Soil Ripping and Herbicides Enhance Tree and Shrub Restoration on Stripmines

I evaluated responses by 16 native woody species to differential soil compaction and density of ground cover. The trees and shrubs studied represent sites in southern Illinois that commonly have restrictions to root growth from soil or drainage conditions. The study site was a restored surface coal mine in southern Illinois with a rooting medium compacted by grading and a dense ground cover of pasture species. Soil compaction was alleviated in half the study area before tree planting by mechanically ripping the soil to a depth of 1.2 m. Roots of half the trees and shrubs were dipped in a Terra® slurry before planting, and the ground cover around all planting spots was afterwards sprayed with herbicide. In year 2 after planting the ground cover in half of the unripped and half of the ripped area was further controlled by repeated application of herbicides. Ripping significantly increased height growth of all trees combined and all species individually in each year of the study. Second-year control of ground cover increased height growth of all trees combined and of seven species individually. Some species were damaged by herbicides. Terra® had little evident effect on species performance. Animal damage reduced early survival and growth, especially of Acer (maple) and Cornus (dogwood) species, and later growth of Quercus rubra (red oak). Removal of ground cover with herbicides tended to increase deer browse. Soil ripping, herbicide application, and choosing tree species unattractive to deer can be recommended to increase success in planting trees for forest restoration.     

Effects of Deer Browsing, Fabric Mats, and Tree Shelters on Quercus rubra Seedlings

We examined the effects of deer browsing, fabric mats, and tree shelters on the growth and survival of Quercus rubra (northern red oak) seedlings planted as part of a reforestation project in southcentral Minnesota. Browsing by white-tailed deer occurred on 68.6% of the seedlings growing without tree shelters, whereas only 3.6% of the trees were browsed by mice or voles and 2.5% were browsed by rabbits. Fabric mats, used to control competition from herbaceous plants for roughly half of the seedlings included in the study, had a detrimental effect overall. Seedlings grown with mats had a greater frequency of deer browsing and a greater chance of dying than seedlings grown without mats. Stem height for seedlings browsed by deer was less overall than for nonbrowsed seedlings, although this pattern varied with use of fabric mats and plot location. The use of plastic tree shelters effectively prevented deer browsing and reduced the mortality rate from 34.6% to 3.2%. Our results indicate that fabric mats should not be used in restoration projects with large deer populations. They also suggest that planting seedlings away from existing forest edges and using seedling protection measures such as tree shelters will increase seedling survival and growth in future restoration projects.     

Population dynamics and growth patterns for a cohort of northern red oak (Quercus rubra) seedlings

Summary I studied the survival and development of a 1986 cohort of northern red oak (Quercus rubra L.) seedlings growing under a variety of overstory and microsite conditions in a northern hardwood forest dominated by northern red oak, red maple (Acer rubrum L.) paper birch (Betula papyrifera Marsh.), and scattered white pine (Pinus strobus L.). Fifty naturally regenerating seedlings of oak were randomly selected in each of three canopy classes: no overstory, partial overstory, and complete overstory. Growth and mortality were measured for six years. Seedling height growth decreased with overstory density, with less growth evident with even a partial overstory. Seedling survival also declined with overstory density and depended on microtopography to a lesser extent. After six years, 92% of the seedlings survived in the open, compared to 54% under the partial overstory, and 36% under the complete overstory. The open environment, in which woody and herbaceous regrowth formed a low canopy reducing light intensities to about 50% of full sunlight, provided a favorable site for the growth and survival of northern red oak.     

TEMPORAL AND SPATIAL PATTERNS OF SPECIFIC LEAF WEIGHT IN SUCCESSIONAL NORTHERN HARDWOOD TREE SPECIES

Temporal and spatial patterns of specific leaf weight (SLW, g/m²) were determined for deciduous hardwood tree species in natural habitats in northern lower Michigan to evaluate the utility of SLW as an index of leaf photosynthetic capacity. No significant diurnal changes in SLW were found. Specific leaf weight decreased and then increased during leaf expansion in the spring. Most species, especially those located in the understory, then had relatively constant SLW for most of the growing season, followed by a decline in SLW during autumn. Specific leaf weight decreased exponentially down through the canopy with increasing cumulative leaf area index. Red oak (Quercus rubra), paper birch (Betula papyrifera), bigtooth aspen (Populus grandidentata), red maple (Acer rubrum), sugar maple (A. saccharum), and beech (Fagus grandifolia) generally had successively lower SLW, for leaves at any one level in the canopy. On a given site, comparisons between years and comparisons of leaves growing within 35 cm of each other showed that differences in SLW among species were not due solely to microenvironmental effects on SLW. Bigtooth aspen, red oak, and red maple on lower-fertility sites had lower SLW than the same species on higher-fertility sites. Maximum CO₂ exchange rate, measured at light-saturation in ambient CO₂ and leaf temperatures of 20 to 25 C, increased with SLW. Photosynthetic capacities of species ranked by SLW in a shaded habitat suggest that red oak, red maple, sugar maple, and beech are successively better adapted to shady conditions.     

Riparian Forest Restoration: Increasing Success by Reducing Plant Competition and Herbivory

The reestablishment of riparian forest is often viewed as “best management practice” for restoring stream ecosystems to a quasi‐natural state and preventing non‐point source contaminants from entering them. We experimentally assessed seedling survivorship and growth of Quercus palustris (pin oak), Q. rubra (red oak), Q. alba (white oak), Betula nigra (river birch), and Acer rubrum (red maple) in response to root‐stock type (bare root vs. containerized), herbivore protection (tree shelters), and weed control (herbicide, mowing, tree mats) over a 4‐year period at two riparian sites near the Chester River in Maryland, U.S.A. We started with tree‐stocking densities of 988/ha (400/ac) in the experimental plots and considered 50% survivorship (i.e., a density of 494/ha [200/ac] at crown closure) to be an “acceptable or minimum” target for riparian restoration. Results after four growing seasons show no significant difference in survivorship and growth between bare‐root and containerized seedlings when averaged across all species and treatments. Overall survivorship and growth was significantly higher for sheltered versus unsheltered seedlings (49% and 77.6 cm vs. 12.1% and 3.6 cm, respectively) when averaged across all species and weed control treatments. Each of the five test species exhibited significantly higher 4‐year growth with shelter protection when averaged across all other treatments, and all species but river birch had significantly higher survivorship in shelters during the period. Seedlings protected from weeds by herbicide exhibited significantly higher survivorship and growth than seedlings in all other weed‐control treatments when averaged across all species and shelter treatments. The highest 4‐year levels of survivorship/growth, when averaged across all species, was associated with seedlings protected by shelters and herbicide (88.8%/125.7cm) and by shelters and weed mats (57.5%/73.5 cm). Thus, only plots where seedlings were assisted by a combination of tree shelters and either herbicide or tree mats exhibited an “acceptable or minimum” rate of survivorship (i.e.,>50%) for riparian forest restoration in the region. Moreover, the combined growth and survivorship data suggest that crown closure over most small streams in need of restoration in the region can be achieved most rapidly (i.e., 15 years or less) by protecting seedlings with tree shelters and controlling competing vegetation with herbicides.     

Inadequate Cold Tolerance as a Possible Limitation to American Chestnut Restoration in the Northeastern United States

The American chestnut (Castanea dentata (Marshall) Borkh.), once a major component of eastern forests from Maine to Georgia, was functionally removed from the forest ecosystem by chestnut blight (an exotic fungal disease caused by Cryphonectria parasitica (Murr.) Barr), first identified at the beginning of the twentieth century. Hybrid‐backcross breeding programs that incorporate the blight resistance of Chinese chestnut (Castenea mollissima Blume) and Japanese chestnut (Castenea crenata Sieb. & Zuc.) into American chestnut stock show promise for achieving the blight resistance needed for species restoration. However, it is uncertain if limitations in tissue cold tolerance within current breeding programs might restrict the restoration of the species at the northern limits of American chestnut's historic range. Shoots of American chestnut and hybrid‐backcross chestnut (i.e., backcross chestnut) saplings growing in two plantings in Vermont were tested during November 2006, February 2007, and April 2007 to assess their cold tolerance relative to ambient low temperatures. Shoots of two potential native competitors, northern red oak (Quercus rubra L.) and sugar maple (Acer saccharum L.), were also sampled for comparison. During the winter, American and backcross chestnuts were approximately 5°C less cold tolerant than red oak and sugar maple, with a tendency for American chestnut to be more cold tolerant than the backcross chestnut. Terminal shoots of American and backcross chestnut also showed significantly more freezing damage in the field than nearby red oak and sugar maple shoots, which showed no visible injury.     

The Use of Asian Ficus Species for Restoring Tropical Forest Ecosystems

Fig (Ficus spp.) trees have been promoted as framework species for tropical forest restoration throughout Asia, because they are considered to be keystone species. This article presents optimal propagation and planting techniques for six Asian dioecious Ficus species, which will enable their inclusion in forest restoration plantings across the Asia‐Pacific region: Ficus auriculata, F. fulva, F. hispida, F. oligodon, F. semicordata, and F. variegata. Nursery experiments compared the growth performance of propagating planting stock from seed and from leafy cuttings, whereas field experiments assessed the cost‐effectiveness and the relative performance of (1) direct seeding, (2) planting stock from seed, and (3) planting stock from cuttings. The most efficient method of producing Ficus spp. was from seed. Propagation from cuttings was much less successful. Seedlings produced from seed had the highest rates of growth and survival both in the nursery and in field trials. In field trials, use of planting stock from seed was also more cost‐effective than direct seeding and vegetative propagation. Establishment costs calculated on the basis of “per plant established” were $1.14 for seed, $6.95 for cutting, and $25.88 for direct seeding.     

Effect of a strong cold storage induced desiccation on metabolic solutes,stock quality and regrowth,in seedlings of two oak species

Bare-root seedlings of pedunculate oak (Quercus robur L.) and northern red oak (Quercus rubra L.) were lifted in January and stored at 1.8°C, at 82% relative humidity, until their fresh weight declined by 33%. Root growth potential (RGP), fine root electrolyte leakage (REL), fine root water content (RWC), shoot tip water content (SWC), starch and metabolic solute contents in root and shoot, were measured just after lifting and after treatment. Survival of treated seedlings was also assessed in a field trial. RWC, SWC, REL, RGP were dramatically affected by desiccation during cold storage. In both species, root soluble carbohydrate level, inositol level and isocitrate level increased, whereas root starch level and shoot soluble carbohydrate level decreased. In northern red oak, treated seedlings had higher root contents of soluble carbohydrates, inositol and proline than in pedunculate oak. Moreover, treatment induced proline accumulation only in northern red oak roots. These differences could explain why field survival of treated seedlings was significantly better in northern red oak than in pedunculate oak.     

Radial growth and climate responses of white oak (Quercus alba) and northern red oak (Quercus rubra) at the northern distribution limit of white oak in Quebec, Canada

Aim The objectives of this study were: (1) to compare radial growth patterns between white oak (Quercus alba L.) and northern red oak (Quercus rubra L.) growing at the northern distribution limit of white oak; and (2) to assess if the radial growth of white oak at its northern distribution limit is controlled by cold temperature. Location The study was conducted in three regions of the Ottawa valley in southern Québec. All stands selected were located at the northern limit of distribution of Q. alba. Methods Twelve mixed red and white oak stands were sampled and increment cores were extracted for radial growth analyses. For each oak species, 12 chronologies were derived from tree‐ring measurement (residual chronologies). Principal components analysis and redundancy analysis were used to highlight the difference between radial growth in both species and to determine their radial growth–climate association. Results There was little difference between the radial growth of each species; Q. alba, however, exhibits more year‐to‐year variation in growth than Q. rubra. More than 65% of the variance in radial growth was shared among sites and species. Both species showed a similar response to climate, which suggested that the limit of distribution of Q. alba might not be determined by effects on growth. Both species had a classic response to climate and drought in the early growing season. Main conclusions The northern distribution limit of Q. alba does not appear to be directly controlled by effects on growth processes as indicated by the similarities in radial growth and response to climate between the two species. The location of the stands on southern aspects suggested that cold temperature could have been a major factor controlling the distribution limit of Q. alba. However, it is speculated that stands growing on southern aspects may be more prone to forest fires or to drought, which would favour the maintenance and establishment of oaks, and of Q. alba in particular. Models relating the northern distribution limits of species to broad climate parameters like annual mean temperature will need to be reviewed to incorporate more biologically relevant information. Such assessments will in turn provide better estimates of the effect of climate changes on species distribution.     

Effects of 76 Hz electromagnetic fields on forest ecosystems in northern Michigan: Tree growth

Since 1984, the possible effects of extremely low-frequency electromagnetic (EM) fields generated by a 76 Hz communication antenna on the growth and productivity of four deciduous and one coniferous species have been studied in the Upper Peninsula of Michigan. Results from two research sites are discussed here: one site near an antenna element and a control site located 50 km from the communication system. Growth models for individual tree diameters were developed for northern red oak (Quercus rubra), paper birch (Betula papyrifera), aspen (Populus tremuloides with a few individuals ofP. grandidentata), and red maple (Acer rubrum). A growth model for individual tree height was developed for young red pine (Pinus resinosa). Average differences between the observed and predicted growth were calculated for each growing season and then compared between the study sites and across the study periods to evaluate changes in growth patterns which could be attributed to EM field effects. For aspen and red maple, the results showed a stimulation of diameter growth at magnetic flux density levels of 1 to 7 milliGauss; height growth of red pine was increased at about the same exposure levels. There are no clear indications of an EM field effect on total annual diameter growth for either of the other two species.     

Species‐Specific Barriers to Tree Regeneration in High Elevation Habitats of West Virginia

Herbaceous competition and herbivory have been identified as critical barriers to restoration of native tree species in degraded landscapes around the world; however, the combined effects of competition and herbivory are poorly understood. We experimentally manipulated levels of herbivory and herbaceous competition and analyzed the response of tree seedling performance over three growing seasons as a function of species and habitat in north‐central West Virginia. Four native tree species were planted in old field and forest experimental plots: Castanea dentata (American chestnut), Quercus rubra (red oak), Acer saccharum (sugar maple), and Picea rubens (red spruce). Red spruce demonstrated the highest growth increment and greatest survival (64%) and most consistent results among treatments and habitats. Red spruce survival was not reduced in the presence of Odocoileus virginianus (white‐tailed deer) browse and herbaceous competition; however, growth was improved by suppression of herbaceous competition. We suspect that this deciduous forest landscape would regenerate to a red spruce dominated forest if seed source was available. In contrast, the other three species tested had very low survival when exposed to deer and were more responsive to competing vegetation and habitat type. American chestnut had low survival and growth across all treatments, suggesting basic climate limitations. Vigorous natural regeneration of Prunus serotina (black cherry) occurred in forest plots where both competing herbs and deer were excluded. Our results demonstrated the importance of testing multiple potential recruitment barriers and species at once and the need for species and habitat‐specific restoration treatments.     

Facilitation of tree seedling establishment in a sand dune succession

Abstract. A nucleated pattern of establishment by pine seedlings of Pinus strobus and P. resinosa around pre-established oak trees of Quercus rubra in a sand dune succession in Ontario, Canada was examined using field observations, seed planting and habitat manipulation. Densities of young pines beneath oak canopies were approximately six times greater than in treeless areas, and densities on the north sides of the trees were significantly greater than on the south sides. However, oaks younger than 35 yr showed no preferential establishment beneath them, while the pine population structures beneath older oaks indicated single periods of successful recruitment. Pine seed planted beneath and beyond oak canopies of three sizes germinated primarily beneath the canopies of medium and large-sized oaks, but subsequent survivorship of seedlings over two growing seasons was poor. Several micro-environmental conditions were changed by oak canopies, but only shade showed a pattern closely corresponding to that of seedling establishment. A habitat manipulation experiment confirmed the primary role of shade in facilitating pine seedling establishment beneath oaks. Failure of pine to continue recruiting successfully beneath facilitating trees is tentatively attributed to intraspecific competition among pine individuals.     

Environmentally-based maternal effects: a hidden force in insect population dynamics?

Summary The nutritional environment of the parental generation of the polyphagous gypsy moth, Lymantria dispar, can significantly influence the growth and reproductive potential of the next generation through environmentally-based maternal effects. In the first experiment, members of the parental generation were reared on red oak trees (Quercus rubra L.) with known defoliation and phenolic levels. Diet in the offspring generation was homogeneous (synthetic diet). With genetic effects accounted for 1) offspring attained greater pupal weights when their mothers fed on trees with higher leaf damage levels, 2) daughters had a shorter prefeeding stage, a trait associated with dispersal tendency, when their mothers experienced higher condensed tannin levels, and 3) sons had lower pupal weights when their mothers experienced greater condensed tannin levels. In the second experiment, members of the parental generation were reared on either red or black oak (Q. velutina) trees. Offspring of each mother were divided among four diets: red oak, chestnut oak (Q. prinus L.), a standard synthetic diet, and a low-protein synthetic diet. The parental host species accounted for 24% of the variation in daughters' development time; offspring diet accounted for 52%. When mothers were reared on black oak rather than red oak, their offspring developed significantly faster when the F1 diet was chestnut oak. Environmentally-based maternal effects can significantly influence the expression of offspring dispersal potential, growth rate, and offspring fecundity. These traits contribute to natality and survival in natural populations and, hence, to population growth potential. Theoretical models of insect population dynamics demonstrate that the presence of a time delay in a density dependent response can induce destabilization. Maternal effects act on a time delay and may participate in the destabilization characteristic of outbreak species.     

Branch growth and leaf numbers of red maple (Acer rubrum L.) and red oak (Quercus rubra L.): response to defoliation

Summary Branch growth and leaf formation from terminal and from lateral buds of red maple (Acer rubrum L.) and red oak (Quercus rubra L.) were measured in response to simulated insect defoliation. A single large branch representative of the crown of each tree was used for enumeration of growth and of bud numbers throughout three successive years of 0, 50, 75, and 100% leaf removal for the entire tree. Leaf number per tree for both species after the last year of defoliation was reduced in direct proportion to the severity of defoliation, in comparison to the predefoliation status of the trees. Bud number per tree for red maple, but not for red oak, was also reduced in proportion to severity of defoliation.Averaged over all defoliation treatments, defoliation reduced branch growth more than leaf production. Furthermore, the reduction in branch growth and leaf production was greater in red oak than in red maple. Three years of successive defoliation reduced the mean lateral plus terminal branch growth by 40% in red oak and by 23% in red maple, while leaf number was reduced 22% in red oak and remained unchanged in red maple. In red maple, 100% defoliation caused greater branch death than the 50 or 75% defoliation treatments, and the amount of death was greater after each successive year of defoliation. In contrast to red maple, undefoliated red oak incurred a substantial amount of branch death throughout the study which was little affected by defoliation treatment.     

Growth and maintenance respiration in leaves of northern red oak seedlings and mature trees after 3 years of ozone exposure

A two-component model of growth and maintenance respiration is used to study the response of northern red oak (Quercus rubra L.) seedlings and 32-year-old trees to sub-ambient (10 μmol h; cumulative dose based on 7 h daily mean), ambient (43 μmol h), and twice-ambient (85 μmolh) ozone. The relative growth rates (RGR) of leaves sampled from seedlings and trees were similar across treatments, as were specific leaf respiration rates (SRR). Growth coefficients estimated from the SRR versus RGR relationship averaged 25-3 mol CO2 kg^?1 leaf dry mass produced for seedlings and 21-5 mol kg^?1 for trees. Maintenance coefficients ranged from 0-89 to 1-07 mol CO₂ kg^?1 leaf dry mass d^?1 for seedlings and from 0-64 to 0-84 mol kg-1 d^?1 for trees. Neither coefficient was affected by ozone. Leaves sampled throughout the growing season also showed little response of respiration to ozone. This occurred despite a 30% reduction in net photosynthesis for trees grown at twice-ambient ozone. These results suggest that growth and maintenance respiration in young northern red oak leaves are not affected by ozone and that in older leaves injury can occur without a parallel increase in so-called ‘maintenance’ respiration.     

Variation in Soil Temperature,Moisture, and Plant Growth with the Addition of Downed Woody Material on Lakeshore Restoration Sites

Downed woody material (DWM) is an important ecosystem component that performs many critical functions including influencing soil temperature and moisture, which affects plant growth and survival. Residential development along lakeshores has increased dramatically in recent decades in the northern Great Lakes region. Such development often leads to reductions in terrestrial and aquatic woody material. Although lakeshore restoration projects have occurred in the past few years in the region, there has been little effort to evaluate success. In 2007, a collaborative lakeshore restoration research project began on two lakes in Vilas County, Wisconsin. We investigated the benefits of the addition of DWM as part of these restoration projects. We randomly assigned three coverage treatments (0, 25, and 50%) of DWM on 3 × 3–m experimental plots (n = 10 per treatment) and monitored soil temperature and volumetric soil water content at a depth of 10 cm. All plots were planted with two native shrub species and five native understory herbaceous species. Mean maximum soil temperature, mean difference in daily high and low soil temperature, and percent change in soil moisture content were significantly lower in the 25 and 50% DWM plots. Plant canopy volume growth for snowberry (Symphoricarpos albus) and Barren strawberry (Waldstenia fragaroides) was significantly greater in the 25 and 50% DWM plots. We conclude that the addition of DWM had a significant positive effect on regulating soil temperature extremes, soil moisture, and plant volume growth for two species of native plants used for restoration projects.     

Northward migrating trees establish in treefall gaps at the northern limit of the temperate–boreal ecotone, Ontario, Canada

Climate change is expected to promote migration of species. In ecotones, areas of ecological tension, disturbances may provide opportunities for some migrating species to establish in otherwise competitive environments. The size of and time since disturbance may determine the establishment ability of these species. We investigated gap dynamics of an old-growth red pine (Pinus resinosa Sol. ex Aiton) forest in the Great Lakes–St. Lawrence forest in northern Ontario, Canada, a transition zone between temperate and boreal forest. We investigated the effects of gaps of different sizes and ages on tree species abundance and basal area. Our results show that tree species from the temperate forest further south, such as red maple (Acer rubrum L.), red oak (Quercus rubra L.), and white pine (Pinus strobus L.), establish more often in large, old gaps; however, tree species that have more northern distributions, such as black spruce (Picea mariana Mill.), paper birch (Betula papyrifera Marsh.), and red pine show no difference in establishment ability with gap size or age. These differences in composition could not be attributed to autogenic succession. We conclude that treefall gaps in this forest facilitate the establishment of northward migrating species, potentially providing a pathway for future forest migration in response to recent changes in climate.     

Development of Tree Regeneration in Fern-dominated Forest Understories After Reduction of Deer Browsing

Dennstaedtia punctilobula (hay‐scented fern) can act as a native invasive species in forests in eastern North America where prolonged deer browsing occurs in stands with partially open overstory canopies. Ferns dominate the understory with a 60‐cm tall canopy, with little regeneration of native tree species. It has been hypothesized that, once established, ferns may continue to inhibit tree regeneration after deer browsing has been reduced. To test this hypothesis, we documented the pattern of recovery of the tree seedling understory in plantations of Pinus strobus (white pine) and Pinus resinosa (red pine) on the Quabbin Reservation watershed protection forest in central Massachusetts, where after 40 years of intensive deer browsing the deer herd was rapidly reduced through controlled hunting. Dense fern understories occur on nearly 4,000 ha of the predominantly oak–pine forest. Three years after deer herd reduction, stands with the highest density fern cover (77% of plots with>90% cover) had significantly fewer seedlings at least 30 cm in height, compared with stands with lower fern density, and those seedlings consisted almost entirely of Betula lenta (black birch) and white pine. Height growth analysis showed that black birch and white pine grew above the height of the fern canopy in 3 and 6 years, respectively. In contrast, two common species, Fraxinus americana (white ash) and Quercus rubra (red oak), grew beneath the dense fern cover for 5 years with height growth less than 5 cm/yr after the first year. A study of spring phenology indicated that the ability of black birch to grow through the fern canopy might have been due to its early leaf development in spring before the fern canopy was formed, in contrast to oak and ash with delayed leaf development. Thus, the ferns showed differential interference among species with seedling development after reduction of deer browse.     

CO2-induced growth enhancements of co-occurring tree species decline at different rates

To elucidate how enriched CO₂ atmospheres, soil fertility, and light availability interact to influence the long-term growth of tree seedlings, six co-occurring members of temperate forest communities including ash (Fraxinus americana L.), gray birch (Betula populifolia), red maple (Acer rubrum), yellow birch (Betula alleghaniensis), striped maple (Acer pensylvanicum), and red oak (Quercus rubra L.) were raised in a glasshouse for three years in a complete factorial design. After three years of growth, plants growing in elevated CO₂ atmospheres were generally larger than those in ambient CO₂ atmospheres, however, magnitudes of CO₂-induced growth enhancements were contingent on the availability of nitrogen and light, as well as species identity. For all species, magnitudes of CO₂-induced growth enhancements after one year of growth were greater than after three years of growth, though species' growth enhancements over the three years declined at different rates. These results suggest that CO₂-induced enhancements in forest productivity may not be sustained for long periods of time. Additionally, species' differential growth responses to elevated CO₂ may indirectly influence forest productivity via long-term species compositional changes in forests.     

Factors that affect compartmentalization and wound closure of Quercus rubra infested with Enaphalodes rufulus

• 1 The mechanisms by which hardwood trees resist invasion by native wood borers are still poorly understood.
• 2 We examined the importance of several host, herbivore and environmental variables in relation to Quercus rubra L., northern red oak, resistance to a native cerambycid, Enaphalodes rufulus (Haldeman) (Coleoptera: Cerambycidae), the red oak borer.
• 3 We employed tree‐ring techniques to date and measure two indicators of host tree physical resistance, vertical lesion size and wound closure of scars left by larval galleries, within 107 Q. rubra hosts from eight sites within the Ozark and Ouachita National Forests of Arkansas.
• 4 Host Q. rubra decline status proved a valuable predictor of resistance, wherein healthy trees exhibited better compartmentalization capacity, as indicated by shorter lesions and faster wound closure, than declining trees and those that died during a recent E. rufulus outbreak.
• 5 Only healthy Q. rubra exhibited a significant declining lesion length trend from 1988 to 2006 and were likely able to re‐allocate resources in favour of maintaining resistance as the feeding pressure from borers increased and then relaxed post‐outbreak.
• 6 Variables related to borer feeding pressure consistently explained more variation in lesion length than host or environmental variables. Hosts were better able to compartmentalize injury from small, unsuccessful E. rufulus galleries than large, successful galleries from which larvae survived to adulthood.
• 7 Healthy Q. rubra, as survivors of the recent outbreak, were able to effectively tolerate feeding pressure from borers at the same time as sustaining growth and maintenance. Tolerance may be an important resistance mechanism for hardwood hosts against the native wood borers that attack them.