Breeding eastern white pine: A world-wide perspective

Eastern white pine (Pinus strobus L.) has wide genetic variability and high productivity in plantations in North America, Europe and the Far East. In regions where neither white pine blister rust nor the white pine weevil are a problem, as in the central states of the U.S.A. and the warmer parts of Europe, primary attention is given to selection and breeding for vigor. Estimates for trees up to age 18 indicate that a first-generation genetic gain in volume of over 20% can be obtained from open-pollinated family selection within stands. Volume gains of 50% or more over local genotypes have been realized at ages 15–20 through the use of seed from selected southern Appalachian stands and in some individual tree crosses of eastern white pine with blue (Himalayan) pine. In the cool, humid regions of eastern North America and Europe, where blister rust resistance is important, breeding strategies include cloning of resistant P. strobus individuals, use of blue and Balkan pines in breeding programs, and exploitation of low-level rust resistance in low-risk localities. Eastern white pine losses from blister rust can be avoided in Japan and Korea by good plantation management without genetic selection. In some regions where the white pine weevil is the major problem, western white pine is useful both for hybridization and as an alternative, more weevil-resistant species. Analysis of relative photosynthesis of SO₂- and O₃-fumigated clones may be useful as a method of screening for pollution tolerance in regions where air pollution is likely to intensify. Efficient techniques for cloning by rooting of cuttings are now available. Efforts are under way to develop technology for micropropagation for mass production of superior clones. Breeding strategies for the major P. strobus plantation regions of the world are summarized.     

White pine (Pinus strobus L.) regeneration dynamics at the species’ northern limit of continuous distribution

The abundance of eastern white pine (Pinus strobus L.) has been significantly reduced across its distribution range over the past few centuries. The species’ regeneration dynamics is well documented in the centre of its range, but is poorly understood at the northern limit of continuous distribution. To address this knowledge gap, we quantified natural white pine regeneration in unmanaged mature stands, identified the most important variables influencing it, and evaluated the impact of damaging agents, namely white pine blister rust (Cronartium ribicola J.C. Fisch.), white pine weevil (Pissodes strobi Peck), and herbivory. We also quantified the influence of remnant stands and residual trees on the spatial distribution of regeneration in logged sites. The results reveal continuous but low recruitment in mature stands. The basal area of balsam fir (Abies balsamea (L.) Miller) had a strong negative effect on white pine regeneration. Regeneration was more abundant than expected on moister substrates, including moss, decaying wood and organic matter. White pine regeneration was noted in recently logged areas, where distance from remnant stands had a significant effect on the abundance of white pine regeneration. The northern limit of continuous distribution holds potential for white pine restoration, for example by preserving remnant white pine stands that can provide seed sources for natural regeneration in a shelterwood cut system, or in adjacent clearcut areas. This study illustrates that different management strategies should be used near northern range limits, where effects of site conditions and disturbance agents are different than in the center of a species’ range.     

Regeneration of Rocky Mountain bristlecone pine (Pinus aristata) and limber pine (Pinus flexilis) three decades after stand-replacing fires

Rocky Mountain bristlecone pine (Pinus aristata) and limber pine (Pinus flexilis) are important high-elevation pines of the southern Rockies that are forecast to decline due to the recent spread of white pine blister rust (Cronartium ribicola) into this region. Proactive management strategies to promote the evolution of rust resistance and maintain ecosystem function require an improved understanding of the role of disturbance on the population dynamics of both species and environmental conditions that favor seedling establishment. We examined patterns of bristlecone and limber pine regeneration across the perimeters of three, 29-year-old, high-severity burns in northern, central, and southern Colorado: Ouzel, Badger Mountain, and Maes Creek, respectively. Both species exhibited a very protracted regeneration response to these fires. Bristlecone pine regeneration was concentrated near burn edges and beneath surviving seed sources. This spatial pattern is consistent with limitations incurred by wind-dispersal, also borne out by the low occurrence of seedling clusters. Relative to unburned stands, the absolute abundance of bristlecone pine generally increased only on plots retaining some surviving trees. Limber pine regeneration pattern varied between sites: high in the burn interior at Ouzel, concentrated at burn edges at Badger, and mostly in unburned stands at Maes. Clark’s Nutcracker dispersal of limber pine in each study area was indicated by high seedling distance from possible seed sources and high frequencies of clustered stems. Except at Ouzel, the absolute abundance of limber pine decreased in burns. Across sites, establishment by both species was boosted by nearby nurse objects (rocks, fallen logs, and standing tree trunks), a relationship that extended out at least as far as the closest three such objects, usually found within 50 cm. Fire decreased the frequency of Pedicularis but increased Castilleja and Ribes species (alternate hosts of white pine blister rust), though only one species, R. cereum, was positively associated with either pine species. We conclude that regeneration of bristlecone and limber pine may benefit from natural disturbance or proactive management creating appropriately sized openings and microtopographic structure (e.g., abundant fallen logs); however, beneficial responses may require many decades to be achieved.     

Regeneration dynamics of a mixed Mediterranean pine forest in the absence of fire

The aim of this study is to determine the competing regeneration and expansion patterns of two co-occurring pine species (Pinus brutia, Pinus nigra ssp. pallasiana), in a transitional montane Mediterranean zone. We measured the regeneration density of all woody species in 102 randomly located stands along an altitudinal gradient on the island of Lesbos, Greece. Individuals of pines were assigned to different size classes. Topographic factors (altitude, aspect, and soil depth) and light availability (through hemispherical photographs) were measured for each stand. Statistical analyses were applied to explore the effect of each factor on recruitment density of the competing pine species, and to elucidate patterns of interaction. Canopy openness was the most important parameter controlling the recruitment of P. brutia, while the regeneration density of P. nigra was mainly related to canopy openness and heat load. An idiosyncratic response of the recruitment vigour of the two species was identified along gradients of shade and drought stress. The decline in P. nigra recruitment density with drought conditions underlines threats to its population maintenance even in the absence of fire. On the other hand P. brutia seems to be a stronger invader in transitional zones. The studied species could be considered typical representatives of the two most widely distributed pine functional types across the Mediterranean basin, and our results agree with the theoretical ability of such species to maintain and expand their populations.     

Conservation and use of the Pinus chiapensis genetic resource in Colombia

The species Pinus chiapensis is under severe pressure from forest conversion to agriculture and fuelwood cutting. These trends are certain to continue and methods for conservation and wise use of the genetic resource must be implemented soon in Mexico and Guatemala as well as in those countries where the species can be grown on a commercial basis. A total of 101 open-pollinated families from nine provenances of Pinus chiapensis have been established in trials and gene conservation banks by Smurfit Carton de Colombia since 1981. The best provenances for growth rate have been Cotzal, La Trinidad and Pohlo. A tree improvement program with grafted seed orchards is underway.     

Regeneration patterns of Eastern white pine (Pinus strobus L.) in hardwood-dominated forests in Virginia, USA

Eastern white pine (Pinus strobus L.) is a moderately shade-tolerant species that co-occurs with hardwood tree species in many forests of the eastern United States, as well as in pure stands. The species is valued for its timber, as well as for wildlife and recreation. Regeneration of this species is somewhat unpredictable and often occurs in patches of similarly-aged cohorts. We described the regeneration patterns of this species and examined their relation to environmental variables within hardwood forests of southwestern Virginia, USA. An average of 5.3 white pine patches per ha were observed in this study. The majority of patches consisted of saplings (85%), with 9% of patches in pole size classes, and 6% in seedling size classes. The average density of patches was 43.5 stems with an average age of 20 years. The size of patches averaged 80.6 m². The total density of seedlings and the number of regeneration patches of all sizes of regeneration (seedlings, saplings, and poles) in plots was related to the surrounding density of large white pine trees (potential seed trees). The density of seedlings or patches was not significantly related to current vegetation cover or soil surface cover variables, but more than half of regeneration patches were located in or adjacent to old canopy gaps, most of which were old logging gaps. While seedling regeneration may occur within the understory of these forests near seed trees, advancement to the sapling and pole stage appears to be associated with canopy gap formation.     

Diversity of northern plantations peaks at intermediate management intensity

Tree diversity is an important component of biodiversity. Management intensification is hypothesized to affect tree diversity. However, evidence to support the relationship between management intensity and tree diversity in northern forests is lacking. This study examined the effects of fertilization, site preparation, and brush control on tree species diversity, shade tolerance diversity and size diversity of jack pine (Pinus banksiana Lamb.), black spruce (Picea mariana [Mill.] B.S.P.), white pine (Pinus strobus L.) and white spruce (Picea glauca [Moench] Voss) plantations, 15 years after planting in Ontario, Canada. Species diversity and shade tolerance diversity were highly correlated, so were diameter size diversity and height size diversity. Fertilization did not affect the tree diversity indices of any plantations. Species diversity and shade tolerance diversity was interactively influenced by site preparation and brush control in the black spruce, white pine, and white spruce plantations, showing that the highest diversity occurred on sites with intensive site preparation without brush control, whereas on sites with brush control, diversity was higher with least intensity of site preparation. However, in the jack pine plantation, neither species diversity nor shade tolerance diversity differed with management intensification, and is attributed to the fast capture of site resources by the planted crop trees of jack pine which minimized establishment of non-crop species. Tree size diversity increased with site preparation intensity in the jack pine and black spruce plantations, while it decreased with brush control in the white pine and white spruce plantations. We concluded that (1) the effects of management intensification on diversity of northern plantations differ with growth habit of planted crop tree species and (2) species diversity and tree size diversity tend to be highest at intermediate levels of silvicultural intensification during the stand establishment phase, supporting the intermediate disturbance hypothesis.     

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

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

Natural Variation in Seed Characteristics of Eastern White Pine (Pinus strobus L.)

Natural variation in seed characteristics of eastern white pine (Pinus strobus L.) trees and its relationship with the size, form, and crown class of parent trees was examined in second-growth, white pine mixedwood stands in Ontario. Cones were collected prior to and following partial harvesting of two stands during 2 years of above-average seed production. Fully enlarged seed extracted from mature cones was X-rayed to determine seed yield per cone, the percentage of sound, partially filled, and empty seed, and the sound seed mass for each tree. Variation in seed yield per cone, the proportion of sound, partially filled, and empty seed, and seed mass was much greater among individuals than between stands or seed years. The strength and significance of the relationship of seed mass with sound seed yield differed with stand and seed year. In uncut stands, seed mass was higher for trees with dominant crown class, larger live crown ratios, and smaller height to diameter ratios, suggesting tree vigour and microsite quality influence seed mass. Proportion of partially filled seeds was significantly reduced in both stands in the second year likely due to increased growing degree days during female gametophyte development. The implications of these results to natural and artificial regeneration of white pine are discussed.     

Radiation use efficiency in adjacent hardwood and pine forests in the southern Appalachians

The efficiency with which trees convert photosynthetically active radiation (PAR) to biomass has been shown to be consistent within stands of an individual species, which is useful for estimating biomass production and carbon accumulation. However, radiation use efficiency (?) has rarely been measured in mixed-species forests, and it is unclear how species diversity may affect the consistency of ?, particularly across environmental gradients. We compared aboveground net primary productivity (ANPP), intercepted photosynthetically active solar radiation (IPAR), and radiation use efficiency (? = ANPP/IPAR) between a mixed deciduous forest and a 50-year-old white pine (Pinus strobus L.) plantation in the southern Appalachian Mountains. Average ANPP was similar in the deciduous forest (11.5 Mg ha⁻¹ y⁻¹) and pine plantation (10.2 Mg ha⁻¹ y⁻¹), while ? was significantly greater in the deciduous forest (1.25 g MJ⁻¹) than in the white pine plantation (0.63 g MJ⁻¹). Our results demonstrate that late-secondary hardwood forests can attain similar ANPP as mature P. strobus plantations in the southern Appalachians, despite substantially less annual IPAR and mineral-nitrogen availability, suggesting greater resource-use efficiency and potential for long-term carbon accumulation in biomass. Along a 260 m elevation gradient within each forest there was not significant variation in ?. Radiation use efficiency may be stable for specific forest types across a range of environmental conditions in the southern Appalachian Mountains, and thus useful for generating estimates of ANPP at the scale of individual watersheds.     

Effects of light/darkness, thermal shocks and inhibitory components on germination of Pinus canariensis, Pinus halepensis and Pinus pinea

Knowledge of the germination characteristics of Pinus species can help in the understanding, prediction and management of the regeneration of pine forests. In the Canary Islands, several exotic pines (Pinus halepensis L. and Pinus pinea L.) were planted with Pinus canariensis Chr. Sm. Ex DC, the only native pine species, and there is now an interest in controlling these exotic species to restore the original forest. The main objective of the present study is to determine the germination response of P. canariensis, P. halepensis and P. pinea to different light regimes (darkness and light/darkness), thermal shocks and the presence of inhibitory substances from the leaves and litter of P. canariensis. P. halepensis seeds showed the highest viability and germination rate. Darkness accelerated P. halepensis germination, while exudates accelerated P. canariensis germination. Only treatments of 200 °C for 5 min and 300 °C for 5 min significantly decreased the germination of all three pine species. In the absence of strong differences in germination among species after treatments, the key for fire management or prescribed burning in this case may be the sprouting ability of P. canariensis. A possible management strategy to control the two obligate seeder exotic species could be to use one intense prescribed fire followed by a low-intensity prescribed fire after seed bank germination in the field, so in this way, only P. canariensis could resprout after fire and it would eliminate only the exotic pines in a mixed stand.     

Blister rust in Romania

In a survey of the incidence of Cronartium ribicola in Romania, this parasite was found to be present all over the country, except in mountain regions. The heaviest attacks occurred in Ribes nigrum and in young plantations of Pinus strobus, whereas old plantations of P. strobus and other pine species were free from blister rust, as were natural populations of Pinus cembra, Ribes alpinum, and R. petraeum in the Carpathians.     

Use of carbofuran for control of eastern white pine cone and seed insects

The major insects that destroy cones and seeds of eastern white pine, Pinus strobus L., were controlled with soil applications of the systematic insecticide carbofuran in orchards at Murphy and Morganton, NC, U.S.A. Carbofuran applied once a year at rates of 4.5, 9.0 or 13.5 g AI (active ingredient) per cm of diameter at breast height protected trees from the white pine cone beetle, Conophthorus coniperda (Schwarz), the leaffooted pine seed bug, Leptoglossus corculus (Say), the shieldbacked pine seed bug, Tetyra bipunctata (Herrich-Schäffer), and the white pine cone borer, Eucosma tocullionana Heinrich. Control was reflected by increased cone survival and increased yields of filled seeds per cone. Increasing the rate of application from 4.5 to 9.0 or 13.5 g generally improved control, but the improvement was not of statistical or practical significance. Carbofuran was ineffective against a seed chalcid, Megastigmus atedius (Walker). White pine clones varied in inherent susceptibility to insect attacks.     

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.     

Importance of Site Type and Tree Species on Disease Incidence of Gremmeniella abietina in Areas with a Harsh Climate in Northern Sweden

The importance of site type and tree species on disease incidence of Gremmeniella abietina was investigated. Blocks of four conventional regenerations of Pinus contorta and Pinus sylvestris, situated on sites formerly supporting pure stands of P. sylvestris and Picea abies respectively, were examined for the occurrence of G. abietina. Trees planted on spruce sites were found to be more severely diseased by G. abietina than trees planted on pine sites. Pinus contorta was more severely attacked by G. abietina than P. sylvestris on both site types. The results show that plantations of P. contorta and P. sylvestris in areas originally covered with pure spruce forests run a high risk of damage by G. abietina.     

Is insecticide spraying a viable and cost-efficient management practice to control pine processionary moth in Mediterranean woodlands?

Insect pests are a major threat to many forests worldwide, from boreal to tropical forest ecosystems. Some pests exhibit periodical outbreaks, after which their populations often crash as a result of natural biological control. To offset such outbreaks, several management techniques are used, including aerial spraying of insecticides. The question remains whether pest decline following an outbreak is the result of management practices or a natural consequence of the insect's population cycle. In this study, we assessed the performance of aerial spraying of insecticides on pine woodland stands to control pine processionary moth Thaumetopoea pityocampa (PPM) outbreaks in southern Spain. To achieve this, we compared the degree to which a forest stand recovers from defoliation from one year of severe damage by PPM to the following year (infestation index) in stands that were treated (i.e. subjected to aerial spraying) and untreated using a 4-years database from the Regional Environmental Council. The results revealed a significant similar recovery from infestation after a PPM outbreak of both sprayed and non sprayed pine woodland stands, for the four most representative pine species (black, Aleppo, maritime, and stone pine). It is concluded that insecticide spraying cannot be considered a prevention for outbreaks if it is applied once the outbreak explodes. Management practices that can help control PPM outbreaks include promoting spatial heterogeneity at the landscape level, fostering biodiversity in pine plantations, and reinforcing parasitoid insect and predatory bird populations that negatively affect the PPM. This study illustrates how simple sampling designs and statistical tests can be useful decision-making tools and can help improve the environmental viability and cost-efficiency of forest management practices.     

Seed dispersal by small rodents favors oak over pine regeneration in the pine-oak forests of the Qinling mountains,China

Abstract

Natural regeneration differences between oak and pine are common in the Qinling Mountains of central China. Thus, we examined whether this difference could be explained by seed dispersal. We investigated the seed dispersal of Quercus aliena var. acuteserrata Maxim. and Pinus tabulaeformis Carr. and the regeneration characteristics of both species were investigated in the pine-oak forest belt (oak stands, pine-oak stands, and pine stands) of Huoditang Forest in the Qinling Mountains in central China. We found that Q. aliena var. acuteserrata acorns were usually scatterhoarded, while seeds of P. tabulaeformis were generally consumed immediately. We found that 55% of the released acorns were transferred to the P. tabulaeformis stands. Moreover, Q. aliena var. acuteserrata regenerated successfully and was recruited in the three forest stands. However, the natural regeneration of P. tabulaeformis was very low in all stands because of high predation by small rodents. We propose, therefore, that the seed dispersal advantage of Q. aliena var. acuteserrata compared with P. tabulaeformis contributed greatly to the regeneration difference between these two species in the pine-oak forest belt of the Qinling Mountains in China.     

The effects of species mixture on the growth and yield of mid-rotation mixed stands of Scots pine and silver birch

The effect of tree species mixture on stand volume yield and on tree-species-specific diameter and height growth rates were analysed in managed mixed stands of Scots pine (Pinus sylvestris L.) and silver birch (Betula pendula Ehrn.).Data were obtained from 14 repeatedly measured stands located in Southern Finland on mineral soil sites with varying admixture of Scots pine and silver birch. Statistical analysis was carried out for studying the effect of species mixture on the development of stand characteristics. For the analysis, the plots were categorised into three groups (plot types) according to the species dominance. In order to analyse species-specific growth rates, individual-tree mixed linear growth models for tree diameter and height growth were developed for both tree species.The results clearly show that the yield of the managed mid-rotation, mixed stands was greater for stands dominated by Scots pine than for stands dominated by birch, and the stand volume increment decreased with an increasing proportion of silver birch. Analysis of diameter and height growth by tree species revealed that the main reason for this pattern is the negative impact of birch competition on the growth of pine trees. The increase in diameter of pine was clearly hampered if the proportion of birch was high. An abundance of birch also slightly decreased the growth in height of Scots pine, although the effect was less than on diameter growth. Species mixture did not affect the diameter growth of birch but did have a significant effect on height development. Height growth of birch was considerably greater in pine-dominated stands than in birch-dominated stands. In pine-dominated mixed stands, the height growth of birch was quite close to that of dominant pine trees, and birches can endure in competition with pines for light.The results apply for even-aged and single-storey managed stands, where stocking density and structure are controlled with pre-commercial and commercial thinnings. The results are not applicable to unmanaged mixed stands undergoing self-thinning. This study provides new information on mixed stands from a silvicultural perspective, which can be applied in decisions involving the management of mixed stands.     

Stand and landscape-level factors related to bird assemblages in exotic pine plantations: Implications for forest management

Plantations cover a substantial amount of Earth's terrestrial surface and this area is expected to increase dramatically in the coming decades. Pinus plantations make up approximately 32% of the global plantation estate. They are primarily managed for wood production, but have some capacity to support native fauna. This capacity likely varies with plantation management. We examined changes in the richness and frequency of occurrence of bird species at 32 plots within a Pinus radiata plantation (a management unit comprising multiple Pinus stands) in south-eastern Australia. Plots were stratified by distance to native forest, stand age class and thinning regime. We also assessed the landscape context of each plot to determine relationships between bird assemblages and stand and landscape-level factors. Bird species richness was significantly higher at plots ≥300 m from native forest and in mature (∼20 years since planting) and old (∼27 years since planting) thinned pine stands. We were able to separate the often confounding effects of stand age and thinning regime by including old stands that had never been thinned. These stands had significantly fewer species than thinned stands suggesting thinning regime, not age is a key factor to improving the capacity of pine plantations to support native species (although an age × thinning interaction may influence this result). At the landscape level, species richness increased in pine stands when they were closer to native riparian vegetation. There were no significant differences in species composition across plots. Our study indicates the importance of stand thinning and retention of native riparian vegetation in improving the value of pine plantations for the conservation of native fauna.