A weeding-duration model for <Emphasis Type="Italic">Larix kaempferi</Emphasis> plantations in Hokkaido,northern Japan

In this study, a weeding-duration model for Larix kaempferi plantations was developed that employs a generalized linear model. The number of years that weeding is necessary is the response variable, and elevation, slope, maximum snow depth, annual precipitation, geological type, soil type, site index, slope aspect, and vegetation type are explanatory variables. Among the explanatory variables, geological type, soil type, slope aspect, and vegetation type are categorical data. We assumed a Poisson distribution for the response variable. The link function was log. Among the models that could be developed from these variables, we chose the model with the smallest Akaike’s information criterion (AIC). The weeding-duration model can be written as follows: years that weeding is necessary = Exp (−0.0172833 × site index + 0.0014053 × maximum snow depth (cm) + 1.7417731). The results of this study imply that weeding of Larix kaempferi plantations is needed for more years as the maximum snow depth increases and fewer years as the site index increases. This model is useful for cost–benefit analyses of afforestation or reforestation with Larix kaempferi.     

Migration of pine wood nematodes in the tissues of <Emphasis Type="Italic">Pinus thunbergii</Emphasis>

The distribution of pine wood nematodes (Bursaphelenchus xylophilus, PWNs) in Japanese black pine (Pinus thunbergii) tissues was investigated by staining with fluorescein isothiocyanate-conjugated wheat germ agglutinin. After PWNs were inoculated to current-year stems of pine seedlings, their distribution at about 5 cm below the inoculation site was confined only to cortical resin canals 1 day after inoculation, and then spread to other tissues, including resin canals of short branches. When PWNs were inoculated onto cross or tangentially cut surfaces of stem segments, maximal PWN migration speed was estimated to be faster through cortical resin canals and xylem axial resin canals vertically (>6.7 and <2.3 mm/h, respectively) than through cortical tissues both vertically and horizontally (<1.2 and <0.2 mm/h). To examine whether PWNs in cortical resin canals could invade surrounding tissues, segments in which PWNs resided only in cortical resin canals were prepared by removing the top portion 6 h after inoculation. Additional incubation of such segments caused extended PWN distribution to xylem axial resin canals and then to other tissues. A similar experiment with top portions of girdled segments removed 12 h after inoculation also showed extended PWN distribution from xylem axial resin canals and pith to cortical resin canals and then to other tissues. These results provided direct evidence that PWNs have the ability to migrate from cortical resin canals and xylem axial resin canals to other tissues.     

An inoculation experiment of Japanese <Emphasis Type="Italic">Bursaphelenchus</Emphasis> nematodes on Japanese black and red pine, <Emphasis Type="Italic">Pinus thunbergii</Emphasis> and <Emphasis Type="Italic">P. densiflora</Emphasis>

The pinewood nematode (PWN) Bursaphelenchus xylophilus is an invasive pathogen that was introduced from North America to Asian countries and Portugal and is devastating native pine forests. Some native European and Asian Bursaphelenchus nematodes also have weak to moderate pathogenicity to native pine species. To evaluate the potential risk of native Bursaphelenchus species, we inoculated ten Japanese Bursaphelenchus species into native pine species (the dominant forest species) in Japan, and evaluated their pathogenicity using mortality and tracheal tissue damage as indices. Inoculation was conducted on August 3, 2007, and the symptoms were observed every 2 weeks until February 1, 2008. None of the inoculated trees, excluding the pathogenic PWN inoculated control, showed external disease symptoms; however, four species [a less pathogenic PWN isolate, B. luxuriosae, Bursaphelenchus sp. NK215 (undescribed), and NK224 (undescribed)] caused tracheal tissue damage in inoculated seedlings and showed weak pathogenicity. Therefore, we conclude that there are some potentially pathogenic native species of nematodes distributed in Japan. Interestingly, two of these weakly pathogenic species, B. luxuriosae and NK215, are not associated with Pinaceae trees, suggesting that nematode pathogenicity may be a pre-adaptive character. More experimental studies under different conditions are necessary to accurately evaluate the potential risk of these pathogens.     

Regeneration dynamics of mixed stands of <Emphasis Type="Italic">Pinus pinaster</Emphasis> Ait. and <Emphasis Type="Italic">Pinus pinea</Emphasis> L. in Central Spain

The dynamics of mixed stands are more complex and less studied than those of monospecific stands. The objective of this work was to analyze the variables involved in seedling occurrence and seedling survival in mixed stands of Pinus pinaster and P. pinea in Mediterranean areas. From 2011 to 2016, regeneration of both species was monitored at two sites located in Central Spain. We installed 72 regeneration plots where seedling dynamics were monitored. All the trees in the study areas were measured and mapped. Additionally, we took hemispherical photographs in each regeneration plot. The average density of P. pinea seedlings over the study period was almost 20 times larger than that of P. pinaster. Our results indicate that the seedlings of both species grow under moderate light conditions. In addition, we found that the occurrence of seedlings of both species was related to the structure of the stand. P. pinea seedlings grew where the density and size of P. pinaster trees were low and where P. pinea trees provided moderately sheltered conditions, whereas the number of P. pinaster seedlings was related to under intermediate densities of P. pinaster trees. Furthermore, seedling survival was positively associated with age of the seedlings and negatively with the August average maximum temperature. The temporal continuity of mixed stands of P. pinea and P. pinaster in the study area is compromised by the observed lack of regeneration of P. pinaster.     

<Emphasis Type="Italic">Bursaphelenchus mucronatus</Emphasis> (Nematoda: Aphelenchoididae) vectored by <Emphasis Type="Italic">Monochamus </Emphasis><Emphasis Type="Italic">urussovi</Emphasis> (Coleoptera: Cerambycidae) in Hokkaido,Japan

Bursaphelenchus mucronatus Mamiya et Enda has been recovered for the first time from adults of the cerambycid beetle, Monochamus urussovi (Fischer), in Hokkaido, Japan. The nematode was also recovered from the inner bark of Picea jezoensis (Siebold et Zuccarini) Carrière and Abies sachalinensis (Fr Schmidt) Masters infested with M. urussovi larvae. PCR–RFLP analysis indicated that B. mucronatus in Hokkaido is the European type.     

Dynamics of ecosystem services in <Emphasis Type="Italic">Pinus sylvestris</Emphasis> stands under different managements and site quality classes

Timber and biodiversity are considered two antagonistic ecosystem services (ES), largely influenced by silviculture and site ecological conditions. In order to address the trade-offs between these two ES over time, we performed a retrospective study at compartment level in the Pinus sylvestris forests of the Spanish Central Mountain System. Archival data from Management Plans for eight forests with contrasting silvicultural systems (uniform shelterwood system, group shelterwood system, irregular shelterwood system,) and three different site quality classes were analysed. Timber production was assessed through stand volume, harvested timber volume and a stand volume index. Biodiversity was examined through structural diversity (Gini index, Shannon tree size diversity index, Simpson’s reciprocal index and evenness index, all applied to diameter classes) and the abundance of large living trees. For all silvicultural systems investigated, stand volume and harvested timber volume have grown since the beginning of the management plans (beginning of the twentieth century in some forests). The largest yields of timber corresponded to the best quality sites with more intensive silvicultural treatments (uniform and group shelterwood systems). The uniform shelterwood system showed lower figures for structural diversity, though not always significant. The best site qualities maintained notable structural diversity values, even under the most intensive management system. The application of the different management systems over decades has revealed a synergy between timber production and structural diversity, particularly in those systems maintaining more than one age class, although results are conditioned by forest harvesting history. The interaction between historical silvicultural treatment and site quality has been identified as an important source of information to understand forest dynamics and functioning of ES provision.     

Time-trajectory of mean component weight and density in self-thinning <Emphasis Type="Italic">Pinus densiflora</Emphasis> stands

The allometric relationships between mean weights of components, such as stems, branches and leaves and tree weight as well as their time-trajectories, were studied with data of self-thinning Pinus densiflora stands with different densities. The allometric relationships existed between the weights of stems, branches and leaves and the tree weight during the course of self-thinning. The stem weight ratio increased with increasing tree weight because the allometric coefficient in stem was higher than unity, whereas the branch weight ratio and the leaf weight ratio decreased because the allometric coefficients in branches and leaves were less than unity. An allometric power relationship existed between mean component weight and mean tree weight during the course of self-thinning. The time-trajectory of mean component weight (w _o) and density (ρ) in the early growth stage was expressed as a mathematical model which incorporates the allometric power relationship into the Tadaki’s model, whereas the model for describing w _o-ρ trajectory in the later growth stage was derived by combining the allometric power relationship with 3/2 power law. The two models, Tadaki’s model and 3/2 power law, showed a good fit to data from P. densiflora stands. The time-trajectories of mean tree weight (w)-density (ρ) or w _o-ρ initially almost moves nearly vertically in the low-density stand, moves along a steep curve and an inclined curve in the medium- and high-density stands, respectively, and gradually approaches self-thinning line in the early stage of stand development, whereas they reached and moved along the self-thinning line in the later stage of stand development. The self-thinning exponents were determined to be 1.71, 1.19 and 1.13 for the trees, 2.38, 1.33 and 1.20 for the stem, 3.16, 1.55 and 1.46 for the branches, 2.66, 1.39 and 1.35 for the leaves in the low-, medium- and high-density stands, respectively. The 3/2 power law of self-thinning is derived on the basis of simple geometric model of space occupation by growing trees, but allometric growth of tree and components can make the slope of the self-thinning line being different from −3/2. The reasons that the self-thinning exponents of components in the low-density stand were greater than those in the medium- and high-density stands were discussed.     

Competition as a predisposing factor of crown dieback in a secondary forest of <Emphasis Type="Italic">Betula maximowicziana</Emphasis> in Hokkaido,northern Japan

We analyzed the probability that Betula maximowicziana Regel (monarch birch) would suffer crown dieback (crown-dieback probability) and the basal area growth rate (GB), which was found to be a predisposing stress factor making birch trees susceptible to crown dieback. First, we analyzed the relationship between the probability that birch trees would suffer from crown dieback in 1999 and GB from a period prior to the occurrence of crown dieback (1985–1987), using a data set of repeated measurements on 217 trees. Logistic regression analysis revealed that monarch birch had a larger crown-dieback probability when GB was low in the preceding period. Hence, there were predisposing stress factors that reduced GB and continued to affect trees for at least a decade. Next, we analyzed GB in the same period in relation to symmetrical and asymmetrical competition between trees and found that GB was reduced by symmetrical competition, suggesting that this was one of the predisposing factors for crown dieback. Based on these results, we used selected models for crown-dieback probability and GB to calculate crown-dieback probabilities for individuals with different initial basal areas and experiencing different intensities of symmetrical competition. The predicted crown-dieback probability decreased with decreasing symmetrical competition between trees. We discuss a possible process of crown dieback to death for monarch birch and the use of thinning as a method to reduce the risk of crown dieback.     

Mortality of planted <Emphasis Type="Italic">Pinus thunbergii</Emphasis> Parlat. saplings subject to coldness during winter and soil types in region of seasonal soil frost

To assess and improve methods for the development of coastal forests in Hokkaido, northern Japan, this study examined the factors affecting mortality of planted Pinus thunbergii Parlat. saplings in a region with seasonal soil frost. The needles of pine saplings at the study site turn red in late spring, suggesting xylem embolism, which can lead to death. Sapling mortality was strongly correlated with the degree of cold just after planting and with the occurrence of repeated severe winters. Volcanic ash soil had been supplied to improve the soil nutrients, and the volcanic ash soil always thawed later than the original coastal sand. Saplings in volcanic ash were significantly more damaged than those in original coastal sand. This suggests that hydraulic stress caused by cuticular transpiration under frozen soil conditions during spring may also accelerate damage due to embolisms. Hence, volcanic ash soil should not be applied to soils in regions with seasonal soil frost. A covering of woodchips can help prevent frost from reaching greater soil depths, and, indeed, fewer saplings at the study site died in areas with such coverage. Planting location can also affect pine sapling mortality. For example, shading caused by wood fencing may affect the soil freezing profile during winter; therefore, planting close to a fence should also be avoided in regions with seasonal soil frost. These findings should be used to improve the development of forests in regions with seasonal soil frost.     

Comparison of postfire mortality in endemic Corsican black pine (<Emphasis Type="Italic">Pinus nigra</Emphasis> ssp. <Emphasis Type="Italic">laricio</Emphasis>) and its direct competitor (<Emphasis Type="Italic">Pinus pinaster</Emphasis>)

• Introduction   

Laricio pine (Pinus nigra J.F. Arn. ssp. laricio (Poiret) Maire var Corsicana Hyl.) is a form of black pine endemic to Corsica, that may now be under threat due to current fire regimes and competition with maritime pine (Pinus pinaster Aiton).     

Seasonal nutrient retranslocation in reforested <Emphasis Type="Italic">Pinus halepensis</Emphasis> Mill. stands in Southeast Spain

Retranslocation, resorption and relocation of nutrients are important adaptive mechanisms developed by plants to acquire the amount of the nutrients required for growth. They are usual mechanisms in deciduous and conifer trees that occur in Mediterranean regions where drought periods are usual. Soil factors, environmental characteristics and species factors are key drivers of nutrient retranslocation in conifers but is not well understood how soil fertility or intraspecific competition influences the process. We studied retranslocation in Pinus halepensis Mill. stands showing different site quality (differences in climate and intraspecific competition) occurring in Southeast Spain. We monitored reforested mature Aleppo pine forests in stands with differences in site quality, climate and intraspecific competition. Stands were characterised, the content of nutrients of soil and green samples (twigs and pine needles) were recorded, and seasonal nutrient retranslocation was obtained. Site characteristics were related to growth rate and nutrient content of foliage and soil. We evaluated whether the retranslocation of nutrients from older to younger foliage was related to the current-year growth rate and to the nutritional status of the plant as influenced by intraspecific competition. Foliar macronutrient concentrations and the amount of retranslocated macronutrients were seasonal, with differences related to site quality and tree density. As a general trend, nutrient concentrations increased after drought (autumn) and decreased during the growth period (spring). However, some micronutrients (mainly Na and Fe) decreased during both periods. The retranslocation pattern in Aleppo pine reinforced the hypothesis that pine adaptations to drought- and fire-prone habitats are linked to the resilience of these forest types. We developed scientific knowledge to assist decision making in adaptive forest management; e.g. fertilizer recommendations or reforestation programmes.     

Aboveground biomass equations for individual trees of <Emphasis Type="Italic">Cryptomeria japonica</Emphasis>, <Emphasis Type="Italic">Chamaecyparis obtusa</Emphasis> and <Emphasis Type="Italic">Larix kaempferi</Emphasis> in Japan

Generic equations are proposed for stem, branch and foliage biomass of individual trees in even-aged pure stands of Cryptomeria japonica, Chamaecyparis obtusa and Larix kaempferi. Biomass data was collected from a total of 1,016 individual trees from 247 stands throughout Japan, and five regression models were assessed by root mean square error, mean bias, fit index (FI), and AIC. The results show that a power equation using diameter at breast height (dbh) and height is the most suitable for all species and components. This equation is more accurate than the familiar power equation that uses ‘dbh² height’, and it expresses the greater volume of branch and foliage mass of trees with a lower height/diameter ratio. A power equation using dbh is more reasonable for models with dbh as the only independent variable and more accurate than a power equation using ‘dbh² height’ for estimating branch and foliage mass. Estimating error for branch and foliage mass is larger than that for stem mass, but the entire aboveground biomass can be estimated with an error of less than 19%, except in the case of small trees with dbh less than 10 cm.     

A growth and yield model for even-aged <Emphasis Type="Italic">Pinus brutia</Emphasis> Ten. stands in Syria

Pinus brutia Ten. subspecies brutia, which occurs in the Eastern Mediterranean region, is the main forest species in Syria and important for multi-purpose forestry. In this study, 6,631 10-year past growth diameter increment measurements were taken in 83 temporary sample plots. The current breast height diameter of all trees was measured and a sample of trees was measured for height and age. The plots were placed so as to capture the whole range of variation in site quality, stand age and stand density. The data were used to develop the following models:

Recessive inheritance of morphological characteristics of utsukushimatsu,<Emphasis Type="Italic">Pinus densiflora</Emphasis> f.<Emphasis Type="Italic">umbraculifera</Emphasis>

Pinus densiflora f.umbraculifera, commonly known as utsukushimatsu, is a distinctively shaped form of Japanese red pine whose growth is restricted to a forest stand in Shiga Prefecture, Japan. The inheritance mode of morphological characteristics of utsukushimatsu was studied to preserve the genetic resource of this pine. As previously reported, F₁ trees grown from open-pollinated seeds harvested from trees inhabiting the native stand showed two phenotypes: one resembling utsukushimatsu, which produces multiple trunks, and the other resembling normalP. densiflora, which produces one or a few trunks. In the present study, controlled pollination was carried out using F₁ and normalP. densiflora trees. Segregation ratios of the two phenotypes observed in the F₂ population showed that the morphological characteristics of utsukushimatsu are inherited recessively. This suggests that the mutation of one gene or a few closely linked genes controls the morphological characteristics of utsukushimatsu. Since multiple trunk formation of utsukushimatsu might be related to a loss of lateral bud inhibition, it follows that a simple gene mutation breaks apical dominance inP. densiflora.     

Adaptive management rules for <Emphasis Type="Italic">Pinus nigra</Emphasis> Arnold ssp<Emphasis Type="Italic">. salzmannii</Emphasis> stands under risk of fire

Key message

We generate flexible management rules for black pine stands, adaptable to alternative stand management situations and entailing thinnings, final-felling, and salvage cuts, based on the results on 270 stand level optimizations.

Context

Forest management instructions often rely on the anticipated prediction of the stand development, which poses a challenge on variable economic and environmental conditions. Instead, an alternative approach to better adapt forest management decisions to changing conditions is defining flexible rules based on thresholds that trigger management operations.

Aims

This article develops rules for the adaptive management of P. nigra stands in Catalonia (Spain) addressing the risk of fire and post-fire forest management.

Methods

The stochastic version of the simulation-optimization system RODAL was used to optimize the management of forest stands in three sites under different fire probability levels. A total of 270 optimizations were done varying site fertility, fire probability, and economic factors. The results of the optimizations were used as the basis of flexible forest management rules for adaptive stand management.

Results

The developed management rules defined the basal area limit for thinning, the thinning intensity, the mean tree diameter at which regeneration cuttings should start, and the basal area below which a salvage cutting should be done. Fire risk was not a significant predictor of the models for thinning and final cutting rules.

Conclusion

The presented rules provide a flexible tool for forest management during the stand development and under changing conditions when the management objective is to maximize economic profitability of timber production.

     

Successful <Emphasis Type="Italic">Agrobacterium</Emphasis>-mediated transformation of <Emphasis Type="Italic">Populus tomentosa</Emphasis> with apple <Emphasis Type="Italic">SPDS</Emphasis> gene

The problem of salinized soils has become one of the most serious constraints to agricultural and forest productivity. With the purpose of enhancing salt stress tolerance of Populus tomentosa, we transformed this tree species with spermidine synthase （SPDS） genes derived from an apple by an Agrobacterium-mediated method. Four transgenic clones were confu＇med by PCR and Southern blot analysis. As well, the expression of introduced SPDS genes was analyzed by real-time quantitative PCR.     

Estimation of stand attributes in <Emphasis Type="Italic">Cryptomeria japonica</Emphasis> and <Emphasis Type="Italic">Chamaecyparis obtusa</Emphasis> stands using QuickBird panchromatic data

This study aims to estimate stand density and stand volume in Cryptomeria japonica and Chamaecyparis obtusa stands from high-resolution satellite data and verify the reliability and uncertainty of the data. Sixty circular sample plots of 0.04 ha each were established. Their stand densities were estimated from the number of tree crowns derived from high-resolution satellite data using the watershed method. Stand densities derived from field surveys in the sample plots were compared with those obtained from high-resolution satellite data by stand age class. As a result, there was a positive correlation between them for sample plots of 41 years of age and over (R = 0.82); however, there was no correlation between them for sample plots of 40 years and under. Individual diameters at breast height (DBH) were estimated from crown areas obtained from high-resolution satellite data for the two species. Using the estimated DBH, individual tree heights were predicted from the height–diameter curves. Stand volumes were estimated from the sum of individual volumes, which were derived from volume formulas having two variables, i.e., DBH and height. Stand volumes derived from the field survey were compared with those obtained from high-resolution satellite data. The correlation coefficient between them for stands of 41 years of age and over was 0.78.     

Developing conservation strategies for <Emphasis Type="Italic">Pinus koraiensis</Emphasis> and <Emphasis Type="Italic">Eleutherococcus senticosus</Emphasis> by using model-based geographic distributions

We described potential changes in the geographic distribution and occurrence probability of Pinus koraiensis Sieb. et Zucc. and Eleutherococcus senticosus (Rupr. et Maxim.) Maxim. in the counties of northeast China. This information was used to identify priority areas for protection and provide protection and management recommendations within each studied county. The two species were mapped in 2884 study plots throughout this region over a 4-year period (38°40′N–53°30′N, 115°05′E–135°02′E). We used the species distribution models (Maxent), systematic conservation planning models (Marxan), and Geographic Information Systems (ArcGIS 10.0). The distributions of two species were correlated in the study area, enabling unique and economically viable joint conservation measures to be implemented. Three models were combined to identify feasible priority conservation sites. We used local spatial statistics to assess all identified conservation areas in relation to potential climate change based shifts in the geographic distribution of the two species. Model-based conservation strategies were used to identify effective measures to protect and utilize these two tree species in the study region. This study presents a novel technique for assessing wild plant distributions, in addition to serving as a model for the conservation of other species within the framework of general forest management, ecological construction, and geographical surveying.     

Fine root biomass and morphology of <Emphasis Type="Italic">Pinus densiflora</Emphasis> under competitive stress by <Emphasis Type="Italic">Chamaecyparis obtusa</Emphasis>

The fine root (diameter ≤2.0 mm) biomass and morphology of Japanese red pines (Pinus densiflora) grown under different aboveground conditions (i.e., high and low competitive environments) were examined in a pine–cypress mixed forest. All P. densiflora subject trees were about 40 years old, and the aboveground condition (i.e., size) of red pines appeared to be influenced by the surrounding Japanese cypress (Chamaecyparis obtusa). Smaller P. densiflora exhibited lower fine root biomasses, shorter root lengths, and lower root tip densities, but longer specific root lengths and higher specific root tip densities relative to larger pines. These results suggest that P. densiflora may adjust the morphological traits of fine roots to the different conditions in biomass allocation to fine roots of individuals with different aboveground growth.     

Fertility variation in endemic populations of Ehrami black pine (<Emphasis Type="Italic">Pinus nigra</Emphasis> Arnold. subsp. <Emphasis Type="Italic">pallasiana</Emphasis> var. <Emphasis Type="Italic">pyramidata</Emphasis>)

Fertility variation and effective number of parents were estimated in the 100 individuals from each of three plantation populations (P1, P2 and P3) of the endangered Pinus nigra. subsp. pallasiana var. pyramidata established in the Isparta–Golcuk district in southern Turkey in 1990. Potential relations between cone yield and growth characters (height, diameter at breast height, diameter at base and crown diameter) were also investigated. Large differences were found among populations and within populations for cone production and growth characters. Mean cone production was eight (ranging from 6.5 in P2 to 9.1 in P1) for the three populations. The 10 most productive individuals for cones produced 32% of the total cones in P1, 39% in P2 and 34% in P3; 21, 16 and 6 individuals, respectively did not produce any cones in the populations. According to the correlation analyses, the relation between cone yield and growth characters changed for the populations, while height, diameter at breast height and crown diameter were positively significantly correlated (r = 0.182, 0.135 and 0.209; p ≤ 0.05) with cone production for the three populations. Fertility variation was 1.81 in total of three populations, ranging from 1.72 in P3 to 3.45 in P1). Effective number of parents was 142 (55.2 of number of parents censused; 29%, P1; 49.8%, P2; 58.3%, P3).