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.     

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.     

Conversion of conifer plantations to native hardwoods: influences of overstory and fertilization on artificial regeneration

Forest tree species in the eastern US such as American chestnut (Castanea dentata (Marsh.) Borkh) and oaks (Quercus spp.) have been negatively impacted by forest changes over the past century. Many mature, introduced pine (Pinus spp.) plantations exist in the Midwest US following establishment 50–60 years ago yet have little economic and ecological value. As oak and chestnut have similar site preferences to pines, these stands may be ideal sites for hardwood restoration plantings. We sought to determine optimal management strategies for converting pine plantations by manipulating their canopies. We underplanted hybrid American chestnut and northern red oak (Quercus rubra L.) seedlings into three canopy treatments (control, shelterwood, clearcut) and included an open field treatment. For each of two growing seasons, 0, 30, or 60 g 19N–6P–12K of controlled-release fertilizer (CRF) were also applied to seedlings. Soil chemical parameters and leaf nutrients were analyzed throughout the study. Chestnut and oak seedlings had significantly greater height after two growing seasons in the clearcut and shelterwood than the control and open field, and chestnut had significantly greater diameter as well. Chestnut height and RCD growth were threefold that of oak after two growing seasons. In general, fertilization increased seedling growth more in the clearcut and open field than shelterwood and control for both species. Soils had significantly higher pH, K, and S in the open field than in pine stands. Results suggest that pine plantations may serve as target sites for restoration of these hardwood species. Shelterwoods and clearcuts are both favorable conversion options for oak and chestnut, and addition of CRF may augment further growth increase, especially in open environments.     

Effects of different soil treatments on growth and net nitrogen uptake of newly planted Picea abies (L.) Karst. seedlings

Five soil treatments in a 4-year-old clearcut in southern Sweden affected biomass increase and net nitrogen uptake by planted Norway spruce (Picea abies (L.) Karst.) seedlings through their on net mineralisation and root growth. The patch soil treatments studied were: (i) soil inversion in an old clearcut; (ii) mineral soil from the clearcut remaining, (iii) mineral soil from the clearcut with fertiliser application during the first season; (iv) mineral soil from a nearby uncut forest replacing the clearcut mineral soil; and (v) an untreated control. Growth increased in seedlings in treatments (i) and (iii), but growth in soil treatments with humus removal was not better than that of seedlings in untreated soil. High N uptake early in the first growing season resulted in increased growth during this season in contrast to late N uptake that resulted in a high N concentration in the seedlings after the first growing season. This in turn led to a high growth rate during the next growing season. Generally, both root growth and net N mineralisation were positively correlated to N uptake in the soil treatments. Therefore, a combination of low net N mineralisation and poor root growth as a result of high soil density appears to explain the low N uptake in seedlings in undisturbed soil. The importance of competition with field vegetation for N and water was not clear. Net mineralisation was larger in soil treatments where the humus layer was retained than where it was removed. Net N mineralisation in soil from old clearcuts was the same as in soil from fresh clearcuts.     

The influence of silvicultural treatments and site conditions on American chestnut (Castanea dentata) seedling establishment in eastern Kentucky, USA

After more than 50 years of research and selective breeding, blight-resistant American chestnut (Castanea dentata) trees will soon be available for planting into the species’ pre-blight range. Increased understanding of the regeneration requirements of pure American chestnut (C. dentata [Marsh.] Borkh.) will increase the success of future efforts to establish blight-resistant chestnut. We quantified survival and initial growth of bare-root American chestnut seedlings at five locations in eastern Kentucky, USA. We used a split-plot design to compare seedlings planted within adjacent mesic and xeric sites treated with either a two-age shelterwood overstory treatment or a midstory removal treatment. The silvicultural treatments and topographic settings allowed us to evaluate chestnut seedling performance under four light and site productivity combinations. Seedling survival was 57% and seedling height averaged 94 cm following two growing seasons. Seedling survival was negatively related to sand and coarse fragment content, but was unrelated to silviculture treatment or topographic position. Chestnut seedlings grew best in shelterwood overstory treatments areas on mesic sites. Seedlings growing in shelterwood overstory treatment areas added 3- and 3.5-times more height and stem increment compared to seedlings planted after midstory removal. Seedling leaf mass and foliar nitrogen (N) content were also greatest in shelterwood plantings on mesic sites. The high-light environment created by shelterwood overstory removal resulted in better initial seedling growth, but the moderate-light of the midstory removal treatment may ultimately provide chestnut seedlings a greater advantage over competing vegetation.     

Interference of bracken (Pteridium aquilinum L. Kuhn) with Scots pine (Pinus sylvestris L.) and Norway spruce (Picea abies L. Karst.) seedling establishment

The nature of interference of bracken with Scots pine and Norway spruce seedling establishment was considered in three field experiments. In a seeding experiment, it was found that Scots pine germination was highest on exposed mineral soil and lowest when intact bracken litter and humus were present, suggesting adverse effects of litter and humus on pine regeneration probably due to phytotoxicity. In a second experiment, smothering by bracken caused high mortality of Scots pine seedlings while Norway spruce seedlings were relatively unaffected. Mortality for both Scots pine and Norway spruce seedlings was low when planted in a adjacent Scots pine-bilberry stand with no bracken. Annual shoot growth of Norway spruce was higher in bracken than in Scots pine-bilberry vegetation while no differences in shoot growth between these two vegetation types occurred for Scots pine. In a third experiment, activated carbon was added to the ground under Norway spruce seedlings planted in bracken to adsorb possible phytotoxic compounds released by bracken. The addition of carbon had no effect on seedling mortality or growth rate, indicating that the seedlings were not susceptible to allelochemicals released by bracken. Since large Norway spruce seedlings were relatively unaffected by bracken interference in this study, artificial regeneration with containerized Norway spruce seedlings is suggested to achieve an acceptable conifer tree establishment on clear-cuts invaded by bracken.     

Effects of granivorous rodents on direct seeding of oak and beech in relation to site preparation and sowing date

Planting bare-rooted seedlings is a rather expensive method to regenerate oak (Quercus robur) and beech (Fagus sylvatica). Direct seeding could reduce artificial regeneration costs substantially, but the method currently suffers from a number of problems, one of which is seed consumption by rodents. In the present study clear-cut areas in southern Sweden were prepared using one of four different mechanical site preparation methods – disc trenching, patch scarification, topsoil removal or mounding – then manually sown with acorns and beech nuts. Acorns were sown in May and July 2006, and both acorns and beech nuts were sown in May 2007. Three years following sowing, best seedling establishment from viable acorns, 60–70%, was achieved after sowing in mounds in May, and the subsequent survival reached 90–100%. After the first growing season for beech, both seedling establishment and subsequent survival was lower than for oak, which coincides with a higher number of lost beech nuts at the excavation to investigate the fate of seeds. Granivorous rodents (Apodemus flavicollis, A. sylvaticus, Mus musculus, and Myodes glareolus) were live-trapped more frequently in the vicinity of slash piles compared to other micro-site features, such as uprooted stumps, snags and stones. In 2006, no connection was found between number of rodents captured and proportion of lost seeds, while in 2007 the two were in good accordance. No clear relationship was found between number of captured rodents, vegetation cover class and mechanical site preparation method. More rodents were captured in 2007 than in 2006, and there was an increase in captures from spring until late summer/autumn for both years, which was mirrored in the seedling establishment result from the three different times of sowing. Based on experiences from the present study the recommendation for successful direct seeding of oak on clear-cuts is a combination of mounding site preparation, removal of slash and sowing in May rather than July. However, the results indicate that successful direct seeding of beech is more difficult to achieve.     

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.     

Influence of woody and herbaceous competition on microclimate and growth of eastern white pine (Pinus strobus L.) seedlings planted in a central Ontario clearcut

The influence of woody and herbaceous plant competition, either alone or in combination, on microclimate and growth of planted eastern white pine (Pinus strobus L.) seedlings was examined over four consecutive growing seasons in a central Ontario clearcut. Treatments that manipulated the comparative abundance of these two plant functional groups significantly affected light availability, soil moisture, and air and soil temperature regimes. These microclimate alterations, coupled with the relative competitiveness of herbaceous and woody vegetation, corresponded to temporal changes in vegetation cover and dominance. The more rapid colonization and growth of the herbaceous plant community, dominated by bracken fern (Pteridium aquilinum) and ericaceous shrubs (Kalmia sp., Vaccinium sp.), resulted in this form of vegetation being a comparatively important early competitor for soil moisture. As the woody plant community, dominated by naturally regenerated trembling aspen (Populus tremuloides Michx.), grew in height and leaf area, it became a comparatively strong competitor for both light and soil moisture. For all vegetation treatments combined, white pine seedling growth responses were strongly correlated with total cover of competing vegetation and its relative influence on above- and belowground microclimatic variables. Higher total cover of competing vegetation was generally associated with lower light and soil moisture availability and cooler soil temperatures. Multiple regression analyses indicated that pine seedling relative height growth increased with soil moisture content and growing season soil heat sum, while seedling relative diameter and relative volume growth increased with light availability.     

Effects of site preparation on soil properties and on morphology of Pinus sylvestris and Picea abies seedlings sown at different dates

The aims of the study are to find out the effects of site preparation on soil properties and on the morphology of three-year-old Pinus sylvestris L. and Picea abies (L.) Karst. seedlings sown at different dates along a slope with variation in soil texture, water, and fertility. Based on the hypothesis that soil texture affects soil water content, water retention capacity, and nutrient content, we studied site preparation of varying intensities: exposed C horizon, mounds (broken O/E/B horizons piled over undisturbed ground), and exposed E/B horizon. Though the growing time difference between sowings was about one month, after three growing seasons, spring-sown seedlings were larger than summer-sown seedlings. P. sylvestris seedlings were largest with mounds and smallest with exposed C horizon, but site preparation did not affect the morphology of P. abies seedlings.     

Effects of Seedbed Substrate on Moisture Conditions and Germination of Pinus sylvestris Seeds in a Clearcut

The objective of the investigation was to describe moisture conditions and germination of Scots pine (Pinus sylvestris L.) seeds in different seedbed substrates (feather mosses, mineral soil, humus and ground organic material) with and without irrigation during 2 weeks, in a clearcut in northern Sweden. Without irrigation no seeds germinated, which could be explained as an effect of the low moisture conditions of the seedbed substrates. This was shown in the seeds' low moisture content, with highest values in mineral soil, 10-20%, and for the other seedbed substrates 5-15%. The moisture conditions were highest during the night and lowest in the afternoon. With irrigation, moisture conditions improved (seed moisture content 20-50%) and diurnal changes were moderated. Seeds germinated in all substrates with most seedlings in mineral soil (60% of sown germinable seeds). The moisture conditions of the seedbed substrates proved to be in poor agreement with the seeds' requirement for germination, especially during warm weather with no precipitation.     

Impacts of burial and insect infection on germination and seedling growth of acorns of Quercus variabilis

Rodents usually exert important role, through their scatter seed hoarding behavior, on plant regeneration in the field. To investigate the effects of burial and insects infection on germination and seedling growth of acorns of Quercus variabilis, perfect and infected acorns were buried in the soil among four depths, 0 cm, 4 cm, 8 cm and 12 cm, to simulate the seed hoarding behavior by rodents in the field. The results showed that (1) the germination rate were both high, under 4 cm burial depth, for perfect and infected acorns (92% and 53% separately), and decreased significantly with increased burial depth; (2) perfect acorns germinated better in 4 cm burial depth group by autumn of first year; (3) there 32% (perfect) and 26% (infected) acorns with 0 cm burial depth, on the soil surface, germinated successfully; for infected acorns, the 4 cm depth group had the best germination or seedling recruitment in both the first year and the second year; (4) acorns of Q. variabilis exhibited dormancy period ca 7 months; (5) burial, infection, and the interaction between these two factors influenced several aspects including stem height, leaf weight, Tannic acid, and biomass within seedling growth; (6) the results from this study suggest that proper burial would be helpful for the germination and seedling growth, and seedlings of shallow buried acorns had an advantage in their early development; and (7) infection by insects will not inevitably influence seedling early development.     

Influence of overstory density on ecophysiology of red oak (Quercus rubra) and sugar maple (Acer saccharum) seedlings in central Ontario shelterwoods

A field experiment was established in a second-growth hardwood forest dominated by red oak (Quercus rubra L.) to examine the effects of shelterwood overstory density on leaf gas exchange and seedling water status of planted red oak, naturally regenerated red oak and sugar maple (Acer saccharum Marsh.) seedlings during the first growing season following harvest. Canopy cover of uncut control stands and moderate and light shelterwoods averaged 97, 80 and 49%, respectively. Understory light and vapor pressure deficit (VPD) strongly influenced gas exchange responses to overstory reduction. Increased irradiance beneath the shelterwoods significantly increased net photosynthesis (P(n)) and leaf conductance to water vapor (G(wv)) of red oak and maple seedlings; however, P(n) and G(wv) of planted and naturally regenerated red oak seedlings were two to three times higher than those of sugar maple seedlings in both partial harvest treatments, due in large part to decreased stomatal limitation of gas exchange in red oak as a result of increased VPD in the shelterwoods. In both species, seedling water status was higher in the partial harvest treatments, as reflected by the higher predawn leaf water potential and seedling water-use efficiency in seedlings in shelterwoods than in uncut stands. Within a treatment, planted and natural red oak seedlings exhibited similar leaf gas exchange rates and water status, indicating little adverse physiological effect of transplanting. We conclude that the use of shelterwoods favors photosynthetic potential of red oak over sugar maple, and should improve red oak regeneration in Ontario.     

The effects of herbaceous and woody competition on planted white pine in a clearcut site

We investigated the effects of herbaceous and woody vegetation control on the survival and growth of planted eastern white pine (Pinus strobus L.) seedlings through six growing seasons. Herbaceous vegetation control involved the suppression of grasses, forbs, ferns, and low-shrubs, and was maintained for 0, 2, or 4 years after white pine seedlings were planted. Woody control involved the removal of all tall-shrub and deciduous trees, and was conducted at the time of planting, at the end of the second or fifth growing seasons, or not at all. Seedling height and basal diameter responded positively and proportionally to duration of herbaceous vegetation control. Gains associated with woody control were generally not significant unless some degree of herbaceous vegetation control was also conducted. Only herbaceous control increased pine crown closure and rate of crown closure. Herbaceous control and the presence of 5000–15,000 stems per ha of young overtopping aspen were associated with reduced weevil (Pissodes strobi Peck.) injury and increased pine height growth. The study suggests that white pine restoration strategies on clearcut sites should focus on the proactive, early management of understory vegetation and the gradual reduction of overtopping cover from woody vegetation to create a seedling light environment that supports acceptable growth with minimal weevil damage.     

Restoration of hard mast species for wildlife in Missouri using precocious flowering oak in the Missouri River floodplain, USA

Increased planting of hard mast oak species in the Lower Missouri River floodplain is critical as natural regeneration of oak along the Upper Mississippi and Lower Missouri Rivers has been limited following major flood events in 1993 and 1995. Traditional planting methods have limited success due to frequent flood events, competition from faster growing vegetation and white-tailed deer herbivory. Results of early growth response of swamp white oak (Quercus bicolor Willd.) seedlings in relation to initial acorn mass and size, and early rapid shoot growth for seedlings produced by containerized root production method (RPM™), are presented. Containerized RPM™ seedlings grown in the greenhouse under optimal conditions demonstrate that seed size had no discernable impact on first-year root or shoot size. Seedling survival for the first two years and acorn production for the first three years after outplanting are presented, comparing use of containerized RPM™ swamp white oak seedlings to nursery stock. Flood tolerant precocious RPM™ oak seedlings in the floodplain provide a source of food for acorn-consuming wildlife ten to fifteen years sooner than oaks originating from natural regeneration, direct seeding or traditional bare root planting. Compared to bare root nursery stock that produced no acorns, some RPM™ swamp white oak seedlings averaged 4.3, 5.2, and 6.3 acorns/seedling in the first three years after fall outplanting. This revised version was published online in August 2006 with corrections to the Cover Date.     

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.     

Etiology of oak decline in Spain

In different areas of Extremadura, Western Spain, soil samples were taken at the bottom of holm oak (Quercus ilex) trees that were showing decline symptoms. Half of each sample was sterilized, and acorns were sown in both sterilized and nonsterilized soil samples. The resulting seedlings were used as baits for the isolation of fungi. Seedlings growing on the natural, nonsterilized substrate were characterized by having a lower vegetative growth than the ones growing on the sterilized soil samples, and most of them died. Phytophthora dnnamomi was consistently isolated from their roots. Fusarium oxysporum was also isolated as well as different species of Pythium, although to a lesser extent. Pathogenicity tests were performed on holm oak seedlings with five different isolates of P. chinnamomi, with F. oxysporum, Pythium and with a mixture of the three fungi. All the inoculated seedlings with P. chinnamomi developed root rot and grew slowly, and 35.7% of them died up to the end of the experiments. P. chinnamomi was consistently isolated from their roots, indicating that this fungus is the causal agent of holm oak decline. However, F. oxysporum caused similar symptoms on oak seedlings as P. chinnamomi, and was isolated also from the roots, although its frequency was lower than that of. P. cinnamomi.     

Effects of roe deer browsing and site preparation on performance of planted broadleaved and conifer seedlings when using temporary fences

Abstract

The effects of fencing and site preparation on performance of birch (Betula pendula Roth), oak (Quercus robur L.), pine (Pinus sylvestris L.) and spruce [Picea abies (L.) Karst.] seedlings were studied over four growing seasons in southern Sweden. The experiment was composed of four browsing treatments, from which roe deer (Capreolus capreolus L.) were excluded for 0, 12, 24 or 42 months, and four soil treatments. Natural browsing was combined with artificial browsing (clipping). It was demonstrated that browsing by roe deer could be isolated from browsing by moose (Alces alces L.) by selective fencing. Except for birch, browsing had little effect on seedling survival. Browsing reduced height growth by more than 100% for oak and pine, and more than 60% of pine seedling developed multiple stems. Except for oak, site preparation increased seedling survival. Inverting site preparation, in combination with or without fertilization, produced positive growth responses in seedlings, whereas patch scarification did not. There was no interaction between browsing and soil treatments on seedling performance. It was concluded that long-term protection against browsing by roe deer is needed for oak and pine, whereas short-term fences might be used for birch, and that spruce is relatively unaffected by browsing.     

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.     

Modifying the microclimate around young oaks through vegetation manipulation: Effects on seedling growth and branching

The objective of this research was to evaluate the influence of vegetation control on the microclimate (light, air temperature, vapor-pressure deficit (VPD)) and on the growth of young planted, sessile oak seedlings. Three types of vegetation control, creating a range of above-ground interaction intensity, were compared. In the open treatment, vegetation was chemically controlled and was maintained at a low height; in the closed treatment, vegetation was manually controlled and a few dominant individuals that overtopped the oak seedlings were cut; and in the sheath treatment, the vegetation in a 0.5-m diameter circle around each seedling was manually cut to a height equal to three quarters of the total seedling height. In all treatments, a strong reduction of incoming radiation was observed, as well as an increase in air temperature and VPD during the day, and a decrease in daily PET, compared to the reference located 4 m above the vegetation. However, the neighboring vegetation did not modify the microclimate around the seedling to a level that could have induced a significant reduction in seedling photosynthesis and, thus, in seedling growth. Seven years after planting, seedlings were shortest in the open treatment and tallest in the closed treatment (334, 372, 378 cm in the open, sheath and closed treatments, respectively). Seedlings in the open treatment allocated proportionally more biomass to diameter increment than to height growth, compared to seedlings in the sheath and closed treatments. The seedlings averaged 1.77, 1.10 and 1.00 forks in the open, sheath and closed treatments, respectively, and the average height of the lowest fork was 171, 206 and 226 cm in the same treatments. The reduction of the number of forks per seedling was not related to a reduction of the appearance of new forks, but rather to a reduction in the persistence of the existing forks. These changes in stem allometry and in fork development seemed to be related to the lateral shade afforded by the vegetation.