Responses of three bottomland species with different flood tolerance capabilities to various flooding regimes

Seedlings of baldcypress (Taxodium distichum), nuttall oak (Quercus nuttalli), and cherrybark oak (Quercus falcata var.pagodaefolia) were subjected to four flooding treatments: control, continuously flooded, intermittently flooded, and partially flooded for 70 days in a greenhouse. The treatments imposed various durations and intensities of soil redox potential (Eh) conditions representing a range encountered by plants in their habitats. Morphological changes and gas exchange responses to the treatments differed among the study species. Rapid development of adventitious root and hypertrophied lenticels were observed in baldcypress and nuttall oak under all flooded treatments. Cherrybark oak had the highest percentage reduction in net photosynthesis ranging from 65–87%, whereas reductions in nuttall oak ranged between 35–68% and in baldcypress between 6–21% in response to various treatments. Recovery of gas exchange was noted in baldcypress but no significant recovery was found in oaks. The recovery in baldcypress contributed to the continued growth and biomass accumulation under various treatments. Little evidence of consistent changes in biomass allocation patterns in response to the treatments was found in baldcypress but total biomass decreased significantly under the continuously flooded treatment. In oaks, total biomass decreased significantly in all flooded treatments. The present findings demonstrated that physiological functions are adversely affected by low soil Eh conditions and the extent of such effects are dependent on the intensity and duration of soil reduction as well as the species' capability to respond to such conditions rapidly. Management plans concerned with regeneration of bottomland forested ecosystems should consider the species flood response capabilities at seedling stages as well as the timing, durations, and intensities of soil reduction at the specific site.     

The Effects of Intermittent Flooding on Seedlings of Three Forest Species

Under greenhouse conditions, seedlings of three forest species, baldcypress (Taxodium distichum), nuttall oak (Quercus nuttallii), and swamp chestnut oak (Quercus michauxii) were subjected to an intermittent flooding and subsequent physiological and growth responses to such conditions were evaluated. Baldcypress showed no significant reductions in stomatal conductance (g _s) or net photosynthetic rate (P _N) in response to flood pulses. In nuttall oak seedlings g _s and P _N were significantly decreased during periods of inundation, but recovered rapidly following drainage. In contrast, in swamp chestnut oak g _s was reduced by 71.8 % while P _N was reduced by 57.2 % compared to controls. Baldcypress displayed no significant changes in total mass while oak species had significantly lower leaf and total mass compared to their respective controls. Thus baldcypress and nuttall oak showed superior performance under frequent intermittent flooding regimes due to several factors including the ability for rapid recovery of gas exchange soon after soil was drained. In contrast, swamp chestnut oak seedlings failed to resume gas exchange functions after the removal of flooding.     

Effects of Flooding on Susceptibility of Taxodium distichum L. Seedlings to Drought

Responses of baldcypress (Taxodium distichum) seedlings to soil moisture were studied to test the hypothesis that flooding may lead to seedling's higher susceptibility to drought. Treatments included a well-watered but drained control (C), continuously flooded (CF), control followed by drought (CD), and flooded followed by drought (FD). Gas exchange values revealed no significant effects on net photosynthetic rate (P _N) in response to flooding. In contrast, after the onset of drought, P _N was significantly reduced in CD and FD plants. Significant growth reductions under mild drought conditions indicated that baldcypress seedlings were drought sensitive. However, comparison of gas exchange rates and growth responses between CD and FD plants indicated that prior flooding had no detectable effect on subsequent sensitivity of baldcypress to drought. These findings explain baldcypress persistence in wetland habitats characterized by periodic flooding and mild drought.     

Impact of waterlogging on the N-metabolism of flood tolerant and non-tolerant tree species

The present study was conducted to characterize the N‐metabolism of important European tree species with different degrees of flooding tolerance. The roots of Fagus sylvatica (sensitive to flooding), Quercus robur (moderately flood tolerant) and Populus tremula × P. alba (flood tolerant) saplings were exposed to different flooding regimes and N uptake, amino acid, protein and chlorophyll concentrations as well as gas exchange were measured. The effects of these treatments on the tree species varied distinctly. In general, the N metabolism of beech was severely affected whereas less impacts were observed on oaks and almost no effects on poplars. The concentrations of amino compounds, particularly of Asp, Asn, Glu and Gln, were lower in the roots of flooded trees than in controls. By contrast, γ‐amino butyric acid concentrations increased. Root protein concentrations remained unaffected in oak and poplar but decreased in beech in response to flooding. The concentrations of pigments remained unaffected by flooding in all tree species investigated. However, photosynthesis and transpiration were severely affected in beech but much less in oak and poplar. The data obtained show a clear correlation between the different flooding tolerances of the trees investigated and the impacts of flooding on N uptake and N metabolism.     

Interactive effects of substrate,hydroperiod, and nutrients on seedling growth of Salix nigra and Taxodium distichum

The large river swamps of Louisiana have complex topography and hydrology, characterized by black willow (Salix nigra) dominance on accreting alluvial sediments and vast areas of baldcypress (Taxodium distichum) deepwater swamps with highly organic substrates. Seedling survival of these two wetland tree species is influenced by their growth rate in relation to the height and duration of annual flooding in riverine environments. This study examines the interactive effects of substrate, hydroperiod, and nutrients on growth rates of black willow and baldcypress seedlings. In a greenhouse experiment with a split-split-plot design, 1-year seedlings of black willow and baldcypress were subjected to two nutrient treatments (unfertilized versus fertilized), two hydroperiods (continuously flooded versus twice daily flooding/draining), and two substrates (sand versus commercial peat mix). Response variables included height, diameter, lateral branch count, biomass, and root:stem ratio. Black willow growth in height and diameter, as well as all biomass components, were significantly greater in peat substrate than in sand. Black willow showed a significant hydroperiod–nutrient interaction wherein fertilizer increased stem and root biomass under drained conditions, but flooded plants did not respond to fertilization. Baldcypress diameter and root biomass were higher in peat than in sand, and the same two variables increased with fertilization in flooded as well as drained treatments. These results can be used in Louisiana wetland forest models as inputs of seedling growth and survival, regeneration potential, and biomass accumulation rates of black willow and baldcypress.     

Gas composition and respiration of water oak (Quercus nigra L.) and green ash (Fraxinus pennsylvanica Marsh.) roots after prolonged flooding

Summary We compared the effects of 9.5 months of a continuous flooding treatment with a drained control treatment on one year-old seedlings of green ash (Fraxinus pennsylvanica Marsh.) and water oak (Quercus nigra L.), two tree species common to the bottomland-hardwood forests of eastern North America. The internal root gas composition of the more flood tolerant species, green ash, maintained higher oxygen and lower carbon dioxide concentrations under the flooding treatment than water oak. This apparently resulted in differences in rhizosphere oxidation. The amounts of Fe and Mn and the Fe/Mn ratio of the root coating extracted from trees in reduced soil conditions were much higher for the green ash than the water oak. It is argued that this reflects differences in the ability of these two species to maintain rhizosphere oxidation under prolonged periods of flooding and to prevent the accumulation of reduced potentially phytotoxic compounds. Alcohol dehydrogenase activity increased in the green ash and decreased in the water oak in the flooded treatment. This indicated that the better adapted species was able to rely upon increased anaerobic respiration in order to compensate for the decreased root oxygen supply, but the water oak was unable to maintain previous levels of respiration, probably as the result of sulfide toxicity.     

Weed control and overstory reduction improve survival and growth of under‐planted oak and hickory seedlings

Weed control and overstory reduction are important silvicultural treatments for improving survival and growth of under‐planted oak and hickory seedlings. Mast‐producing trees in the bottomland forests of the blackland prairie and Post Oak Savannah ecoregions of Texas have declined in abundance. Oaks and hickories have been replaced by more shade‐tolerant species, including green ash (Fraxinus pennsylvanica Marshall) and sugarberry (Celtis laevigata Willd.), which do not produce significant hard mast for priority wildlife species. A split‐plot experiment design was installed on three sites at Richland Creek Wildlife Management Area in Freestone County, Texas, studying the effects of canopy coverage and competition control on survival and growth of bur oak (Quercus macrocarpa Michx.), Shumard oak (Quercus shumardii Buckl.), and pecan (Carya illinoinensis (Wagenh.) K. Koch) seedlings. Uprooting by hogs shortly after planting resulted in greater than 90% mortality of pecan on the two lower elevation sites. Year one survival of Shumard oak was significantly higher than bur oak. However, bur oak was more preferred by hogs than Shumard oak. Year one growth of bur oak was significantly greater than Shumard oak. Severe flooding during the second growing season caused complete mortality on the lower two sites. None of the species were well suited to such prolonged (3–4 months) inundation as seedlings. On the remaining site, density reduction and weed‐barrier mats improved growth and survival while herbaceous weed control with herbicides actually reduced both growth and survival.     

EFFECTS OF FLOODING AND NUTRIENT ENRICHMENT ON BIOMASS ALLOCATION IN ACER RUBRUM SEEDLINGS

A greenhouse experiment was conducted on Acer rubrum seedlings to evaluate the effects of flood frequency on production and allocation of biomass and to test the effects of N and P fertilization on production and allocation. Seedlings from the Dismal Swamp were subjected to three flood treatments (no flooding, intermittent flooding, and continuous flooding) and four enrichment treatments (no enrichment, N additions, P additions, and N + P additions). More continuous flooding resulted in less biomass production. Biomass increased during the study in all treatments except for root mass in the continuously flooded treatment. However, production of abundant adventitious roots compensated for the lack of normal root growth. Root/shoot ratios exhibited the greatest decreases in the continuously flooded plants. Plants with N + P added had significantly more leaf, stem, and total mass than the nonenriched plants four months into the study. The N + P additions had apparently compensated for the effects of flood stress in the continuously flooded plants by the end of the study. The fertilized seedlings accumulated higher concentrations of N and P, but their nutrient use efficiency (biomass production per unit nutrient absorbed) was lower than in the nonenriched plants. Acer rubrum seedlings survive flooded conditions through several adaptations; however, theirgrowth is slowed by continuous flooding.     

Comparative studies on physiological and biochemical adaptation of Taxodium distichum and Taxodium ascendens seedlings to different soil water regimes

Responses of baldcypress (Taxodium distichum) and pondcypress (Taxodium ascendens) seedlings in leaf gas exchange and root metabolism to a wide range of hydrological regimes were studied. Four water treatments included control (C), mild drought (MD), wet soil (WS) and flooding (FL). Both species showed no significant change in net photosynthetic rate (Pn) in response to flooding in contrast to significant reduction in Pn under mild drought conditions. In baldcypress, Pn was also maintained at control level compared to significant increase of Pn in pondcypress when subjected to the wet soil treatment. In flooding, both species showed significant increase in contents of malate and shikimate in their lateral roots and further in total roots, as compared to control. However, different adaptation to flooding was demonstrated in their taproots in contents of malate and shikimate. Furthermore, baldcypress seedlings showed significant reductions in lateral and total root biomass, in contrast with no significant change in that of pondcypress seedlings in response to flooding. In wet soil, no significant effects were revealed in contents of malate and shikimate in roots (including different root portions) of baldcypress seedlings in contrast to significant increase of their contents in lateral roots of pondcypress seedlings. No significant effects on root biomass were detected in either baldcypress or pondcypress seedlings in response to wet soil. When subjected to mild drought, baldcypress seedlings displayed no significant change in contents of malate and shikimate in roots, whereas pondcypress seedlings exhibited significant reduction in content of shikimate in lateral roots and total roots. In addition, root biomass of baldcypress seedlings showed no significant change, while lateral and total root biomass of pondcypress seedlings demonstrated significant increase when mild drought was imposed. These findings indicate that baldcypress and pondcypress appear to be promising candidates for reforestation in the Three Gorges Reservoir region due to their characteristics of water-tolerance and mild drought endurance. However, pondcypress is more resilient than baldcypress to the continuum of water availability conditions expected in the Three Gorges Reservoir region.     

Flooding and salinity effects on growth and survival of four common forested wetland species

The survival, growth, and biomass of baldcypress (Taxodium distichum (L.) Rich.), water tupelo (Nyssa aquatica L.), Chinese tallow (Sapium sebiferum (L.) Roxb.), and green ash (Fraxinus pennsylvanica Marsh.) seedlings were examined in an experiment varying water levels (watered, flooded) and salinity levels (0, 2, and 10 ppt, plus a simulated storm surge with 32 ppt saltwater). All seedlings, except for those flooded with 10 ppt saltwater, survived to the end of the experiment. In 10 ppt saltwater, flooded baldcypress, water tupelo, and green ash survived two weeks whereas Chinese tallow survived for 6 weeks. However, a second set of slightly older baldcypress, water tupelo, and Chinese tallow seedlings survived eight weeks of flooding with 10 ppt saltwater. When carried through the winter to the beginning of the second growing season, flooded baldcypress and Chinese tallow seedlings from the 0 and 2 ppt treatments leafed out, but only Chinese tallow recovered from the saltwater surge treatment. The diameter and growth (height) of each species was not affected when watered with 2 ppt saltwater, except for the effects of the height growth of baldcypress. Growth was reduced for all species when watered with 10 ppt saltwater. The diameter growth of green ash was reduced by freshwater flooding. The diameter growth of baldcypress and water tupelo was greater when flooded with fresh water. Flooding with 2 ppt saltwater caused a significant reduction in diameter growth in water tupelo, green ash, and Chinese tallow, but not in baldcypress. Root and stem biomass values were not significantly different for any species between the 0 and 2 ppt salinity watering treatments. However, seedlings watered with 10 ppt saltwater had significantly lower root and stem biomass values, except for baldcypress roots and green ash stems. Baldcypress was least affected by flooding with 0 and 2 ppt saltwater, although there were slight reductions in root biomass with increasing salinity. Because of the susceptibility of the seedlings of these four species to increases in flooding and salinity, their regeneration may be limited in the future, thereby causing shifts in species composition.     

Photosynthesis and water relations of four oak species: impact of flooding and salinity

 As global climate changes, sea level rise and increased frequency of hurricanes will expose coastal forests to increased flooding and salinity. Quercus species are frequently dominant in these forest, yet little is known about their salinity tolerance, especially in combination with flooding. In this study, 1-year-old seedlings of Quercus lyrata Walt. (overcup oak), Q. michauxii Nutt. (swamp chestnut oak), Q. nigra L. (water oak), and Q. nuttallii Palmer (Nuttall oak) were chronically (simulating sea level rise) and acutely (simulating hurricane storm surge) exposed to increased flooding and salinity, individually and in combination. The four species demonstrated two response patterns of photosynthesis (A), conductance, and leaf water potential, apparently related to their relative flood tolerance. In Q. lyrata, Q. nuttallii, and Q. nigra (moderately flood-tolerant), A was not immediately reduced after the initiation of the freshwater flooding, but was reduced as the duration of flooding increased. In the second pattern, demonstrated by the weakly flood-tolerant Q. michauxii, A was immediately reduced by freshwater flooding with an increasing impact over time. Watering with 2 parts per thousand (ppt) saline water did not consistently reduce A, but flooding with 2 ppt reduced A of all species, similar to the response with freshwater flooding. Photosynthesis of all species was reduced by 6 ppt watering or flooding, with the latter treatment killing all species within 8 weeks. When acutely exposed to 30 ppt salinity, A was quickly and severely reduced regardless of whether the seedlings were watered or flooded. Acutely flooded seedlings exposed to high salinity died within 2 weeks, but seedlings watered with 30 ppt saline water recovered and A was not reduced the following spring. As saline flooding of coastal areas increases due to sea level rise, photosynthesis of these species will be differentially affected based primarily on their flood tolerance. This suggests that increased flooding associated with sea level rise will impact these tree species to a greater extent than small increases in soil salinity. High salinity accompanying storm surges will be very harmful to all of these species. Received: 20 October 1997 / Accepted: 2 December 1998     

Effects of flooding and salinity on photosynthesis and water relations of four Southeastern Coastal Plain forest species

The influence of flooding and salinity on photosynthesis and water relations was examined for four common coastal tree species [green ash (Fraxinus pennsylvanica Marshall), water tupelo (Nyssa aquatica L.). Chinese tallow (Sapium sebiferum (L.) Roxb.), and baldcypress (Taxodium distichum (L.) Richard)]. Both chronic (as might be associated with sea level rise) and acute (similar to hurricane storm surges) exposures to these stresses were examined. Chronic freshwater flooding of green ash, water tupelo, and Chinese tallow seedlings reduced photosynthesis (A) relative to that of watered seedlings, while baldcypress was unaffected. Chinese tallow A declined with increasing length of flooding. A salinity increase of the floodwater to 2 ppt decreased A of baldcypress and water tupelo, but not A of green ash and Chinese tallow, which was already severely reduced by freshwater flooding. All seedlings of the four species died within 2 to 6 weeks when flooded with 10 ppt saltwater. Photosynthesis of all four species did not differ between 0 and 2 ppt watering. Watering with 10 ppt salinity initially reduced A of all four species, but the seedlings recovered over time. Photosynthesis was severely decreased for all species when flooded with 21 ppt salinity for 48 hours. Reduced A continued following the treatment. Photosynthesis of only green ash and water tupelo was reduced by watering with 21 ppt salinity for 48 hours. Flooding of low-lying areas with increased salinity would lead to shifts in species composition of coastal forests due to these differential tolerances.     

Tree shelters facilitate brown oak seedling survival and establishment in a grazing‐dominant forest of Bhutan,Eastern Himalaya

Brown oak (Quercus semecarpifolia) forest is essential for ecological and socioeconomic functions, mainly grazing in the Himalayas. The tree has failed to regenerate naturally and is a threatened species. Restoration of brown oaks is crucial to ensure sustainability while maintaining livestock grazing in these habitats. Achieving this requires cost‐effective restoration techniques that are practicable and sympathetic to the multiple uses of the forest. We assessed the combined effect of grazing (control) and three tree shelters (Protex tubes, mesh wires, and wooden frames) on the field performance of oak seedlings in a forest with heavy grazing pressure. Seedling survival and morphological indicators, including seedling height, collar diameter, sturdiness quotient (SQ), and leaf mass per area (LMA) indices, were measured. More than 90% of control seedlings without protective shelters suffered severe browsing and demonstrated significantly lower survival rates compared to tree shelter seedlings, indicating that grazing was the primary factor governing regeneration success. Seedling survival in tree shelters was three times higher, while the height increase was two times higher than the control. Additionally, locally made mesh wire and wooden tree shelters were more effective than Protex and control in producing quality seedlings reflected by the SQ and LMA values. We suggest that tree shelter is a promising option to restore brown oaks due to its efficacy to defend grazing and support the local community's rights to forest grazing. Our finding is expected to support Bhutan's forest policy of incorporating grazing and tree regeneration into forest management.     

Acorns containing deeper plumule survive better: how white oaks counter embryo excision by rodents

Several squirrel species excise the embryo of acorns of most white oak species to arrest germination for long‐term storage. However, it is not clear how these acorns counter embryo excision and survive in the arms race of coevolution. In this study, we simulated the embryo excision behavior of squirrels by removing 4 mm of cotyledon from the apical end of white oak acorns differing in embryo depths to investigate the effects of embryo excision on acorn germination and seedling performance of white oak species. The embryo depth in the cotyledons was significantly different among white oak acorns, with Quercus mongolica containing the embryo most deeply in the acorns. We found that artificial embryo excision significantly decreased acorn germination rates of Quercus variabilis, Quercus acutissima, Quercus aliena, Quercus aliena var. acutiserrata, Quercus serrata. var. brevipetiolata but not Q. mongolica. Artificial embryo excision exerted significant negative impacts on seedling performance of all oak species except Quercus aliena. Our study demonstrates the role of embryo depth of acorns in countering embryo excision by squirrels and may explain the fact that squirrels do not perform embryo excision in acorns of Q. mongolica with deeper embryos. This apparent adaptation of acorns sheds light on the coevolutionary dynamics between oaks and their seed predators.     

Rosellinia aquila Among Fungi on Branches of Sessile Oak with Symptoms of Decline

Rosellinia aquila was found for the first time in crowns of sessile oaks, Quercus robur (Mattuschka) Liebl., with symptoms of decline. The fungus occurred mostly on diseased branches but also colonized twigs with healthy appearance. Its anamorph and teleomorph stages were produced abundantly on dead branches after long incubation in moist conditions. Colonization of oak trees by R. aquila was likely to have been facilitated by higher humidity and temperature in the oak stands.     

Effects of Flooding Regime and Seedling Treatment on Early Survival and Growth of Nuttall Oak

Effects of flooding on survival and growth of three different types of Nuttall oak (Quercus texana Buckl.) seedlings were observed at the end of third and fifth growing seasons at Yazoo National Wildlife Refuge, Mississippi, U.S.A. Three types of seedlings were planted in January 1995 in a split‐plot design, with four replications at each of two elevations on floodprone, former cropland in Sharkey clay soil. The lower of the two planting elevations was inundated for 21 days during the first growing season, whereas the higher elevation did not flood during the 5‐year period of this study. The three types of 1‐0 seedlings were bareroot seedlings, seedlings grown in containers (3.8 × 21–cm plastic seedling cones), and container‐grown seedlings inoculated with vegetative mycelia of Pisolithus tinctorius (Pers.) Coker. Survival of all the three seedling types was greatest at the lower, intermittently flooded elevation, indicating that drought and related effects on plant competition were more limiting to seedling survival than flooding. At the lower elevation, survival of mycorrhizal‐inoculated container seedlings was greater than that of noninoculated container seedlings. Survival among bareroot seedlings and inoculated container seedlings was not significantly different at either elevation. At the higher, nonflooded elevation, however, bareroot seedling survival was greater than the survival of container seedlings without inoculation. Differences were significant among the inoculated and the noninoculated container seedlings, with higher survival of inoculated seedlings at both elevations, though differences were only significant in year 3. At the end of the fifth year, height of bareroot seedlings was significantly greater than the heights of both types of container‐grown seedlings at both planting elevations. Because seedlings grown in the plastic seedlings cones did not survive better than the bareroot seedlings at either planting elevation, the bareroot stock appear to be the economically superior choice for regeneration in Sharkey soil.     

Leaf gas exchange,chlorophyll fluorescence and growth responses of <Emphasis Type="Italic">Melaleuca alternifolia</Emphasis> seedlings to flooding and subsequent recovery

Periodic flooding of trees in tropical floodplains and reservoirs where water levels fluctuate is a common phenomenon. The effects of flooding and subsequent recovery on gas exchange, chlorophyll fluorescence and growth responses of Melaleuca alternifolia seedlings, a tall shrub species used in floodplain and reservoir forest restoration in southern China, were studied during a grow season (from March to December in 2007). M. alternifolia seedlings were flooded for 180 days, drained and left to recover for another 60 days. Survival rates of the seedlings were 100% during the 180-day flooding period. Chlorophyll (Chl) content, net photosynthetic rate (P _N), stomatal conductance (g _s), and transpiration rate (E) of the flooded seedlings were all significantly lower than those of the control. Significant reductions of photochemical quenching coefficient (q_p) and increases of nonphotochemical quenching (NPQ) in the flooded seedlings were observed. However, there were no significant differences in the maximal quantum yield of PSII photochemistry (F_v/F_m) between treatments. All seedlings survived during the two-month recovery period after the flooded treatment was drained, and the biomass and height of the recovered seedlings approached those of the control at the end of the experiment. During the first-month recovery period, Chl content, P _N, g _s and E in the recovered seedlings were all obviously low, then increased gradually and rose to the levels similar to the control by the end of the experiment. Quenching analysis revealed significant reductions of q_p and increments of NPQ in the recovered seedlings at the beginning of draining, and a nearly complete recovery for both parameters by the end of the experiment. However, F_v/F_m of the recovered seedlings did not differ significantly from the control during the recovery period. Our study demonstrated that M. alternifolia seedlings can survive and grow through 180 days of flooding with a subsequent 60-day recovery period in drained conditions, indicating that seedlings of this species would be suitable for afforestation in areas exposed to intermittent flooding.     

Effects of periodic flooding and root pruning on Quercus nuttallii seedlings

In the southern United States, much of the emphasis in bottomland restoration is placed on establishing an oak-dominated forest. Artificial regeneration is an alternative for restoration on cleared lands and where a desirable seed source is not present. Currently the standard procedure for seedling preparation is to prune the roots prior to transplanting in the field. It is not fully known what effect(s) root pruning has on transplanted seedlings. In addition, bottomland restoration efforts inherently take place on floodplains. The potential interaction between root pruning and flooding on seedling performance is not known. This study consisted of two separate but related laboratory experiments. The purpose of the first experiment was to quantify the effects of various percentages of root removal and varying soil moisture regimes on transplanted Nuttall oak seedlings (Quercus nuttallii Palmer). Root pruning treatments consisted of removal of roots at 0%, 25% and 75% while soil moisture regime was maintained at non-flooded or periodically flooded conditions. Plant gas exchange, growth, and survival were measured. Root pruning alone had adverse effects on height growth during the first 72 days following transplanting. Periodic flooding also produced adverse effects on stomatal conductance (p = 0.0002), height growth (p = 0.005), and survival (p = 0.02). Photosynthetic data indicated that as pruning intensified in the periodically flooded seedlings, photosynthetic rates decreased. In contrast, as pruning intensified in the non-flooded seedlings, photosynthesis increased. This demonstrated that pruning rate had a varying effect on photosynthesis dependent upon soil moisture condition. Experiment 2 focused on the effects of varying degrees of root pruning on new root formation. The seedlings were grown under laboratory conditions, harvested at 0, 10, 20, and 30 days after treatment initiation, and analyzed for new root formation. Results of Experiment 2 indicated no difference in new root formation, root length, or root biomass due to the pruning treatment. Overall, our results from both experiments indicated that root pruning had no detectable long-term adverse effects on growth and survival of seedlings under drained soil conditions; however, as results from Experiment 1 demonstrated, if seedlings were planted in periodically flooded conditions, root pruning produced adverse effects. Thus, in restoration efforts utilizing Nuttall oak seedlings, the planting strategy and pruning rate should be carefully evaluated based on the knowledge of sites' hydrology. Alternatively, on sites with unpredictable flooding both pruned and unpruned seedlings may be utilized to ensure survival.     

Effects of Inoculum Density and Wounding on Stem Infection of Three Eastern US Forest Species by Phytophthora ramorum

Seedlings of three Eastern US forest species Quercus rubra (northern red oak), Quercus prinus (chestnut oak) and Acer rubrum (red maple) were inoculated by applying Phytophthora ramorum sporangia to stems at different inoculum densities with and without wounding. Disease occurred in all treatments involving wounds, and no disease was observed in unwounded treatments. Younger seedlings (2–3 years old) did not differ significantly from older seedlings (5–6 years old) in disease incidence, but older seedlings sustained smaller lesions compared with younger seedlings. For both old and young seedlings, disease on wounded stems was observed down to the lowest sporangia concentration utilized (500 sporangia/ml for old seedlings and 100 sporangia/ml for young seedlings). The results show that in the presence of wounding, even very low sporangia concentrations can result in disease, and further suggest that wounding caused by insects and other factors may play an important role in P. ramorum epidemiology in forest environments.     

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.