stands. We proposed a hypothesis, that once beech has reached dominance in forest community, its recruitment could become limited due to the gradual accumulation of pathogens attacking seeds and seedlings. We employed data on seed production and germination along with a field experiment to estimate the germination success of beech in two old-growth forests. Beech produced more seeds than the co-occurring coniferous trees, but less than 1% of beechnuts germinated in the next season. In the field experiment, the percentage of decayed beechnuts was 57% in the Carpathians and 61% in the Alps. Most of the dead germinants and decayed beechnuts were infested by fungi. The average number of fungal colonies per one sample in the Carpathians was significantly higher after mast year than one year before, while the differences between the Alps and Carpathians after mast years were statistically not significant. Fungi have been isolated from practically all dead beechnuts and dead germinants. The number of beechnuts per seed trap, the number of germinants around it and the relative number of fungal colonies obtained from plastic boxes placed in the same sample plot were not significantly correlated. The mortality of germinants continued throughout the spring; the number of life germinants in the middle of May amounted to 0.87% of the initial number of beechnuts in the Carpathians and only 0.28% in the Alps. High rates of beechnut and germinant mortality could probably offset the huge reproductive effort of European beech in old-growth stands and limit the possibility to attain absolute dominance by that species. However, our hypothesis that the build-up of fungal pathogens on the forest floor old-growth stands is able to stop the regeneration of beech still needs to be tested using larger data sets.

Fine-scale spruce mortality dynamics driven by bark beetle disturbance in Babia Góra National Park, Poland

Abstract Bark beetle outbreaks have had major impacts on Norway spruce forests in Europe. The large majority of these forests are located in areas under forest management; thus, few studies have investigated outbreak-driven spruce mortality patterns unaffected by humans. Our study examined spruce mortality resultant from a beetle outbreak in a high-elevation, unmanaged forest over a 17-year span. We analyzed three tree-level survivorship and DBH datasets collected during pre-, mid-, and post-outbreak conditions to evaluate long-term mortality dynamics. We measured changes in mortality severity, topographic and stand structure characteristics, and stand complexity using ANOVAs, and we assessed five topographic and stand structure mortality predictors by employing boosted regression trees. Our results showed that though spruce mortality increased significantly over time, such increases were disproportionate with spatial synchrony. Moreover, the outbreak did not significantly alter the living stand structure and had little effect on stand complexity, exhibiting the effects of an outbreak that spread throughout the forest without causing major stand-level damage. Larger trees at higher elevations on south-facing slopes were targeted most frequently, particularly during the later stages of the outbreak. Aspect, elevation, and slope were the best predictors of mortality, demonstrating moderate forecasting ability. We showed that bark beetle outbreaks can operate on patch-scale gradients, affecting microhabitat conditions, without resulting in sweeping, stand-altering mortality. Small-scale outbreaks may increase forest resilience against more severe outbreaks in the future by creating canopy gaps that facilitate regeneration, which leads to more complex age and size structures within the stand.

Species composition, elevation, and former management type affect browsing pressure on forest regeneration in the Tatra National Park

Abstract Effects of ungulate pressure on the development of young generation of trees is one of the most important issues in ecology and forestry. Ungulate pressure influence on the development of natural regeneration has been also reported from several national parks. Our study on the effects of ungulate browsing on the young generation of trees was conducted on more than 500 sample plots controlled during one growing season. The overall browsing pressure ranged from 7.6% in seedlings to 20.3% in low saplings. The pressure of ungulates on the regeneration of Picea abies, the dominant species in the Tatra National Park, was by and large below 1%. Broadleaved species were browsed more frequently. The relationship between the plot altitudes and browsing intensity was statistically significant for seedlings and low saplings; at the higher altitudes, the browsing pressure was greater. There was also observed a statistically significant relationship between the type of former management and the browsing degree in seedlings; in the areas subjected to “landscape protection”, the intensity of browsing was higher when compared to strictly protected areas. Pressure exerted by ungulates on tree regeneration was very unevenly distributed, i.e. some plots were heavily browsed and many others - not browsed at all. The most affected tree species were Salix caprea and Sorbus aucuparia, although the percentage of browsed individuals rarely exceeded 50%. Other species favored by ungulates was Acer pseudoplatanus; despite the high browsing pressure, this species was present among seedlings and tall saplings, suggesting that it would be able to recruit to the tree layer. Abies alba was browsed less frequently than the deciduous trees; however, among the tall saplings it was the third most browsed species.