Canopy gaps and regeneration in old-growth Oriental beech (Fagus orientalis Lipsky) stands, northern Iran

Virgin beech Fagus orientalis forests in northern Iran provide a unique opportunity to study the disturbance regimes of forest ecosystems without human influence. The aim of this research was to describe characteristics of natural canopy gaps and gap area fraction as an environmental influence on the success of beech seedling establishment in mature beech stands. All canopy gaps and related forest parameters were measured within three 25 ha areas within the Gorazbon compartment of the University of Tehran’s Kheyrud Experimental Forest. An average of 3 gaps/ha occurred in the forest and gap sizes ranged from 19 to 1250 m² in size. The most frequent (58%) canopy gaps were <200 m². In total, canopy gaps covered 9.3% of the forest area. Gaps <400 m² in size were irregular in shape, but larger gaps did not differ significantly in shape from a circle. Most gaps (41%) were formed by a single tree-fall event and beech made up 63% of gap makers and 93% of gap fillers. Frequency and diversity of tree seedlings were not significantly correlated with gap size. The minimum gap size that contained at least one beech gap-filling sapling (<1.3 m tall) was 23.7 m². The median gap size containing at least one beech gap-filling sapling was 206 m² and the maximum size was 1808 m². The management implications from our study suggest that the creation of small and medium sized gaps in mixed beech forest should mimic natural disturbance regimes and provide suitable conditions for successful beech regeneration.     

Long-term effect of fire on herbaceous species diversity in oriental beech (Fagus orientalis Lipsky) forests in northern Iran

We studied the long-term impacts of natural burning on herbaceous species diversity 37 years after a fire occurred in the Roudbar forests of Guilan Province, northern Iran. Numerous studies have examined short-term changes in understory vegetation following wildfire; however, very few long-term studies are available or changes inferred from retrospective studies based on chronosequences. For this study, 170 ha of forest (85 ha, burned areas; 85 ha, unburned areas) were surveyed. Because the 1000 m 2 plots were too large for detailed measurements of herbaceous species, we determined a sub-sample size according to the Whittaker’s nested plot sampling protocol and minimal areas method. Hence, sub-plots of 32 m 2 were used for herbaceous species measurements, which consisted of percent cover of each species based on the Domin criterion. We measured plant diversity (Shannon-Wiener index), species richness (Margalef’s index), and evenness (Smith-Wilson index). Mean percent cover, together with diversity, richness and evenness, increased markedly in burned areas compared to unburned controls. This suggests that the biodiversity of these forests could be restored within 37 years after fire. However, the abundance of invasive species such as Rubus fruticosus and Bromus benekenii increased significantly in burned areas, but these could be controlled by relevant silvicultural operations.     

Natural regeneration of oriental beech(Fagus orientalis Lipsky)trees in canopy gaps and under closed canopy in a forest in northern Iran

For developing nature-based silvicultural practices in a beech forest to promote, abundance, height,vitality, and preferred growth form, regenerated trees growing in gaps were compared with those under closed canopies.A systematic 50×50 m grid was plotted in a beech stand in the Kheyrud Experimental Forest for selecting trees to measure variables in gaps and under closed canopies.Abundance and mean height of regenerated beech trees were significantly higher in closed canopies than in canopy gaps.Beech seedlings with excurrent growth were significantly taller within regeneration patches under closed canopy.Moreover, regenerated trees with high vitality were more abundant in closed canopy areas than in gaps.Thus, beech regeneration should improve under closed canopies; hence, gaps in a near natural forest should be created only after adequate regeneration and appropriate growth under the parent tree in a closed canopy area is ensured.The results of this research have profound implications for the sustainable management of the forest and for ensuring sustainable beech regeneration.The presence of a closed canopy cover likely will reduce potential stresses on oriental beech regeneration.     

Effect of livestock grazing and human uses on herbaceous species diversity in oriental beech （Fagus orientalis Lipsky） forests,Guilan, Masal,northern Iran

Plant diversity plays key ecological roles in forest ecosystems, including influencing succession, resilience and nutrient cycling. This study was conducted to investigate the effect of livestock grazing and human uses on herbaceous species diversity. We surveyed 50 ha of protected area and 50 ha of unprotected area to evaluate herbaceous species diversity in oriental beech （Fagus orientalis Lipsky） forests in northern Iran. We calculated and compared three indices each of diversity and evenness, and species richness between the two areas. Herbaceous cover was higher in the unprotected area while leaf litter depth and tree canopy cover were higher in the protected area. The diversity indices, H （Shan- non-Wiener index ）, N~ （McArthur index）, N2 （Hill＇s index）, EQ （Modified Nee index）, E,ar （Smith-Wilson index）, E5 （modified index of Hill） and R=S （species richness） and species richness R=S were greater in the protected area than in the unprotected area, suggesting that protection from grazing results in increased numbers of plants and species. The effect of land protection on plant diversity was more pronounced for evenness than for species richness and the positive correlation between diversity and evenness indices was higher than that between diversity and richness.     

A silvicultural management tool for the Oriental beech (Fagus orientalis Lipsky) forests

Stand density management tools help forest managers and landowners to more effectively allocate growing space so that specific silvicultural objectives can be met. Due to the economic importance of Oriental beech (Fagus orientalis Lipsky) forests in Turkey, a stand density management tool was developed for this species to optimize regeneration success rate and tree growth. For the development of this tool, named stand density management diagram (SDMD), we utilized forest inventory data from the Kastamonu Regional Forest Directorate in Turkey. Previously published forest management approaches and models were employed during the development of the tool. The SDMD illustrates the relation among four forest indexes: the basal area per hectare, number of trees per hectare, forest stand volume per hectare, and quadratic mean diameter of the beech stands. The stand stocking percent (SSP) can be determined based upon any two of these four measurements. The results suggest that SSP is a better predictor of tree growth than BA in Oriental beech forests. The newly developed SDMD allows for a more effective use of the growing space to achieve specific silvicultural objectives including tree regeneration, timber production, thinning planning, and wildlife protection in Oriental beech forests.     

Pit and mound influence on soil features in an Oriental Beech (Fagus orientalis Lipsky) forest

Windthrow, i.e. the felling of trees by wind, occurs continually in forest ecosystems. The uprooting of old trees creates multiple microsites (e.g. pit and mound landscape) that are the main source of soil heterogeneity. To determine the impact of pit and mound landscapes on soil features, a beech forest of the Langa district (Mazandaran province, Northern Iran) was studied. An area of 25 ha was considered for this study wherein three microsites were distinguished, including pit bottom (pit), mount top (mound) and level areas (closed canopy). In this area, 22 uprooted trees were also found. Soil samples were collected at different depths (i.e. 0–15, 15–30 and 30–45 cm) from all microsites and analysed. Our study shows that windthrow events should be considered as an important factor in influencing forest ecosystem, as they affect physical (i.e. density, texture and water content), chemical (i.e. pH, organic C, total N, cation exchange capacity and available nutrients) and biological (i.e. soil microbial respiration and earthworm density/biomass) characteristics of soil, thus resulting in pit and mound microsites that may strongly differ with respect to the closed canopy.     

Considering and comparing regeneration in natural and man-made stands in northern Iran

In this study the regeneration diversity of Syahkal forests afforestated and natural stands in north of Iran was studied and compared from the point of view of evenness and diversity index.In order to accomplish this study two natural and man-made stands that almost are located in the same height above the sea level were chosen.The area of each stand was 30 ha and the inventory was done by the random-systematic method with a 5R land measurement(500 m2)selected.Also,in order to study the regeneration in the center of each sample piece 125 square meters micro plots were formed.The results show that regeneration diversity index in natural stands is more than that in man-made stands which in this case Mc-Arthur index with 2.41 has the most amount while Simpson index with 0.543 has the least amount.But the evenness indexes in man-made stands are more than that in natural stand which the main reason is the purity of the man-made stand.     

Stand characteristics of mixed oriental beech (<Emphasis Type="Italic">Fagus orientalis</Emphasis> Lipsky) stands in the stem exclusion phase,northern Iran

The structure of forest stands changes through developmental phases. This study is carried out in the unmanaged, oriental beech (Fagus orientalis Lipsky) stands in the north of Iran. The aim of this research was to quantify structural characteristics of stands in the stem exclusion phase using common structural indices, which include mingling, tree–tree distance, stem diameter, and tree height differentiation. According to our measurements from three stands, naturally regenerated stands tend to be mixed in species composition have slightly heterogeneous diameter distributions and uniform tree height. The average distance between trees was 3.3 m. Stocking volume of the stands had an average of 540 m³ ha^?1 and 412 stem ha^?1. Dead wood volume was 24 m³ ha^?1, and as a standing volume, the most frequent species in dead wood pool was oriental beech (F. orientalis) (48 %). The common form of dead trees was snag (41 %). The mean value of mingling and tree-to-tree interval indices revealed that beech was mixed intensively with hornbeam and appears to be a more successful competitor for space and light compared with hornbeam; moreover, we found relatively high evidence of inter-species competition in this phase. A better understanding of stand characteristics in the stem exclusion phase as a critical part of the natural dynamics of forest ecosystems could facilitate predictions about the future changes within the stand.     

Characterization,pathogenicity and host range studies of Paecilomyces formosus associated with dieback of Christ's thorn trees (Paliurus spina-christi Mill.) in Iran

Christ's thorn (Paliurus spina-christi Mill.) tree is an important and valuable forestry species and has significant potential in afforestation and reforestation systems. In recent years, dieback symptoms were observed on Paliurus spina-christi trees in Gilan-e Gharb region of Kermanshah province, west Iran. During 2020 to 2021, Christ's thorn trees that exhibit branch dieback symptoms were sampled for fungal isolations. The pathogenicity tests, biochemical assay of growth, acid production on creatine sucrose agar (CREA) medium and morphological and molecular investigations (ITS and β-tubulin regions) identified Paecilomyces formosus as a dieback causal agent. This is the first report of P. formosus pathogenicity on Paliurus spina-christi trees. Furthermore, disease severity was assayed on 11 tree species under three different temperatures 15, 25 and 35°C. Disease severity caused by all isolates on detached branches was greater at 35°C than at 15 and 25°C. This study presents the host range of this pathogen and showed that these potential hosts are prone to this pathogen under high temperatures, which forest trees experienced in recent decades.     

Control of rhododendron (Rhododendron ponticum and R. flavum) in the eastern beech (Fagus orientalis) forests of Turkey

Because purple-flowered rhododendron (Rhododendron ponticum L.) and yellow-flowered rhododendron (R. flavum Don.) significantly reduce seedling growth and regeneration of eastern beech (Fagus orientalis Lipsky) as well as local floristic diversity in northern Turkey, effective and cost-efficient woody control is needed. Various manual (cut and grub) and herbicide (foliar and cut-stump spray) woody control techniques were used in two different beech-rhododendron sites in the Black Sea Region (BSR) of Turkey. Two years later, biological and economic effectiveness were substantially greater with foliar-applied herbicide and grubbing treatments than with the cutting, cut-stump, and check treatments. Yet, grubbing showed a great potential for sprout crown reduction. Cut-stump spray was intermediate in effectiveness. Cutting had the least woody control and economic effectiveness among all of the treatments. Foliar-applied Arsenal SL (imazapyr) had significantly greater rhododendron control than foliar-appliied Garlon 4 (triclopyr ester). Increasing rates did not enhance herbicide efficacy. Reducing rates further may provide the same level of control at less cost.     

Spatial and seasonal variations in photosynthetic properties within a beech (Fagus crenata Blume) crown

The light response curve and the intercellular CO₂ concentration response curve of CO₂ assimilation rate were investigated together with the light conditions at the four different heights within the beech crown from 1995 to 1997 on Mt. Fuji in Japan. On the seasonal fluctuation, the CO₂ assimilation rate at light saturated condition increased rapidly in May, and attained to the maximum between the end of June and July, thereafter, slightly decreased until the middle of August and rapidly decreased in September and October. The daily sum of photosynthetic photon flux density attenuated with deeping within the crown, and particularly, the relative value on 2nd position dropped to only 30%. TheA _max decreased from 10 to 5μmol m⁻² s⁻¹, approximately, with deeping within the crown. The light saturation point, quantum yield, light compensation point and dark respiration rate also varied with deeping. These results suggest that the photosynthetic properties vary gradually from sun to shade leaves along the light attenuation within a beech crown. At light saturated condition, the stomatal conductance and mesophyll conductance were strongly correlated withA _max among the four different heights (r > 0.96, respectively). TheC _i/C _a ratio was around 0.8, and there were no remarkable differences among the four different heights. These results suggest that the vertical gradient ofA _max depends on the variation of mesophyll conductance. The stomatal conductance may be also one of the major factor in the vertical gradient ofA _max. However the intercellular CO₂ concentration doesn’t influence the vertical gradient ofA _max within the crown. This work is supported by the Sasagawa Scientific Research Grant from The Japan Science Society and Grant-in-Aid for Scientific Research (C).     

Notes on beech (Fagus crenata Blume) seed and seedling mortality due to rodent herbivory in a northernmost beech forest, Utasai, Hokkaido

We investigated the mortality of post-dispersal seeds and current-year seedlings of a beech species (Fagus crenata Blume) at the northernmost extent of its geographical range at Utasai (Kuromatsunai), Hokkaido, Japan. Introduced seeds that were protected from rodent predation by wire cages had a higher survival rate from winter to the following spring (mean ± SD: 84 ± 16.7%,n = 10) than controls without cages (mean ± SD: 12 ± 17.9%,n = 10). Unprotected transplanted current-year seedlings with almost fully opened primary leaves never survived more than three days (0%,n = 30), while more than 80% (n = 30) of seedlings within cages survived to the end of the growing season. These results indicate that rodent herbivory is a major mortality factor that strongly prevents the establishment of beech seedlings. In a natural population, we observed that 90% of seedlings (n = 197) disappeared within 10 days after their emergence and rodent herbivory caused this heavy mortality. A herbivorous rodent, the gray-backed vole,Clethrionomys rufocanus bedfordiae, was remarkably abundant in late June (101.5 voles/ha), suggesting that this species strongly affects the formation of the seedling bank. The presence ofC. rufocanus bedfordiae may be one of the reasons for the scarcity of beech seedlings and saplings and the rarity of recruitment in this northernmost beech forest.     

Ectomycorrhizal communities within beech (<Emphasis Type="Italic">Fagus sylvatica</Emphasis> L.) forests that naturally regenerate from clear-cutting in northern Spain

The objectives of the work described here were to evaluate the diversity of ectomycorrhizal (ECM) fungi within Spanish beech (Fagus sylvatica L.) forests subjected to clear-cutting and natural regeneration and to elucidate the extent to which the carbohydrate concentration in roots of trees of different ages and sizes is related to ECM colonization. The study concerned an unmanaged forest, a stand clear-cut in 1996 and another clear-cut in 2001. ECM colonization of beech roots showed seasonal dynamics in the disturbed areas, but the percentage of roots colonized by ECM was not always related to the accumulation of non-structural carbohydrates. The composition of ECM communities differed between different stands and 40% of ECM morphotypes only occurred in disturbed ecosystems. However, comparable numbers of different ECM morphotypes (24, 25) were found in the three beech stands. This finding indicates that ECM diversity was quite high and similar within disturbed and unmanaged areas. This finding suggests that (1) ECM diversity was not affected by the size and age of trees and (2) the potential of ECM inocula remained high within clear-cut areas. Consequently, the introduction of ECM inocula by silvicultural practices would not be needed to improve the regeneration of clear-cut areas described in our study.     

Gap formation in Danish beech (Fagus sylvatica) forests of low management intensity: soil moisture and nitrate in soil solution

Soil moisture content (0–90 cm depth) and nitrate-nitrogen (NO₃-N) concentrations in soil solution (90 cm depth) were monitored after gap formation (diameter 15–18 m) in three Danish beech-dominated forests on nutrient-rich till soils. NO₃-N drainage losses were estimated by the water balance model WATBAL for one of the sites. Two forests were non-intervention forests (semi-natural and unmanaged), the third was subject to nature-based management. The study was intended to assess the range of effects of gap formation in forests of low management intensity. In the unmanaged and the nature-based managed forest, soil solution was collected for 5 years and soil moisture measured in the fourth year after gap formation. Average NO₃-N concentrations were significantly higher in the gaps (9.9 and 8.1 mg NO₃-N l⁻¹, respectively) than under closed canopy (0.2 mg l⁻¹). In the semi-natural forest, measurements were carried out up to 29 months after gap formation. Average NO₃-N concentrations in the gap were 19.3 mg NO₃-N l⁻¹. Gap formation alone did not account for this high level, as concentrations were high also under closed canopy (average 12.4 mg NO₃-N l⁻¹). However, the gap had significantly higher N concentrations when trees were in full leaf, and NO₃-N drainage losses were significantly increased in the gap. No losses occurred under closed canopy in growing seasons. Soil moisture was close to field capacity in all three gaps, but decreased under closed canopy in growing seasons. In the semi-natural forest, advanced regeneration and lateral closure of the gap affected soil moisture levels in the gap in the last year of the study.     

Phenological phases of common beech (Fagus sylvatica L.) and their dependence on region and altitude in Southern Germany

Long-term observational data of the phenological network of the German Weather Service (DWD) were used to derive the altitudinal dependence of leaf unfolding (LU), leaf discolouration (LD) and the length of the vegetation period (LD-LU) of common beech (Fagus sylvatica L.) in different regions of Southern Germany. Depending on altitude, the vegetation period starts in Bavaria between the 2nd and 24th May with a delay of around 2 days per 100 m altitude increase (averaged over the last 30 years). In Southern Bavaria, LU starts a few days earlier as well as increases slightly faster into higher altitudes than in the northern regions. This can be explained by a more favourable supply of warmth and radiation in the south. The end of the vegetation period, defined here as the beginning of LD, showed no altitudinal dependence, which reflects an interaction of endogenous and exogenous factors as triggers for LD. In the Bavarian Forest region, altitudinal dependences are strongly influenced by sites which are affected by cold-air tailback (‘Kaltluftstau’) was obtained. Hence, phenological gradients should only be derived and applied above a phenological inversion zone. The calculated vertical gradients for LU can be used together with available observational data to estimate the beginning of the forest vegetation period of beech for different altitudes of Bavaria.     

Dieback and decline of wild almond (Amygdalus scoparia Spach) in the Harat protected forest of Yazd Province,Iran

Wild almond (Amygdalus scoparia) is a dominant shrub species in mountain forests of the Irano‐Turanian region. Dieback and decline symptoms of wild almond shrubs were first observed in Harat protected forest (Yazd, Iran) in the autumn of 2014. Since then, the incidence and severity of the disease have increased. To study the aetiology and estimate losses from the disease, field studies were conducted in 2016 and 2017. For this purpose, eight stands were selected, in which 24%–99% of the trees were wild almond in their species composition. In total, 50 cankers and 50 infected twigs of wild almond shrubs were collected. Fungal isolates were identified based on morphological and cultural characteristics, as well as sequence data of ITS1 + 5.8S + ITS2 rDNA. Pathogenicity of fungi was checked on detached shoots and evaluated after 40 days by measuring the discoloured lesion length at the inoculation site. Among 681 wild almond shrubs checked, 84.44% showed decline symptoms with different severities; only 15.56% did not show observable disease symptoms. The most frequent colonizers of infected tissues taken from cankers were Wilsonomyces carpophilus (56%), Thyrostroma cornicola (40%) and Collophorina paarla (30%). Infected twigs were extensively colonized by W. carpophilus (36%), Th. cornicola (24%) and Ulocladium consortiale (24%). Other species, such as Endoconidioma populi, Microsphaeropsis olivacea, Trichoderma asperellum, Paecilomyces formosus, Saccothecium rubi, Preussia sp. and Chaetomium globosum, had lower isolation frequencies. W. carpophilus, Th. cornicola, C. paarla and U. consortiale were found as pathogens on detached wild almond shoots. Based on the frequency of the isolates and the pathogenicity tests, four fungi, such as W. carpophilus, Th. cornicola, C. paarla and U. consortiale, are considered serious contributing agents playing a significant role in the dieback and decline of wild almond. All isolated species are reported for the first time on the wild almond shrubs of the world.     

Photosynthesis and biomass allocation of beech (Fagus crenata) and dwarf-bamboo (Sasa kurilensis) in response to contrasting light regimes in a Japan Sea-Type beech forest

Regeneration of beech (Fagus crenata) forests depends on the formation of canopy gaps. However, in Japan Sea-type beech forests, a dwarf bamboo (Sasa kurilensis) conspicuously occupies sunny gaps. Therefore,F. crenata seedlings must escape the severe interference ofS. kurilensis in the gaps and persist beneath a closed canopy of the beech forest. We hypothesized that the growth ofF. crenata seedlings in the understory would be favored by their being more plastic thanS. kurilensis in photosynthetic and morphological traits, which would support the matter production ofF. crenata seedlings in a wide range of light availabilities. To examine this hypothesis, the photosynthetic-light response of individual leaves and the biomass allocation in aboveground parts (i.e., the culm/foliage ratio) were surveyed at sites with contrasting light availabilities in a Japan Sea-type beech forest in central Japan. InF. crenata, photosynthetic light utilization efficiency at relatively low light was greater, and the dark respiration rate was smaller in the leaves of seedlings (10 cm in height) beneath the closed canopy than in the leaves of saplings at the sunny forest edge. The culm/foliage (C/F) ratio of theF. crenata seedlings at the shady site was small, suggesting effective matter-production beneath the beech canopy. On the other hand,S. kurilensis both in the gap and beneath the beech canopy showed low plasticity in photosynthesis and the culm/foliage ratio. Because the shoot density ofS. kurilensis was smaller beneath the beech canopy than in the gap, the light availability at the bottom of theS. kurilensis layer was greater beneath the beech canopy. These results suggest that the photosynthetic productivity of theF. crenata seedlings would be enough for the seedlings to survive in the understory with a low density ofS. kurilensis shoots beneath the closed beech canopy.