Restoring longleaf pine (Pinus palustris Mill.) in loblolly pine (Pinus taeda L.) stands: Effects of restoration treatments on natural loblolly pine regeneration

Historical land use and management practices in the southeastern United States have resulted in the dominance of loblolly pine (Pinus taeda L.) on many upland sites that historically were occupied by longleaf pine (Pinus palustris Mill.). There is currently much interest in restoring high quality longleaf pine habitats to such areas, but managers may also desire the retention of some existing canopy trees to meet current conservation objectives. However, fast-growing natural loblolly pine regeneration may threaten the success of artificially regenerated longleaf pine seedlings. We evaluated the establishment and growth of natural loblolly pine regeneration following different levels of timber harvest using single-tree selection (Control (uncut, residual basal area ∼16 m²/ha), MedBA (residual basal area of ∼9 m²/ha), LowBA (residual basal area of ∼6 m²/ha), and Clearcut (complete canopy removal)) and to different positions within canopy gaps (approximately 2800 m²) created by patch cutting at two ecologically distinct sites within the longleaf pine range: Fort Benning, GA in the Middle Coastal Plain and Camp Lejeune, NC in the Lower Coastal Plain. The density of loblolly pine seedlings was much higher at Camp Lejeune than at Fort Benning at the end of the first growing season after harvesting. Following two growing seasons, there were no significant effects of canopy density or gap position on the density of loblolly pine seedlings at either site, but loblolly pine seedlings were taller on treatments with greater canopy removal. Prescribed fires applied following the second growing season killed 70.6% of loblolly pine seedlings at Fort Benning and 64.3% of seedlings at Camp Lejeune. Loblolly pine seedlings were generally less than 2 m tall, and completeness of the prescribed burns appeared more important for determining seedling survival than seedling size. Silvicultural treatments that include canopy removal, such as patch cutting or clearcuts, will increase loblolly pine seedling growth and shorten the window of opportunity for control with prescribed fire. Therefore, application of prescribed fire every 2-3 years will be critical for control of loblolly pine regeneration during restoration of longleaf pine in existing loblolly pine stands.     

Root growth and hydraulic conductivity of southern pine seedlings in response to soil temperature and water availability after planting

Comparison of the root system growth and water transport of southern pine species after planting in different root-zone environments is needed to guide decisions regarding when, and what species to plant. Evaluation of how seed source affects root system responses to soil conditions will allow seed sources to be matched to planting conditions. The root growth and hydraulic conductivity of three sources each of shortleaf, loblolly and longleaf pine seedlings were evaluated for 28 days in a seedling growth system that simulated the planting environment. Across species, an increase in root-zone temperature alleviated limitations to root growth caused by water stress. In the coldest temperature, longleaf pine maintained a higher hydraulic conductivity compared to shortleaf and loblolly pine. Without water limitation, the root growth and hydraulic conductivity of shortleaf and loblolly pine were superior to that of longleaf pine, but as water availability decreased, the root growth of longleaf pine surpassed that of loblolly pine. Hydraulic conductivities of the seed sources differed, and differences were attributed to either new root growth, or an increase in the efficiency of the root system to transport water.     

Aluminum sensitivity of loblolly pine and slash pine seedlings grown in solution culture

To probe variation in Al sensitivity of two co-occurring pine species, seedlings from six full-sib families of loblolly pine (Pinus taeda L.) and slash pine (Pinus elliottii Engelm.) were grown in solution culture containing 4.4 mM (high-Al) or 0.01 mM (low-Al) AlCl(3) at pH 4 for 58 days. On average, both pine species had 41% less total dry weight in the high-Al treatment than in the low-Al treatment. Stem volume growth of slash pine was more sensitive to the high-Al treatment than that of loblolly pine. In both species, the high-Al treatment inhibited root dry weight more than shoot dry weight. Within-species variation in Al sensitivity among families was greater in loblolly pine (24 to 52% inhibition of seedling dry weight) than in slash pine (35 to 47% inhibition of seedling dry weight). Foliar Al concentration was positively correlated with Al sensitivity in slash pine but not in loblolly pine; however, in both species, the concentration of Al in roots was 20-fold greater than in foliage.     

Comparing morphology and physiology of southeastern US <Emphasis Type="Italic">Pinus</Emphasis> seedlings: implications for adaptation to surface fire regimes

Key message

The suite of traits expressed as seedlings by coastal and mountain longleaf pine and south Florida slash pine suggest they can survive fire in the seedling stage. In contrast, loblolly pine and typical slash pine tolerate fire when mature but do not exhibit traits that allow them to survive fire when young, representing a different strategy for survival in frequently burned communities.

Context

Fire is an important driver in the distribution and abundance of southern US pine species, and seedling fire tolerance often determines individual survival under frequent fire regimes.

Aims

We investigated seedling growth, biomass allocation, needle distribution, bark thickness, and total non-structural carbohydrate (TNC) storage in taproots and related them to the expression of fire-tolerance for five species or types, including loblolly pine (Pinus taeda L.), two longleaf pine (P. palustris Mill.) types representing two distinct ecological communities (coastal and mountain) and two slash pine (P. elliottii Englem.) varieties.

Methods

We analyzed the relationship of seedling growth, biomass characteristics, and total non-structural carbohydrate storage between species by using analysis of variance.

Results

Both coastal and mountain longleaf pines had thick bark, long, densely arranged needles, and a grass-stage. South Florida slash pine shared the same suite of traits but, contrary to previous reports, displayed reduced height growth rather than a grass-stage. In contrast, loblolly pine and typical slash pine had faster height growth, more branching, lower needle density, and thinner bark. Both longleaf pines and south Florida slash pine also had higher TNC storage in taproots than either loblolly or typical slash pines.

Conclusion

The relative strength of expression of these fire-adaptation traits among the five species types generally matches the fire-return intervals associated with each species’ habitat, suggesting the importance of fire regimes in determining the distribution and abundance of the studied species.

     

Comparison of container-grown Pinus taeda L. seedlings raised outdoors and in a growth chamber in sunlight

Two annual crops of container-grown loblolly pine (Pinus taeda L.) seedlings raised outdoors differed from those grown inside a covered chamber with a clear top. After a 90-day growth period, outside-grown container seedlings had shorter heights, smaller root collar diameters, and lower root, stem, foliage, shoot, and total dry weights than chamber-grown seedlings. Fall-planted, outside-grown seedlings raised the first year had up to 10% higher field survival than those from the chamber. However, chamber-grown container seedlings that survived retained their initial height and diameter advantages following one complete growing season. Fall-planted seedlings raised the second year showed no differences due to growth treatments when favorable weather for seedling growth occurred after field-planting. Container-grown seedlings raised outdoors during the summer months are well suited for fall-outplanting.     

Use of waste materials as nursery growing media for <Emphasis Type="Italic">Pinus halepensis</Emphasis> production

Peat moss is gradually being replaced by other materials as a growing medium in forest nurseries due to economic and ecological constraints. In this study, six different mixtures were tested, mixing peat moss (P) and pine bark (B) with digested sewage sludge (S) activated sewage sludge (A) and paper mill sludge (M), as growing media for Pinus halepensis seedlings; three different waste doses were applied. Seed germination percentage, seedling growth and foliar nutrient content after 1 year in a greenhouse and percentage survival after transplanting were recorded. The influence of base substrate (P or B) on germination percentage changed in different ways according the type of waste. The order of the different applied mixtures by suitability (germination rate and seedling growth) from best to worst was as follows: activated sludge with peat, activated sludge with pine bark, sewage sludge with peat, sewage sludge with pine bark, paper mill sludge with pine bark and finally paper mill sludge with peat.     

Container seedlings outperform barefoot stock: Survival and growth after 10 years

Ponderosa pine, Jeffrey pine, and Douglas-fir seedlings were planted in container or bareroot form at three elevations in northern California. At the lowest elevation (762 m), container seedlings of ponderosa pine were significantly taller than bareroot seedlings at ages 4 and 10, and had breast-height diameters that were significantly larger than bareroot counterparts at age 10. Survival of Douglas-fir container seedlings was significantly greater than that of barefoot seedlings for all ages tested. At the mid-elevation site (1220 m), container seedlings of ponderosa pine and Douglas-fir were significantly larger in breast-height diameter than bareroot seedlings at age 10. Douglas-fir container seedlings survived significantly better at all ages than barefoot seedlings. At the highest elevation (1662 m), seedling height and diameter did not differ significantly, but survival of container seedlings was significantly higher than barefoot seedlings for both pine species at all ages tested.     

Root and shoot allometry of bareroot and container Douglas-fir seedlings

Detailed root and shoot development of bareroot and container Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco) seedling stocktypes were compared during the first growing season after outplanting. The study was installed in raised beds with ideal environmental conditions and at a field reforestation site. Survival at both sites was 98% and did not differ between stocktypes. Seedlings were excavated in spring (5, 8, 12, and 16 weeks after planting) and in fall (35, 40, and 45 weeks after planting). In spring, container seedlings had more numbers of new roots and greater new root and shoot biomass than bareroot seedlings at both sites. In fall, bareroot seedlings consistently averaged more new root growth (though nonsignificant) than container seedlings suggesting that stocktype differences may not continue long-term. Container seedlings had significantly greater water percent than bareroot seedlings at the field site (all sample dates) and the raised bed site (weeks 5, 8, and 40 only). Regardless of environmental conditions or season, seedlings at both sites maintained water percent between 60 and 70% of fresh weight. Seedlings grown in the raised beds had much greater growth than those grown in the field. However, relative growth patterns for the two stocktypes were very similar on each site. The data generated establish baseline differences between stocktypes for root initiation, growth, and allometry during the first year after planting. Challenges associated with root development research are discussed.     

Morphological characteristics associated with tolerance to competition from herbaceous vegetation for seedlings of jack pine,black spruce,and white pine

Tolerance of bareroot and container-grown seedlings of black spruce (Picea mariana (Mill.) B.S.P.), jack pine (Pinus banksiana Lamb.), and eastern white pine (Pinus strobus L.) to competition from herbaceous vegetation was examined in the first five years after planting on a site in the Great Lakes/St. Lawrence forest of Ontario, Canada. Shoot and root morphological characteristics of various stocktypes were measured before planting and correlated with 5-year survival and growth following control and no control of herbaceous vegetation. For black spruce and jack pine, medium-sized bareroot stocktypes had greater relative 5-year stem volume growth in the presence of herbaceous vegetation than did container stock of either species or large bareroot stock of spruce. Relative volume growth was measured as the ratio of the cumulative stem volume increment in the presence of vegetation (Veg) to that in the absence of vegetation (NoVeg), i.e., the Veg:NoVeg ratio. In white pine, the Veg:NoVeg ratio of volume increment of medium container and large bareroot stocktypes exceeded that of small container and medium bareroot stocktypes. In jack pine, root collar diameter at planting and number of first-order lateral roots were positively correlated with 5-year Veg:NoVeg ratio of volume increment. In white pine, the Veg:NoVeg ratio was also positively correlated with root collar diameter at planting and with root volume. In black spruce, the ratio was not related to pre-plant morphology. Thus, for white pine and jack pine, certain pre-plant morphological features may be useful in forecasting the relative ability of different stocktypes to grow under herbaceous competition conditions in the field.     

火炬松和湿地松幼苗蔗糖含量与生长潜势关系的研究初报

火炬松(Pinux taeda L.)和湿地松(Pinus elliottii Engelm)是我国南方重要的外引用材树种,经过多年的试验研究,已选出一批较好的种源、家系和单株,并建立了种子园。当前早期鉴定和早期选择已成为必不可少的一项研究内容,近几年来越来越多的研究者指出,林木幼龄期的某些性状或生理生化指标与成熟期性状之间存在一定的相关性,认为树木生长的早期选择是可能的,尤其从生理生化方面来探索林木早期预测的研究已引起人们的高度重视。但     

Effect of uninucleate Rhizoctonia on Scots pine and Norway spruce seedlings

One‐year‐old container‐grown seedlings were planted in spring on clear cut areas: the Norway spruce (Picea abies) on a moist upland site (Myrtillus‐type) and Scots pine (Pinus sylvestris) on a dryish upland site (Vaccinium‐type). While still in the nursery, half of the seedlings of each species had been inoculated during the previous summer, with a uninucleate Rhizoctonia sp., a root dieback fungus. At outplanting all the seedlings appeared healthy and had a normal apical bud, although the height of the inoculated seedlings was less than that of the uninoculated control seedlings. At the end of the first growing season after planting, the mortality of inoculated Scots pine and Norway spruce seedlings was 25 and 69%, respectively. After two growing seasons the mortality of inoculated seedlings had increased to 38% for Scots pine and 93% for Norway spruce. The mortality of control seedlings after two growing seasons in the forest was 2% for Scots pine and 13% for Norway spruce. After outplanting the annual growth of inoculated seedlings was poor compared with the growth of control seedlings. These results show that, although Rhizoctonia‐affected seedlings are alive and green in the nursery, the disease subsequently affects both their survival and growth in the forest.     

A summary of pine straw yields and economic benefits in loblolly, longleaf and slash pine stands

Forest landowners in the southeastern United States have the opportunity to manage their loblolly, longleaf and slash pine stands for pine straw (fresh undecomposed needles; the litter layer) for non-timber revenues. Pine straw is used primarily as mulch in landscaping and has grown in revenues paid to landowners in Georgia from $15.5 million in 1999 to $81 million in 2009. Pine straw is typically sold by the acre or by the bale. Selling pine straw by the acre may be advantageous to absentee landowners. Selling pine straw by the bale can generate more annual income, but bale counts need to be accurate and bale dimensions defined. For both methods, recent (2005–2010) pine straw multi-year revenues range from $50 to $150 per acre annually. Longleaf pine straw commands the highest price per bale, followed by slash pine, and lastly loblolly pine. Per rake yields from loblolly stands tend to be 15–30?% greater than slash and longleaf pine. Pine straw raking typically starts at canopy closure continuing to the first thinning, generating from $300 per acre to over $1000 per acre in new income. This paper summarizes pine straw yields and economics in loblolly, longleaf, and slash pine stands.     

Influence of different nursery container media on rooting of Scots pine and silver birch seedlings after transplanting

Scots pine (Pinus sylvestris L.) and silver birch (Betula pendula Roth.) seedlings were grown in containers filled with growth media based on medium-textured sphagnum peat, coarse perlite and fine sand. The seedlings were then planted into fine and coarse sandy soils in 2.2 l pots, which were subjected to two water-content treatments (only one for birch). After the seedlings had grown five weeks in a greenhouse, rooting into the surrounding soil and shoot growth were measured. Addition of perlite and sand to peat medium slightly affected rooting; thus suggesting minor effects on seedling establishment. However, nitrogen concentration of the seedlings varied between growth media and correlated positively with rooting into the soil. The particle size and water content of the soil affected considerably rooting of the seedlings. Seedling height at the time of planting did not affect rooting or shoot growth. The fact that the fewest out-grown roots occurred in the dry fine sandy soil, suggests that dry soil together with high strength and resistance to root penetration reduce rooting and water uptake by container seedlings most and may thus cause water and nutrient stresses to seedlings after outplanting.     

Container characteristics influence Pinus pinea seedling development in the nursery and field

The growth and development of Pinus pinea seedlings grown in different containers was followed through one growing season in the nursery and 3 years following outplanting in the field. The variables studied in the nursery were height, diameter, biomass of shoots and roots, nutrient uptake and root density. The measured field variables, height and diameter increment and survival, were correlated with the nursery variables. Container volume had the greatest influence on plant morphology. Containers with larger rooting volume had seedlings with larger height and diameter, greater nutrient content, and better field performance. Growing density was correlated with seedling morphology and nutrient concentration in the nursery. Among the variables that influenced container volume, the diameter of the container was the most important, while the depth of the container had a minor influence on seedling morphology.The best indicator of seedling development in the nursery was the ratio of container depth to container diameter, and the optimum ratio was 4. All containers produced seedlings with some root spiralling, including those containers with ribs. There was no relationship between either the number of spiralling roots or the angle of spiralling and container characteristics. Furthermore, root spiralling did not influence seedling performance following outplanting. Root density (root biomass/cm³) was inversely correlated with container volume but there was no correlation with either depth or growing density. The largest plants were produced with container volumes of 300–400 cm³, depth/diameter ratios of 4, and growing densities of 200–300 seedlings/m². These growing conditions will result in larger Pinus pinea seedlings coming out of the nursery, which will increase growth following outplanting.     

Light quality treatments enhance somatic seedling production in three southern pine species

Embryogenic cultures of loblolly pine (Pinus taeda L.), slash pine (Pinus elliottii Engelm.), longleaf pine (Pinus palustris Mill.) and slash pine x longleaf pine hybrids were initiated from immature seeds on an initiation medium containing 13.57 microM 2,4-dichlorophenoxyacetic acid and 2.22 microM benzylaminopurine. Embryogenic cultures proliferated and somatic embryos developed, matured and germinated following a modified protocol and media originally developed for radiata pine (Pinus radiata D. Don.) somatic seedling production. A discrete, light-sensitive pre-germination stage and a later germination (radicle emergence) stage were identified by the differential response of somatic embryos to light of different wavelengths. Different light quality treatments were applied during the pre-germination and germination steps, using cool white fluorescent bulbs or light-emitting diodes (LEDs), or both. In general, red wavelengths provided by LEDs during these steps resulted in higher frequencies of somatic embryo germination (up to 64%) and conversion (up to 50%), longer tap roots and more first-order lateral roots than the standard cool white fluorescent treatments or treatment with blue wavelengths from LEDs. In addition, exposure to red light allowed germination of somatic embryos of some clones that failed to produce germinants under fluorescent light. Germination and conversion were further enhanced by sequential application of cool white fluorescent light and red light, resulting in up to 100% germination and conversion in one experiment. Longleaf pine somatic embryos were especially responsive to the light quality treatments, resulting in the first report of somatic seedling production for this species.     

Carbohydrate reserves in Pinus sylvestris seedling needles as an attribute of seedling vigor

The carbohydrate reserve of bareroot Scots pine (Pinus sylvestris L.) seedlings was evaluated as means of describing seedling quality and capacity to withstand stress during nursery lifting, outplanting and acclimatization at the planting site. Carbohydrate concentrations in needles were followed through four spring storage treatments and the postplanting success and growth of seedlings monitored at two sites. The reduction in total glucose concentration in needles was affected by storage time and temperature. Depletion of carbohydrate reserves decreased needle growth, reduced the number of terminal buds, disturbed leader shoot formation, and consequently reduced shoot length. The depletion of reserves below a threshold concentration of ～2 % total glucose during storage resulted in significant seedling mortality. A simple model for estimating the depletion of carbohydrate reserves based on dark respiration was evaluated as a measure of the accumulated strain during the storage. The application of carbohydrate analyses as a routine test of seedling quality in a nursery is not currently feasible because of the insensitivity and laboriousness of the method.     

The effect of weed control and fertilization on survival and growth of four pine species in the Virginia Piedmont

The growth response of loblolly pine (Pinus taeda L.), shortleaf pine (Pinus echinata Mill.), Virginia pine (Pinus virginiana Mill.), and white pine (Pinus strobus L.) to weed control and fertilization in the Piedmont of Virginia was assessed. Four different silvicultural treatments were evaluated: (1) check (no treatment); (2) weed control; (3) fertilization; (4) weed control plus fertilization. The weed control treatment included a series of herbicide and mechanical treatments to eliminate competing hardwoods. The fertilizer treatments added N, P, K, and S. Survival and growth was measured annually through age 5. There were significant differences in survival and growth among species. Survival was greatest for loblolly pine, lower in shortleaf and Virginia pine, and lowest in white pine. Fertilization without controlling the competing hardwoods decreased survival in all planted pines due to the increased hardwood competition. Loblolly pine was tallest through the 5-year period, shortleaf and Virginia pine were shorter and white pine was shortest. Silvicultural treatments had no impact on tree height but significantly affected DBH. Weed control increased DBH while fertilization did not. When applied in combination with weed control, there was no additional increase in growth of the pines due to fertilization beyond that from weed control only. Fertilization stimulated the growth of the competing hardwoods which were significantly taller in the fertilized plots.     

Effects of Acidic Precipitation and Ectomycorrhizal Inoculation on Growth,Mineral Nutrition,and Xylem Water Potential of Juvenile Loblolly Pine and White Oak

Abstract

Individual and interactive effects of simulated acidic rainfall and mycorrhizal inoculation on growth and nutrient and water relations of loblolly pine (Pinus taedaL.) and white oak (Quercus albaL.) grown in a loam soil were examined. Seedlings of each species inoculated with basidiospores of the ectomycorrhizal fungus Pisolithus tinctorius(Pers.) Coker and Couch, a known my-cobiont of both loblolly pine and white oak, and uninoculated control seedlings received two simulated rains per week of either pH 3.6, 4.2, or 4.8 for 26 weeks. Higher acidity rainfall reduced the growth but increased mycorrhizal colonization of loblolly pine, while both loblolly pine and white oak exposed to these rains exhibited greater foliar injury. Inoculation with P. tinctoriusincreased growth and reduced foliar injury of both species. Foliar concentrations of P, S, and Cu in loblolly pine and white oak, Ca in loblolly pine, and Fe and Zn in white oak decreased with increasing rain acidity while the Al concentration of both species increased. Higher rainfall acidity also reduced soil pH and Ca and Mg concentrations while increasing soil AI. Foliage of inoculated seedlings of both species had higher N and P concentrations and lower Al concentrations than control seedlings. Following the final rain applications, a drought cycle was simulated by withholding irrigation for two weeks during which seedling xylem pressure potential and soil water potential were measured. One day after cessation of irrigation, xylem pressure potential of loblolly pine that had received pH 3.6 rains was lower than that of other treatments. Thereafter, xylem pressure potential and soil water potential of the inoculated treatment decreased below those of the control treatment in both species. These results suggest that acid deposition is detrimental to juvenile loblolly pine and white oak, but the magnitude of this effect is less than the positive response to ectomycorrhizal inoculation.     

Pine growth response to management of the subterranean clover understory

Management of subterranean clover (Trifolium subterraneum L.) understory grown with loblolly pine (Pinus taeda L.) and slash pine (Pinus elliottii Engelm.) resulted in differences in both clover yield and pine diameter. Loblolly and slash pines were planted into a subterranean clover pasture in 1984. Suppression of warm season herbaceous vegetation by applying herbicides or disking the site in late summer resulted in significantly greater subterranean clover production than on the control site where no additional understory management was applied. Pine species had no effects on clover yield. The N content of the pine foliage on the blocks managed with herbicide application or disking generally was significantly greater than the control from the 5th through 7th growing seasons (1988–1990). The dbh of the pine trees in the disked and herbicide treatments was greater than the control after the 7th growing season (1990).     

Aluminum fractions in root tips of slash pine and loblolly pine families differing in Al resistance

Aluminum (Al) distribution among several cellular fractions was investigated in root tips of seedlings of one Al-resistant and one Al-sensitive family of slash pine (Pinus elliottii Engelm.) and loblolly pine (Pinus taeda L.) grown in nutrient solution containing 100 microM AlCl3 (pH 4) for 167 h. Aluminum present in 5-mm-long root tips was fractionated into cell-wall-labile (desorbed in 0.5 mM citric acid), cell-wall-bound (retained after filtering disrupted cells through 20-microm mesh) and symplasmic (filtrate following cell disruption) fractions. When averaged across both species, 12% of Al absorbed by root tips appeared in the symplasmic fraction and 88% in the apoplasmic fraction (55% as cell-wall-labile, and 33% as cell-wall-bound). On a fresh mass basis, total Al in root tips was lower in loblolly pine than in slash pine, lower in the Al-resistant slash pine family than in the Al-sensitive slash pine family, and lower in the Al-resistant families than in the Al-sensitive families across species. Although the data support the hypothesis that Al-resistant plants limit Al uptake to root apices, they do not exclude other mechanisms of Al resistance. Differential Al resistance between the species and between slash pine families may also be associated with the size of the total non-labile and cell-wall-labile Al fractions, respectively. We were unable to identify the basis for differential Al resistance in loblolly pine.