Rodent impact on establishment of direct-seeded Fagus sylvatica,Quercus robur and Quercus petraea on forest land

Abstract

Seed predation by granivorous rodents constitutes a major problem during reforestation using direct seeding. Acorns or beech nuts were sown at 14 different sites in Denmark and southern Sweden, and seedling establishment was inventoried during the first three growing seasons. Around the time of sowing, rodents were either snap-trapped or live-trapped. At some sites, perches for raptors were installed, and at other sites, signs of predation were investigated or predation was monitored by camera surveillance. Granivorous rodents had a profound negative influence on sowings, and the removal of beech nuts occurred rapidly following sowing. Establishment of oak was better when surroundings consisted of mixed forests rather than broadleaved forests. Fewer rodents were caught, and establishment was better, in large reforestation areas. Rodent captures indicated that forest edges, slash piles and stone wall remains were suitable rodent microhabitats. Snap-trapping or raptor perches did not result in increased oak establishment. Neither sowing in summer nor increased seeding depth decreased predation by rodents on beech nuts. In conclusion, seeds need protection from rodents immediately at the time of sowing. Choosing large regeneration areas combined with removal of suitable rodent habitats seems to be a practical alternative for the development of successful strategies for direct seeding.     

Direct seeding of Quercus robur L. for reforestation: The influence of mechanical site preparation and sowing date on early growth of seedlings

Compared with conventional planting, direct seeding of Quercus spp. is a method that can reduce the costs of reforestation considerably. However, interference from natural vegetation and predation on acorns contribute to regeneration failures. Mechanical site preparation has the potential to reduce both these problems. To study the influence of sowing date and of different mechanical site preparation treatments on early seedling growth, an experiment was carried out in storm-felled areas, formerly Norway spruce forests, in southern Sweden from May 2006 to September 2008. Five treatments were applied and acorns were planted in May and July 2006. The treatments were: disc trenching, patch scarification, topsoil removal, mounding and an undisturbed control. A delay of less than two months in seeding resulted in the equivalent of one year's growth reduction with respect to oak seedling's height and biomass. This could be explained by delayed emergence of seedlings, and by the dry conditions during summer seeding. The seedlings only exhibited a weak positive biomass growth response following disc trenching, patch scarification and top soil removal. In the latter case this might be explained by increased soil compaction and extensive removal of the humus layer. Site preparation by mounding resulted in good vegetation control, an increased biomass growth response and deeper seedling roots. In this treatment, we found high relative light levels near seedlings, reduced soil moisture and dry bulk density of the soil and higher soil temperatures. One or more of these environmental factors could have influenced the growth response. Whether this increased early growth rate persists throughout young stand development is, however, uncertain.     

Flying vs. climbing: Factors controlling arboreal seed removal in oak-beech forests

Nuts are heavy and nutritious seeds that need animals to be successfully dispersed. Most studies address nut removal by a single animal species once seeds fall onto the ground. However, nuts are also accessible before the seed drop and usually to a wide guild of seed foragers. This study examines the factors controlling arboreal seed removal in oak-beech forests within the whole guild of nut foragers. We found that seed-dispersing rodents (Apodemus sylvaticus) were the main acorn removers in the oaks (up to 3.75 m height), with a rapid seed encounter and a high removal rate. However, rodents did not climb the beech trees, probably due to their smoother bark in comparison to oak bark and/or the lower nutritional value of beechnuts with regard to acorns. Jays (Garrulus glandarius) were more abundant in oak stands (both dense and scattered) and clearly preferred acorns to beechnuts whereas nuthatches (Sitta europaea) were more abundant in beech stands and preferred beechnuts to acorns. Non-storing birds such as great tits (Parus major) also removed acorns and beechnuts, especially in the stands where oaks are dominant. Jays and rodents preferred sound seeds over insect-infested seeds but such a preference was not found for nuthatches. This study highlights that pure beech stands showed a reduced guild of arboreal nut foragers in comparison to oak stands. This different guild could probably affect the spatial patterns of seed dispersal, with a proportionally higher number of long dispersal events for acorns (mostly jay-dispersed) than for beechnuts (mostly nuthatch-dispersed). Long-distance dispersal of beechnuts (by jays) is determined by the presence of other preferred species (oaks) and their frequency of non-mast years. Seed location in different habitats strongly determines the contribution of different arboreal removers (including climbing rodents) and their removal speed, leading to a differential seed fate that will eventually affect tree regeneration. As nuthatches are sedentary birds, it is important to maintain old and dead trees where they can breed (crevices), forage (arthropods) and store seeds in order to favor beechnut dispersal and gene flow. By maintaining or favoring oak trees within beech stands we will ensure a wider guild of arboreal nut dispersers.     

Diversification of Pinus halepensis forests by sowing Quercus ilex and Quercus pubescens acorns: testing the effects of different vegetation and soil treatments

The goal of this study was to develop management strategies favouring establishment and survival of holm oak (Quercus ilex L.) and downy oak (Quercus pubescens Willd.)??two species co-occurring in Southern France??in mature Allepo pine forests (Pinus halepensis Mill subsp. halepensis). An experimental design was assessed in a partially cut mature stand of Aleppo pine in which five soil and vegetation treatments??chopping, chopping followed by scarification in one or two directions, prescribed burning, control??and two slash treatments (presence/absence) were applied. A total of 1,600 sowing points, each composed of 3 Q. ilex or Q. pubescens acorns, were installed in the different treatments in November of two consecutive years at 6 and 18 months, after the end of treatments. Survival was monitored 3 and 2 years after sowing, soil surface at the sowing points was characterized at different dates, and predawn leaf water potentials were measured during the dry season. High mortality occurred after the first summer, but survival after 1 year was 2.3?C5.2 higher in Q. ilex than in Q. pubescens, confirming that Q. ilex was better adapted to the drier parts of the Mediterranean area. Survival was significantly influenced by the treatments, but there was a variable response between the two sowing years under most of the treatments. Only intense fire proved the most beneficial treatment for seedling survival in both years. The micro-local soil cover conditions induced by the treatments played a major role in explaining oak survival. In particular, grass cover (mainly Brachypodium retusum) proved to be largely unfavourable to seedling survival and growth, and this detrimental effect was also confirmed by lower predawn leaf water potential values with increasing grass cover. Acorn introduction designed to diversify mature Aleppo pine forest after soil and vegetation treatments therefore has to be considered for treatments that most efficiently impair the pre-existing competing grass cover such as prescribed high-intensity fire treatment.     

Germination and early growth of sown Quercus robur L. in relation to soil preparation, sowing depths and prevention against predation

A field experiment was established in the autumn of 1992 in order to study predation of acorns, germination and early growth of oak seedlings. The experiment was established on a clearcut and an adjacent oak shelterwood in southern Sweden. In the experiment, a total of 6 840 acorns were sown. The sowing was carried out with different vegetation control and soil preparation treatments and with different planting depths.The germination percentage was highest for acorns sown at five cm depth and lowest for acorns sown on top of the soil, and lower for acorns sown in areas where the humus layer had been removed than in areas where the humus layer was retained. There was no difference in the percentage germination after two growing seasons between acorns sown inside the shelterwood and acorns sown on the clearcut. The height of the seedlings was lowest when the humus was removed while the number of leaves per seedling was lowest for seedlings in untreated areas after two growing seasons. Furthermore, seedlings in untreated areas showed the lowest relative height growth rate during the second growing season. In spite of higher biomass of ground vegetation in undisturbed areas compared to treated areas, no effect of the soil and vegetation control treatments could be found on soil water potentials. The soil temperature and photosynthetic active radiation at seedling level were higher in areas where soil preparation and vegetation control had been performed. However, it was concluded that neither low soil water availability nor low light levels could be the only cause of lower relative height growth rate during the second growing season for seedlings in undisturbed areas.     

Tree Seed Production and Food Conditions for Rodents in an Oak Wood in Southern England

Seed production and seed losses were monitored in an oak (Quercusrobur) wood in southern England between 1975 and 1988. In additionto acorns, seeds from several large beech (Fagus sylvatica)and sweet chestnut (Castanea sativa) trees which occurred withinthe wood, and from the understorey of hazel (Corylus avellana)trees, were also monitored. The seeds from these four tree speciesare all subject to heavy predation by rodents, as well as otherseed-eaters such as birds and insects. Seed fall tended to be irregular and three years, 1976, 1985and 1987 were considered oak mast years. There were too fewyears with large acorn crops to examine whether resource depletionin these years resulted in poor crops in subsequent years. Therewas no correlation in seed production between the four treespecies which does not agree with one of the predictions ofthe predator satiation theory of masting behaviour. The effectsof weather on seed production were investigated but few significantcorrelations were found. In general the results tended to supportprevious findings. Between 1980 and 1982 a serious outbreakof Tortrix moth caterpillars occurred within the wood and mayhave affected acorn production. A negative relationship wasfound between oak defoliation and acorn production in the subsequentyear. The loss of seeds from the forest floor was initially rapidand in 9 years no seeds survived into the following year andfood conditions were poor for granivores. In 3 of the remaining4 years acorns survived well into spring and early summer providinggood food conditions for forest rodents through to the timeswhen alternative food supplies became available. Although oaktrees contributed most seed energy in good years, the othertree species, especially beech, became important when acornproduction was poor. Predispersal losses due to predators werefrequently high and averaged 80 per cent, 40 per cent, 38 percent and 33 per cent in hazel, sweet chestnut, oak and beechrespectively. On average, hazel nuts tended to fall first, some6 weeks before acorns which tended to be the last seeds to fallof the four tree species. Knopper galls and weeviled acornsfell earlier than sound acorns. A key-factor analysis indicatedthat a failure to mature, predispersal predation and insectinfestation from weevils and Knopper galls contributed equallyto changes in numbers of acorns among years.     

Effect of midstory and understory removal on the establishment and development of natural and artificial pin oak advance reproduction in bottomland forests

Regenerating oaks (Quercus L.) on mesic and hydric sites has remained a problem largely because of inadequate density and poor distribution of large oak advance reproduction prior to harvesting. We examined the effect of midstory and understory removal on the establishment and 3-year development of natural and artificial sources of pin oak (Q. palustris Muenchh.) advance reproduction in bottomland forests in southeastern Missouri, USA. Midstory and understory removals increased the photosynthetically-active radiation (PAR) reaching the seedling layer from about 3 to 15%. This increased light did not increase the density of natural pin oak advance reproduction compared to control, but it increased the survival and nominally increased the growth of the natural pin oak advance reproduction. Where the midstory and understory had been removed, underplanted RPM^® container stock and bareroot pin oak stock maintained high survival, but of the two only the RPM^® stock maintained positive height and diameter growth while bareroot stock suffered some growth reductions. Pin oaks originating from the direct seeding of stratified acorns sown in the spring had low germination and survival, but the survivors had growth rates similar to those of natural seedlings in thinned stands. Applying triclopyr to competitors in the ground flora layer only nominally increased PAR but reduced the percent survival and marginally increased the growth of natural and artificial pin oak. We conclude that artificial reproduction may be used to further increase the probability of achieving adequate numbers of the desired species in the future. Bareroot seedlings may not perform as well as RPM^® seedlings and natural seedlings already present. However, bareroot and RPM^® seedlings remained significantly larger than the natural seedlings after 3 years.     

Acorn defenses to herbivory from insects: Implications for the joint evolution of resistance,tolerance and escape

Acorn predation by insects and its effects on seedling establishment were investigated among three co-occurring oak species (Quercus variabilis, Q. serrata and Cyclobalanopsis glauca) in a subtropical evergreen broadleaved forest, Southwest China. All oak species had high tannin concentration (over 10%) but differed in acorn mass and germination schedule. We hypothesized that the defensive traits in acorns (e.g. seed mass, tannins and germination schedule) act together to reduce damage from insect seed predators. Q. variabilis had significantly lower predation (51.2%) than either Q. serrata (71.4%) or C. glauca (73.8%). Seedling establishment decreased with increasing injury for all oak species studied. As much as 43% of insect infested acorns germinated and established as viable seedlings despite extensive acorn mortality due to insect seed predators. The larger seed mass of Q. variabilis appears to tolerate insect infestation and retain enough stored reserves for seed germination and early growth of seedlings. Autumn germination also benefited Q. variabilis and Q. serrata, allowing escape from both direct and ancillary effects of insect feeding by fast reserve shifting to immediate germination of mature acorns. Our results indicate that large seed mass, tannin and autumn germination act together to ameliorate effects of insect seed predation, and the joint evolution of resistance and tolerance (as well as escape through autumn germination) in acorns may be the selective consequences driven in part by interactions with insect seed predators.     

The development of mice and voles in an oak forest with a surplus of acorns

130?kg of acorns of Common Oak (Quercus robur L.) were distributed on an area of 1.44?ha in an oak forest near Zagreb (Croatia), where acorns were lacking. The influence of this surplus of food on the development of mice and voles was observed from September 1999 till February 2000 on the experimental plot and on an untreated control area before and after offering the acorns. The Redbacked Vole Clethrionomys glareolus was the most abundant and dominant species in both observed areas. After distribution of the acorns, all species of mice, e.?g. the Yellownecked Fieldmouse Apodemus flavicollis, the Longtailed Fieldmouse A. sylvaticus, and the Striped Field Mouse A. agrarius, increased up to 4 times in number on the area treated with a surplus of acorns. On the same area, however, all species of voles, e.?g. the Redbacked Vole C. glareolus, the Field Vole Microtus agrestis, and the Common Vole M. arvalis, decreased. The numbers of captured and recaptured rodents before and after offering acorns on the treated and the untreated control area were compared by statistical analysis (Chi²-test). The results show how food can influence the competitive relations between mice and voles.     

Edge type effect on germination of oak tree species in the Highlands of Chiapas,Mexico

The purpose of this study was to examine the effect of habitat edges on the probability of acorn germination of three oak species (Quercus crassifolia Humb. and Bonpl., Quercus rugosa Née and Quercus laurina Humb. and Bonpl.). The effects of edge type (hard and soft), habitat type (grassland, edges and forest) and leaf litter cover (covered or uncovered acorns) on acorn germination was evaluated by the experimental establishment of acorns along transects crossing habitat edges. More acorns developed into seedlings in grasslands (38%) than in the forest edge (18%) or the forest interior (15%). In sites with soft edges, a higher number of seedlings emerged from acorns covered by litter compared with acorns sowed in the adjacent forest edge and forest interior (P < 0.05). In sites with hard edges, fewer seedlings emerged in the edge (14%) compared with the adjacent grassland (38%), and the adjacent forest (20%) presented intermediate values. However, in sites with soft edges significant differences in seedling emergence were recorded between the grassland (38%) and the forest (10%), whereas the edge presented intermediate values (23%). The effect of leaf litter cover on acorn germination was only significant in grasslands in sites with soft edges (P < 0.05). Acorns in grasslands received relatively little insect damage (10%) compared with the edge (19%) and the forest (30%, P < 0.05), emphasising the importance of acorn dispersal for successful oak establishment. The implications for regeneration of these oak species and the dynamics of montane oak forests in Mexico are discussed.     

Establishment and growth of oak (Quercus alba, Quercus prinus) seedlings in burned and fire-excluded upland forests on the Cumberland Plateau

Recurrent problems with regeneration of oaks (Quercus spp.) have been documented across a wide range of ecosystems. In oak-dominated forests of the central and Appalachian hardwood regions of the United States, a lack of competitive oak regeneration has been tied, in part, to fire suppression in these landscapes, and managers throughout the region are using prescribed fire to address this concern. To examine fire effects on oak regeneration, researchers have generally relied on inventories or population studies of existing seedlings. These studies are valuable but do not permit examination of the role of fire in enhancing the establishment and growth of new oak seedlings stemming from oak mast events. In this study, white (Quercus alba) and chestnut oak (Quercus prinus) acorn mast crops serendipitously occurred in year three (fall 2005) of a landscape-scale prescribed fire experiment. We examined establishment, survival, height and diameter of new seedlings on sites on the Cumberland Plateau in eastern Kentucky. Treatments were fire exclusion, a single prescribed fire (1x-burn; 2003), and repeated prescribed fire (3x-burn; 2003, 2004, and after acorn drop in 2006), all conducted in late spring. Initial densities of newly established chestnut and white oak seedlings were statistically similar across treatments (P = 0.42), despite fires on the 3x-burn site having occurred after acorns were on the ground. Oak seedling density was significantly predicted by oak basal area on all sites (R² = 0.12–0.46), except for chestnut oak on fire-excluded sites (R² = 0.04). Litter depth was less on 3x-burn sites compared to 1x-burn and fire-excluded sites, whereas canopy openness was greater on both burn treatments compared to fire-excluded sites. Seedling mortality was generally higher on fire-excluded sites compared to burn sites, especially for white oak. Oak seedling mortality in the first two growing seasons was significantly predicted by initial litter depth and open sky, with greater litter depth and lower percent open sky leading to higher mortality. In the third growing season none of the measured variables predicted chestnut oak seedling survival; for white oak, percent open sky remained a significant predictor of mortality. Initially, seedlings on the fire-excluded sites had similar height but smaller diameter; after three growing seasons there were few differences in seedling height or diameter among treatments. Our findings suggest a potential role for prescribed fire in establishing forest floor and light conditions that may enhance the success of new oak germinants, although different responses among species may suggest the need to target management for individual oak species.     

Spatiotemporal patterns in seedling emergence and early growth of two oak species direct-seeded on abandoned pastureland

? Fine-scale spatial and temporal establishment patterns of direct-seeded oaks on abandoned agricultural land have been little studied despite their potential importance for long-term stand structure.

? Here we periodically monitored seedling emergence and early growth of bur oak (Quercus macrocarpa Michx.) and red oak (Q. rubra L.) on an abandoned pasture, and tested the effects of herbaceous competition, rodents, and soil physicochemical properties.

? Herbaceous competition slightly decreased diameter growth, but rodents had little impact on establishment. Red oak seedlings emerged earlier than bur oak and in a greater proportion (92% vs. 56%). Seedling emergence and early growth of both species showed significant spatial structures that were partly explained by variation in soil physicochemical properties. Bur oak was more responsive to microenvironmental heterogeneity than red oak, yet much of the variation in emergence and growth of both species remained unexplained.

? This suggests that other factors, such as acorn size or genetic variability, may exert equal or greater control than microenvironmental heterogeneity over seedling emergence and early growth of these two oak species on abandoned pastureland.

     

Development of oak and beech seedlings planted under varying shelterwood densities and with different site preparation methods in southern Sweden

Effects of site preparation, shelterwood density and planting depth on the survival and growth of planted beech and oak seedlings were studied. Experiments were performed in one oak and one beech stand in southern Sweden. Two areas with different densities of shelterwood and one clearcut were established in each stand. Growth, damage and survival of the planted seedlings were observed for three years. Soil water potential was recorded weekly and radiation and soil temperatures were recorded continuously during the growing seasons.Neither site preparation methods nor planting depth affected oak seedling growth, when planting was carried out on fresh clearcuts or in shelterwoods, while growth of beech seedlings was positively affected by mounding. Growth of oak seedlings was inhibited by the shelterwood treatments. In beech seedlings, growth was lowest in dense shelterwood, while there was no difference in growth between seedlings on the clearcut area and in the shelterwood of low density. Differences in growth may be explained by differences in radiation and soil water potential.When planting was carried out on a one-year-old clearcut, site preparation improved the subsequent growth of oak and beech seedlings.     

Sowing methods and mulch affect 1+0 northern red oak seedling quality

The effects of sowing depth, seedling density and mulches on northern red oak seedling survival and growth were evaluated in Wilson State Forest Nursery in southwest Wisconsin, USA. Sowing depths between 2.2 and 6.3 cm, combined with sowing densities of 75 and 150 acorns · m^-2, made up five sowing method plots. Mulch treatments of ground corncobs aged 1 year, hardwood sawdust aged 2 years, and no mulch made up 3 subplot treatments. Treatments resulted in a range of densities from 18 to 148 seedlings · m^-2. Mulch delayed emergence and increased seedling survival. Increasing sowing depth also delayed emergence. Corncob mulch increased root collar diameter; however, hardwood sawdust, aged for 2 years, decreased both root collar diameter and the number of permanent first-order lateral roots. Increasing sowing depth decreased root dry mass but increased shoot dry mass. Increasing density from 18 to 148 seedlings · m^-2decreased root dry mass in this study.     

Date of sowing and emergence timing affect growth and development of loblolly pine seedlings

Loblolly pine (Pinus taeda L.) seeds were sown from February to June in Alabama, U.S.A. If sowing was conducted in May or June, seedling emergence was reduced and seedling mortality increased. Seedlings from earlier sowings grew to greater sizes, but seedlings from later sowings grew later into the fall. Within any given sowing date, the earliest seedlings to emerge grew largest. Seedlings from the February sowing formed buds in mid-summer, flushed, and in some instances flushed yet again. However, by early October, not all seedlings from May and June sowings had formed an initial bud.     

Mechanizsed microsite preparation and direct seeding of Pinus sylvestris in boreal forests — a way to create desired spacing at low cost

The main objective of this study was to examine the cost, flexibility, and appropriate scale of mechanized microsite preparation (MP), in combination with mechanical direct seeding of Pinus sylvestris L. with orchard seed. This technique was tested at four boreal forest sites in Northern Sweden. Orchard and stand seeds were sown with and without MP. The use of orchard seed increased seedling establishment by 41% and the use of MP increased seedling establishment by 47%, respectively, after two years. The best substrates for sowing when using MP were OA_h-, E- and BC-horizon, in ranked order. The use of orchard seed compared to stand seed increased mean seedling height by 25% after four years. These trials suggest that to obtain a density of 5,000 stems ha^-1 four years after seeding, 61,000 viable stand seeds or 41,000 orchard seeds ha^-1 should be sown if MP is not used. If MP is used, seeding rate could be reduced by about 32%. By using MP, and by further improving scarification technique so that all scarified area is thin OA_h-horizon, we predict that only 32,000 stand seeds or 22,000 orchard seeds ha^-1, i.e., half the dosage, should be needed. Under these optimal conditions, it would be necessary to sow about six and four germinable stand and orchard seeds, respectively, to ensure one seedling after four years. Furthermore, regeneration cost would be less than a third that of planting.     

The effects of four repellents on bank vole consumption and germination of beech nuts and acorns

New Forests - Consumption and removal of buried seeds by granivorous rodents is one of the major problems when direct seeding is applied for restoration of oak (Quercus robur) and European beech...     

The effects of microhabitat,plant litter,and seed burial on the regeneration of Quercus wutaishanica and Pinus tabulaeformis

The effects of microhabitat (shrubs and herbs), plant litter, and seed burial on the regeneration of Liaodong oak (Quercus wutaishanica Mayr) and Chinese pine (Pinus tabulaeformis Carr.) were studied in three typical stands (Liaodong oak forest, Chinese pine plantation, and grassland) in the Loess Plateau, China. We monitored the establishment and growth of seedlings of these two woody species in sown experimental plots, in which shrubs and herbs, plant litter, and seed burial were manipulated. In the grassland, shrubs and herbs facilitated Liaodong oak establishment, with no effect on the establishment of Chinese pine. In the two forest stands, shrubs and herbs primarily had an inhibition effect on the establishment of these trees. The effects of plant litter were facilitation or inhibition, depending on the target species and the habitat. Seed burial had a positive effect on seedling establishment. In all three habitats, shrubs and herbs had inhibition effects on seedling growth of both tree species. Plant litter and seed burial did not influence seedling growth in either species. Liaodong oak and Chinese pine use different regeneration strategies during early stages of succession and similar strategies during late stages of succession.     

Germination and seedling growth of Quercus vulcanica: effects of stratification, desiccation, radicle pruning, and season of sowing

Effects of stratification, desiccation, radicle pruning, and season of sowing on Quercus vulcanica germination and growth were studied to identify optimum nursery procedures for artificial regeneration of this species. Following stratification (0, 2, 4 and 8 weeks), acorns were germinated, and acorn moisture content and germination performance were also determined at various times during desiccation. In early December unsprouted acorns were planted in containers, and in early April sprouted acorns, which had been stored in polyethylene bags at 4 °C, with radicles left intact or with radicles pruned back to 1.0 cm were planted in containers.Stratification for 2 and 4 weeks did not increase germination percentage but significantly increased germination rate. Germination percentage of the seeds dropped when the moisture content of the seeds was reduced by desiccation, and the critical minimum moisture content of the recalcitrant Q. vulcanica acorns was found to be 11–16%. Spring sowing of sprouted acorns altered the morphology the containerized seedlings and caused the formation of significantly more main roots, but resulted in significantly less shoot height and shoot dry weight. Spring-sown sprouted acorns had also a significant advantage over the fall-sown unsprouted acorns in seedling survival, and thus nursery personnel should not be concerned if acorns sprout before sowing.     

Silvicultural methods of oak regeneration with special respect to shade tolerant mixed species

An overview is presented of the silviculture of pedunculate oak (Quercus robur L.) and sessile oak (Q. petraea Liebl.) in Germany. This presentation is confined to less dry to moist and stagnic gleysol sites, where the intention is to produce primary timbers in long rotation periods. Incorporation of a shade tolerant species as an admixture species is indispensable to the suppression of epicormic branches. The most frequent and recommended admixture species for this purpose is beech (Fagus sylvatica L.). Because beech is competitively stronger than the oaks on these sites, silvicultural measures must be taken to keep it in check. According to the classic silviculture which has taken shape in the Spessart and Pfälzer forests, the desired goal of a two-storeyed stand with oak in the overstorey and beech in the understorey is achieved by means of heavy seeding beneath an open and rapidly cleared canopy. Because this procedure resembles clear cutting and entails its recognized disadvantages, trials have recently been undertaken to regenerate oaks in a silviculture with permanent canopy cover. Trials with young oaks show that they still achieve satisfactory growth at 15–20% of full light. Accordingly, successful regeneration is possible in beech stands under an open canopy or in gaps, and under an approximately closed canopy cover in pine stands due to their more penetrable crowns. The problematic aspect of this, especially in beech stands, is that young beeches become competitively stronger than oaks as canopy cover increases. This requires great effort in restraining the beech during cleanings and thinnings. Furthermore, browsing by wild animals must be minimized as it exclusively affects oak. As a result, however, there will be significantly fewer oaks and more beeches in the dominant layer than when using classic methods.