Regeneration patterns of Eastern white pine (Pinus strobus L.) in hardwood-dominated forests in Virginia, USA

Eastern white pine (Pinus strobus L.) is a moderately shade-tolerant species that co-occurs with hardwood tree species in many forests of the eastern United States, as well as in pure stands. The species is valued for its timber, as well as for wildlife and recreation. Regeneration of this species is somewhat unpredictable and often occurs in patches of similarly-aged cohorts. We described the regeneration patterns of this species and examined their relation to environmental variables within hardwood forests of southwestern Virginia, USA. An average of 5.3 white pine patches per ha were observed in this study. The majority of patches consisted of saplings (85%), with 9% of patches in pole size classes, and 6% in seedling size classes. The average density of patches was 43.5 stems with an average age of 20 years. The size of patches averaged 80.6 m². The total density of seedlings and the number of regeneration patches of all sizes of regeneration (seedlings, saplings, and poles) in plots was related to the surrounding density of large white pine trees (potential seed trees). The density of seedlings or patches was not significantly related to current vegetation cover or soil surface cover variables, but more than half of regeneration patches were located in or adjacent to old canopy gaps, most of which were old logging gaps. While seedling regeneration may occur within the understory of these forests near seed trees, advancement to the sapling and pole stage appears to be associated with canopy gap formation.     

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.     

Wild pistachio tree (Pistacia mutica) in the Qalajeh forest region of western Iran

Wild pistachio tree (Pistacia mutica) is considered the most important species in the Zagros forests. It can adapt to unfavorable environmental conditions. To find the suitable ecological conditions for pistachio species, we investigated different environmental factors such as gradient, aspect and position of the slopes through the distribution area. Frequency of pistachio trees in the north and northwest of Qalajeh forests was 36.4% and 1%, respectively. Maximum (32%) and minimum (13%) number of wild pistachio trees were in 30%–60% and >120% slope classes, respectively. The most number and least number of pistachio trees were found at elevations of 1100–1200 m and 2200–2300 m, respectively. The percentage of pistachio canopy coverage was 10. The average percentage of herbaceous coverage was 6.5. Pistachio trees of diameter per hectare at breast height (DBH) >20 cm numbered 5 while average number of pistachio trees was 8 per hectare. Pistachio seedlings per hectare averaged 3.5. The number of other species per ha was 7 trees. The effect of aspect on other species was not significant (p=0.151). Slope class and geographical aspect did not affect regeneration (p=0.275 and p=0.111, respectively). Pistachio plays an important economic role in semi-arid areas, therefore it is essential to protect and restore Qalajeh forests through participation by government and local people.     

Effective height development of four co-occurring species in the gap-phase regeneration of Douglas fir monocultures under nature-oriented conversion

Natural regeneration in gaps in Douglas fir forest stands in the Netherlands mainly consists of Betula pendula (Roth.), Pinus sylvestris (L.), Larix kaempferi (Carr.), and Pseudotsuga menziesii (Mirb. Franco). Even though these species are well known, the autogenic development of these species in an unmanaged plant community falls outside of traditional silvicultural experience. We therefore present an explorative study, aimed to map out differences between these species under conditions of co-occurrence, and develop hypotheses on the further development of these sapling communities. We focused on differences in height, and particularly the architectural contribution to height.We analyzed interspecific differences in the height–mass and height–DBH relationship, biomass partitioning, and slenderness for a total of 177 saplings up to 13.7 cm DBH and 11.8 m high. Average annual height and diameter growth rates of an additional 48 saplings are presented for comparison with the static architectural data.Betula had a high stem biomass, high stem slenderness, and low leaf mass. The reverse was true for Pseudotsuga. Larix behaved as an intermediate species. Pinus however deviated from the classical pioneer–climax dichotomy in having a low slenderness and low height per unit biomass while being a highly light-demanding species. The static height-mass relationship shows a similar species ranking when compared to average annual height growth rates, but not when compared to diameter growth rates. This underlines the importance of a species’ architecture for its height development and suggesting a predictable stratification between species.Based on the results regarding species’ stratification in heights and a priori information on light demand, it is hypothesized that in the studied gap-phase regeneration Betula, Larix and Pseudotsuga will partition the available light. Pinus however will be overtopped and given its need for high-light environments, is expected to be competitively excluded.     

The effects of gap size and age on natural regeneration of Picea mongolica in the semi-arid region of Northern China

The aim of this work was to study the effects of gap size and age on natural regeneration of endangered tree species Picea mongolica W. D. Xu, which is only found in the semi-arid region of Northern China. For this purpose, six random transect belts of varying size were established in Baiyinaobao Natural Reserve. The number, height and age of P. mongolica saplings were inventoried. Results show that the total number of saplings per hectare and corresponding number and relative share of saplings in height class I (0.1–0.99 m) decreased along with increasing gap size, opposite to height class III (6–9 m). In height class II (1–5.99 m) the number of saplings per hectare and their relative share increased first slightly and then decreased with the increase of gap size. The number of saplings and their relative share in each height class changed similarly with the increase in the distance to the edge of the mature stand (p < 0.05). Gap age also significantly affect the number of saplings per hectare or their height distribution except in height class II. The distance to the mature stand edge played a very important role in the success of natural regeneration for species P. mongolica.     

印度加瓦尔喜马拉雅山的亚热带的宽叶榆绿木(Anogeissus latifolia)林恢复状况(英文)

研究了位于印度加瓦尔海平面(950～1100m)上的亚热带森林的再生生长状况。对生长在不同方位(东、西、北、南)研究地的植被进行了定量分析。研究结果表明:宽叶榆绿木(Anogeissus latifolia)的树木、幼树和幼苗层在研究地点的所有方位显生长优势。长叶松(Pinusroxburghii)和毛白杨(Terminalia tomentosa)的树木、小树在研究地点的北部显生长优势。树木层的最高(380株·ha-1)和最低密度(260株·ha-1)分别出现在南坡和北坡。幼树的最高(1790株·ha-1)和最低密度(970株·ha-1)分别出现在东坡和西坡。树木和灌木层的多样性分别为0.846～1.710和1.943-2.847。与其它种的相比较(4%～33%),宽叶榆绿木(Anogeissus latifolia)的相对剪枝强度较高(45%-57%),此种是在北坡中最重要的树种。目前研究表明,如果不合理开采继续进行,宽叶榆绿木(Anogeissus latifolia)就有可能被其他种代替,在树种的成分和森林重建方面会发生剧烈的变化。人类活动的压力,地理方位和土壤营养已引起了树种的成分和森林重建方面的变化。     

Comparison of ecological condition and conservation status of English yew population in two Austrian gene conservation forests

English yew Taxus baccata L. has been catalogued as endangered tree species and prone to extinction in Austria as well as many other parts of Europe. The present work is based on the comparison of the natural population of two gene conservation forests from different geographic locations in Austria where the spatial structure, regeneration status and possible conservation measures are examined. The pole stand distribution varied distinctly in each sites. The total no of individuals per ha （DBH ≥5 cm）, average DBH and average height were 492 n.hm^-2, 8.8 cm and 6.3 m in Stiwollgraben whereas in Leininger Riese 45 n.hm^-2, 16.3 cm and 7.6 m respectively. Over 79% of the Stiwollgraben population were represented the good health condition, while in Leininger Riese it was less then 49 % which means population of Stiwollgraben is in better condition compared to Leininger Riese. The sites differed considerably in the pattern of regeneration but pattern were consistent with the dynamics depicted by the age distribution. Considering the one-year-old seedlings Stiwollgraben contains 13 019 individuals.hm^-2 whereas Leininger Riese only 1 368. Surprisingly there were no any saplings in respect of 51 to 150 cm height classes in both sites and 30 to 50 cm in Stiwollgraben. In that context the conservation of English yew on the forest level may require well-managed reserves and long-term rotations between harvest events, protection from the herbivore and reduction of competition, which will enhance the long-term viability of the species.     

Leaf conductance of primary and mature leaves ofPinus roxburghii: A comparison

We used a null-balance porometer to measure leaf conductance of mature and primary leaves of natural seedlings, saplings, and trees ofPinus roxburghii (chir) during autumn at four sites at 1,320–1,930 m elevation in the central Himalaya of India. Our hypothesis that primary leaves had higher leaf conductance than mature leaves (needles in fascicles), based on measurements in other pines, was rejected. Comparisons on the same saplings and seedlings showed lower leaf conductance for primary leaves than for mature leaves. Primary leaves on seedlings and saplings also did not consistently have higher leaf conductance than mature leaves on near-by trees. Mature leaves on seedlings, however, did often have higher conductance than mature leaves on nearby trees. Mean values for leaf conductance (mmol m⁻² s⁻¹, based on total leaf surface area) ranged from 42–82 for mature leaves on trees, 60–121 for mature leaves on seedlings, and 28–67 for primary leaves on seedlings. Compared to published values for other pine species, conductance of mature leaves ofP. roxburghii is relatively low. Funding for this study was provided by grants from the United States National Science Foundation, INT-9312052 and INT-9404043. We appreciate the review of an early draft by Barbara Bond.     

Land-use impacts on the composition and diversity of the Baikiaea– Guibourtia–Pterocarpus woodlands of north-western Zimbabwe

Variation in floristic composition of recovering Baikiaea–Guibourtia–Pterocarpus woodlands was studied in different development stages (from early regrowth to mature woodland) under different land-use systems (protected areas, timber harvesting, pole and firewood collection, and abandoned crop fields), in the Gwayi and Tsholotsho areas in north-western Zimbabwe. A total of 150 nested circular plots were sampled representatively in four different development stages related to the land-use systems. The DBH (stem diameter at 1.3 m above ground level) and tree height were recorded by species for all stems of tree species with DBH ≥ 15 cm in a 30-m-radius plot (0.283 ha) and for trees with DBH 5.0–14.9 cm in a 11.3-m-radius plot (0.04 ha) (both centred around the same midpoint). Stems with DBH < 5 cm were counted by species in an inner sub-plot of 5.65 m (0.01 ha) radius. Tree data (stem DBH ≥ 5 cm) and regeneration data (stem DBH < 5 cm) by stem counts per species per plot, were used separately to run TWINSPAN (TWo-way INdicator SPecies ANalysis) classifications of species assemblages. Importance values were calculated for all tree species per community. Shannon–Wiener diversity indices were calculated for each community and tested for differences using one-way ANOVA in SPSS version 21. Detrended correspondence analysis (DCA) implemented in the CANOCO ordination software was used to determine the extent of variation amongst the identified communities. The classification identified 12 tree communities and 13 regeneration communities, clustering plots from different land uses together. Baikiaea plurijuga was the most important tree species in all tree communities, except where Combretum collinum, C. apiculatum, Commiphora mossambicensis and Pterocarpus angolensis were dominant. Pterocarpus angolensis showed low importance in most communities, except for communities from undisturbed sites (mostly mature trees) and abandoned fields (mostly young trees). Baikiaea plurijuga was most important in most regeneration communities, except in communities dominated by Baphia massaiensis, C. collinum, C. apiculatum and P. angolensis. Species diversity differed significantly (p < 0.05) amongst tree communities. The DCA ordination showed little variation amongst the communities. The cumulative contribution of environmental factors explaining variation in species composition was 22.6% for tree communities and 26.1% for regeneration communities, suggesting that recovery from disturbance after different land uses may explain more of such variation.     

Phytosociology,structure and dynamics of Pinus roxburghii associations from Northern Pakistan

We investigated the phytosociology, structure and dynamics of Pinus roxburghii in 40 stands in northern areas of Pakistan by using cluster analysis （Ward’s agglomerative clustering） and ordination （Non-metric Multidimensional Scaling）. Cluster analysis revealed three major groups associated with specific environmental characteristics： （1） P. roxburghii （2） Pinus-Quercus baloot and （3） Pinus-Olea ferruginea community types. NMS-ordination showed the major gradient as an amalgam of elevation （r2=0.441, p＆lt;0.01） and slope （r2=0.391, p＆lt;0.05） as the two topographic factors correlated with species distribution. The first ordination axis also showed positive correlation with soil variables like pH and electrical conductivity, suggesting that soil chemistry was related to topographic characteristics and probably acted as a secondary gradient. We also examined size class distributions, age structures and growth rates of the three communities in order to describe community development and dynamics. Total tree density was 14700 plants/ha, with P. roxburghii having a relative density of 82%to 100%. Density of juve-nile and total density and basal area of the subordinate tree species were low. The low density of trees in the smallest diameter size-class sug-gested that the recruitment of small P. roxburghii plants into the adult population may be lower than the required replacement rate for the stands. Pooled size-class distributions for the species showed a multimo-dal pattern with some regeneration gaps. Browsing, heavy logging and＆amp;nbsp;other anthropogenic activities were the overriding factors responsible for the poor recruitment of P. roxburghii. We concluded from the age struc-ture that the forests were characterized by the dominance of young trees. Growth rate analysis revealed that P. roxburghii was the fastest growing species among the conifers species in Pakistan. In view of its relatively fast growth and longevity, P. roxburghii seems to be a suitable choice for short-t     

Scots pine stand establishment with special emphasis on uncertainty and cost-effectiveness,the case of northern Finland

The effect of uncertainty on the cost-effectiveness of alternative chains of stand establishment in northern Finland was examined. The data were from a reforestation study of Scots pine (Pinus sylvestris L.) consisting of 288 sample plots, which were measured with respect to regeneration success. The study design included four site-preparation methods (patch scarification, ploughing, prescribed burning and disk trenching) combined with three reforestation methods (sowing, planting with containerized seedlings and planting with bare-rooted transplants). Initial reforestation density was 2,500 spots or seedlings per hectare, and the regeneration success was modeled as probability with two thresholds, namely 500 and 1,100 saplings. On formerly spruce-dominated as well as pine-dominated sites the most cost-effective chain was ploughing and planting with containerized seedlings, when threshold was 1,100 saplings per hectare. However, with threshold of 500 saplings the best performer was ploughing and direct sowing on both sites.     

Restoring silvopastures with oak saplings: effects of mulch and diameter class on survival,growth, and annual leaf-nutrient patterns

In Southwestern Spain, multifunctional silvopastoral systems consisting of pastureland and open oak woodlands are known as Dehesas. These, and other similar systems of the Mediterranean basin, are currently threatened by increasing intensive land use. As a consequence, oak regeneration is declining and is in need of adequate management and active restoration. Traditional restoration practices outplant one-year-old, nursery-produced oak seedlings grown in 250–350 cm³ containers, but establishment and growth results are typically poor. This work examines holm oak (Quercus ilex L. subsp. ballota (Desf.) Samp.) grown in a non-conventional container size (24 l) and age (6–7 years) with three mulch treatments (control, stone, and straw). In an open Dehesa of SE Spain, 106 oak saplings were planted in March 2010, and survival, diameter at breast height (DBH), and leaf-nutrient concentrations were analyzed. Forty months after planting, all treatments showed high survival (81 %) but only straw-mulched saplings differed significantly (94 %) from control (74 %). DBH increased over time but showed no significant differences among mulch treatments. Saplings with high initial DBH showed the greatest growth and change in DBH at the end of the study period. Leaf-nutrient concentrations changed significantly in the year following outplanting. Bi-monthly foliar nutrient concentration data show decreases in P, K, Zn, and B and sharp increases in Ca and Fe. In this work, we provide some evidence concerning the viability of non-conventional oak size for restoring, regenerating, or building up new agroforestry or silvopastoral systems. A combination of saplings with more than 10 mm of DBH and straw mulch is recommended.     

子午岭油松林林隙更新特征研究

利用样地调查资料,以林下为对照,分析研究子午岭油松林的林隙更新特征,结果表明:1)子午岭油松林林隙内和林下更新树种在组成和数量上有较大差异,其中,油松在林隙内的重要值远远大于其在林下的重要值,是林隙更新的主要树种;2)油松幼苗幼树在林隙内生长良好,7年生以上的幼树在林下分布已很少,地径在1.0cm以上、高度在50cm以上的幼树数量分布在林隙内也远远大于林下,说明随着年龄的增长,油松幼苗幼树在林隙内存活的概率大于林下;3)方差分析表明,油松幼苗幼树在林隙内与林下的高度生长差异极为显著;4)相关性分析表明,林隙内油松幼苗幼树的更新数量与光照强度呈显著正相关,与植被总盖度呈显著负相关。     

Effects of management, environment and landscape conditions on establishment of hardwood seedlings and saplings in central Japanese coniferous plantations

There is an increasing need to restore natural hardwood forests in landscapes dominated by monocultural conifer plantations. A convenient restoration approach is to exploit natural regeneration processes. Natural regeneration, however, is affected by diverse interacting factors, for which better understanding is required, in order to optimize restoration programs. To identify optimal management practices for improving natural regeneration of hardwood trees in coniferous plantations, we examined the effects of multiple factors on the abundance of seedlings, small saplings and large saplings (height <0.3, 0.3-1.3 and ?1.3 m, respectively) of hardwood tree and shrub species in both line thinned (LT) and unthinned (UT) plantations of sugi (Cryptomeria japonica) and hardwood forests (HF) in central Japan. The effects of management practices (number of the times of weeding and cleaning, thinning method, years after thinning and forest age), environment (slope position, slope angle and canopy openness), and landscape conditions (distance from nearest hardwood forest, altitude and landuse before planting) on the number of hardwood individuals were examined by using the data obtained from the LT plantations. We also compared hardwood density between LT and UT plantations to examine the effect of line thinning. Finally, we examined species composition of LT plantations and HF to identify hardwood forest components in the thinned plantations. The effects on hardwood regeneration of environmental conditions, landscape factors and management practices applied in the plantations varied, depending on the size class and life form of the regenerating species. The abundance of large saplings of tall tree species was affected by several management factors, especially number of the times of weeding. Landscape conditions (distance from the nearest hardwood forest and altitude) affected the abundance of small saplings and seedlings of tall tree species, but not the other classes. Seedlings and small saplings of many tall tree species that contribute to hardwood forest canopies were less abundant in the LT plantations. The results show that numerous factors affect the establishment and abundance of naturally regenerating hardwood tree species, and suggest that successful establishment during early plantation stages can have long-lasting effects on natural regeneration of tall tree species.     

Regeneration pattern of four threatened tree species in Sudanian savannas of Burkina Faso

Research was carried out on four threatened tree species across north and south Sudanian phytogeographical zones in Burkina Faso. A quantitative inventory of regeneration populations was completed and sapling dynamics were monitored through surveys of survival and growth in permanent plots. Laboratory and nursery experiments were carried out to test germination and the performance of transplanted seedlings. Results revealed difficulties that saplings faced during their development. In natural stands of Sterculia setigera and Afzelia africana, saplings were rare or irregularly distributed. A greater number of individuals of Bombax costatum were observed, which was the only species with significant density variation between phytogeographical zones (p?=?0.021). Size class showed irregular population structures. A high mortality between the first and second year, followed by a stabilization trend, was observed. The initial height of saplings was highly correlated to their survival rate. The rate of new sapling spontaneous recruitment was better in Bombax costatum and Boswellia dalzielii stands, but heir growth was unpredictable in all species. Most saplings grew from lignotubers and root suckers; however, the potential for sexual reproduction was good. Sterculia setigera and Bombax costatum seedlings showed potential for use in silviculture, as evidenced by the fact that both showed high post-transplantation rates of survival and growth.     

Foliar pathogen and insect herbivore effects on two landslide tree species in Puerto Rico

To better understand pathogen/herbivore interactions and landslide regeneration, percent leaf area lost to disease and herbivory on two Puerto Rican trees over a 1-year period was sampled. Cecropia schreberiana saplings lost from 1 to 3% leaf area to pathogens and from 1 to 7% to herbivores. For Inga vera, both sapling and seedling losses to pathogens were minimal, but Inga herbivory losses reached 25% for saplings and 34% for seedlings. The most common fungi on Cecropia leaves were species in the genera Phoma and Phyllosticta, and on Inga leaves was Colletotrichum gloeosporioides. Percent survivorship after 1 year in the field varied among species and life-form (46% for Cecropia saplings, 15% for Inga saplings, 0% for Inga seedlings). There were no effects of pathogens or herbivores on survivorship or growth, but increased levels of herbivory did significantly correlate with total phenolics and condensed tannins in both Inga seedlings and saplings. For both seedlings and saplings of two trees on the neotropic island of Puerto Rico: (1) leaf herbivory was modest and leaf losses to pathogen disease were small; (2) these mechanisms did not affect survivorship or growth; (3) a neotropical tree (I. vera) displayed increased levels of secondary chemicals in its leaves, correlated with increased levels of insect herbivory.     

Regeneration of Norway spruce in canopy gaps in Sphagnum-Myrtillus old-growth forests

Gap-associated spruce (Picea abies (L.) Karst.) regeneration in Sphagnum-Myrtillus stands of south taiga forests (Central Forest Biosphere reserve, Tver region, Russia) was studied to evaluate the role of different disturbances in spruce dynamics. Sampled gaps (n=70) ranged from 40 m² to 1.7 ha in size, and from 1 to 70 years since disturbance moment. Formation of gaps lead to increase in the number of stems per ha in all gap size classes (small: 40–200 m², medium: 200–3000 m², and large: >3000 m² gaps). Spruce was the most important species in gap refilling, although its role was not the same in different gap classes. The highest values of relative abundance (compared to other species) were recorded in small gaps, and much lower values – in middle and large gaps. However, as refilling of gaps proceeded, spruce showed rather active regeneration in middle and large gaps and partly regained its abundance in middle-age disturbances. In general, all types of gaps studied supported spruce regeneration into the forest canopy. Almost perfect correlation between predicted outcome of spruce dynamics in gaps and its current role in the canopy of Sphagnum-Myrtillus stands suggests a good adaptation of this species to the current disturbance regime and a steady state of the these forests.     

浸泡和冷预处理对喜马拉雅长叶松不同种源种子萌发的作用

对喜马拉雅长叶松种子进行24小时GA3和H2O2浸泡预处理和15d的2-3℃低温预处理后,研究了在20℃,25℃和30℃萌发条件下,21个不同种源的喜马拉雅长叶松种子的萌发情况。结果表明,H2O2（1%v/v）和GA3（10mg/L）浸泡预处理,种子萌发率分别是82.39%和78.19%,而未经预处理的种子平均萌发率为70.79%。GA3和H2O2浸泡预处理分别使种子萌发时间缩短了8d和10d。在超过21天的20℃萌发条件下,湿冷处理提高了种子萌发率和缩短了萌发时间 而在25℃和30℃萌发条件下,总的萌发率未受到影响。喜马拉雅长叶松种子因缺少休眠而表现出很好的萌发,但是因为越来越多的造林项目需要大量的种子,播种预处理有利于提高种子萌发率和萌发速率,有助于满足种子需求。     

Browsing damage on broadleaved trees in semi-natural temperate forest in Sweden,with a focus on oak regeneration

Absence of, or poor, oak (Quercus spp.) regeneration is a problem in uneven-aged, mixed closed-canopy broadleaved forests. Browsing by ungulates on small trees may contribute to poor oak regeneration in such forests. This possibility was investigated in 25 Swedish stands, and browsing damage was analysed in relation to landscape and stand factors. The proportion of browsed small (<20 cm tall) oak seedlings and other seedlings was low, and apparently a minor mortality factor. For saplings (20–130 cm tall), accumulated browsing damage was generally higher on oak than on five major competing tree species: Fraxinus excelsior, Corylus avellana, Tilia cordata, Acer platanoides and Sorbus aucuparia. Leaf removal was rare in late summer, except for rowan. The amount of cover (shelter) for ungulates near plots was positively correlated with oak browsing intensity; within plots, a high density of ash saplings may reduce browsing on oak saplings. In these forests, browsing probably retards growth of oak saplings relative to competing trees. Oak may persist as a minor stand component, but monitoring is needed to study future changes.