Regeneration dynamics in aging black pine (<Emphasis Type="Italic">Pinus nigra</Emphasis> Arn.) plantations on the south slopes of the Middle Balkan Range in Bulgaria

To study the regeneration dynamics in mature black pine (Pinus nigra Arn.) plantations in the Middle Balkan Range, Bulgaria, a total of 48 research plots (sized 0.1 ha) were established; 26 plots were established in the foothills of the mountains (foothill zone) and 22 in the main massive (mountainous zone). Pine overstorey as well as advance regeneration in subplots were recorded. Analysis of variance and regression analysis showed that in general relative stand basal area (RSBA) was the most important predictor of deciduous advance regeneration density, top height as well as age span of the regeneration layer. In the mountainous zone soil depth (SD) and the presence of mature deciduous trees (MDT) explained a significant amount of the variation in regeneration features. Lower RSBA induced by earlier thinnings resulted in higher regeneration density, larger top heights and wider age span of deciduous species. Under the pine canopy oak regeneration can persist longer compared to natural broadleaved canopies. In the studied plantations pine regeneration was confined to few scattered seedlings and regeneration groups. Based on these findings silvicultural concepts for the further management of black pine plantations in the Middle Balkan Range are discussed.     

Resistance of wood from black pine (<Emphasis Type="Italic">Pinus nigra</Emphasis> var. <Emphasis Type="Italic">austriaca</Emphasis>) against weathering

At present tapped wood of black pines (Pinus nigra var. austriaca) is mainly combusted. The priority of material use over thermal recycling has led to some considerations regarding the utilization of tapped wood. The high content of extractives suggests a higher natural durability, and therefore, the suitability for outdoor applications. Tapped and not tapped wood boards from black pine (sapwood and heartwood) were subjected to weathering tests to find out its resistance against abiotic stress. Additionally tapped wood particles with a high content of resin were exposed to weathering and to composting. Weathering caused roughness to increase. Infrared spectral characteristics revealed the differences before and after tapping and weathering. Principal component analyses supported the grouping according to the chemical changes. In heartwood the lignin band at 1510 cm^??1 disappears and the typical resin band at 1688 cm^??1 decreases considerably. The lignin band of resinous parts is affected neither by weathering nor by composting. However, the resin band shows an intensity decrease and broadening due to weathering and disappears during composting.     

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).     

Interactions between climate,growth and seed production in Spanish black pine (<Emphasis Type="Italic">Pinus nigra</Emphasis> Arn. ssp. <Emphasis Type="Italic">salzmannii</Emphasis>) forests in Cuenca Mountains (Spain)

Tree growth is regulated by a combination of exogenous and endogenous factors. Such factors also interact with each other, complicating the understanding of causal links. IN particular, resource allocation is sensitive to reproductive investment, especially in masting species, which in turn is regulated by climatic variables. Both resource allocation and seed production patterns are also sensitive to tree age. This study aims to (1) evaluate the effects of tree age and local and regional climate on tree ring width and seed production by Spanish black pine (Pinus nigra Arn. ssp. salzmannii) forest in Cuenca Mountains (Spain), and (2) assess the relationship between seed production and secondary growth of Spanish black pine. Seed fall was estimated using 60 rectangular seed traps (40 × 50 × 15 cm) from 2000 to 2014, randomly distributed across the study area. Standardized tree-ring chronologies were calculated using a random sample of 106 trees stratified into three age classes (> 80 years; 26–80 years, and ≤ 25 years). Local climate data was obtained from a meteorological station, and regional climate data from the CRU-TS 3.1 dataset. Average seed production ranged over time from 2 to 189 seeds m^?2 (coefficient of variation = 157%). We identified four masting years (2000, 2003, 2006, and 2014) using a classification based on percentile seed production. Seed production was regulated by climate of the previous 2–3 years, while tree growth responded to precipitation and temperature in the previous and current year. Independent of climate, high seed production had a negative effect on tree ring width and weakened climate growth relationships, indicating resource depletion. Tree age modulated climate sensitivity, increasing correlations between climate and tree-ring index in older trees. P. nigra has been showed to be a climate sensitive species with a bimodal masting behaviour, which should be taking into account for management purposes and silvicultural guidelines under climate change scenarios.     

Influence of region of provenance and climate factors on wood anatomical traits of <Emphasis Type="Italic">Pinus nigra</Emphasis> Arn. subsp. <Emphasis Type="Italic">salzmannii</Emphasis>

Region of provenance is defined as an area with uniform ecological conditions where stands with similar phenotypic or genetic features are found. This study assesses the effect of differing climate conditions of eight Spanish regions of provenance of Pinus nigra Arn. subsp. salzmannii on earlywood anatomical traits measured in samples from basal discs from mature trees. Results showed that variation in wood biometry between provenances was high and more pronounced than intrapopulation variation. When comparing P. nigra with other Mediterranean pines, high intertracheid wall strength values are associated with better adaptation of pines to arid conditions. However, the intraspecific variations of this parameter in P. nigra did not follow the same pattern, due to the influence of mechanical support requirements. Trees subject to greater aridity were characterised by short tracheids, apparently resulting from their poorer growth, and high frequency of rays and ray parenchyma cells, which would allow trees to store greater amounts of starch, which is the source of metabolites invested in minimising the limitations imposed by water stress. Severe winter cold spells were strongly associated with high axial resin canal frequency and large radial resin canals, creating a powerful, preformed defence system. Increased tracheid lumen involved an increase in the size of bordered pits, favouring sap flow between tracheids, in addition to an increase in the maximum diameter of cross-field pits, favouring the flow of water and metabolites between the axial and radial systems. The high influence of region of provenance on structural variation in P. nigra shows the importance of provenance in the selection of seed origin for reforestation.     

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).     

Species proportions by area in mixtures of Scots pine (<Emphasis Type="Italic">Pinus sylvestris</Emphasis> L.) and European beech (<Emphasis Type="Italic">Fagus sylvatica</Emphasis> L.)

Scots pine (Pinus sylvestris L.) and European beech (Fagus sylvatica L.) dominate many of the European forest stands. Also, mixtures of European beech and Scots pine more or less occur over all European countries, but have been scarcely investigated. The area occupied by each species is of high relevance, especially for growth evaluation and comparison of different species in mixed and monospecific stands. Thus, we studied different methods to describe species proportions and their definition as proportion by area. 25 triplets consisting of mixed and monospecific stands were established across Europe ranging from Lithuania to Spain in northern to southern direction and from Bulgaria to Belgium in eastern to western direction. On stand level, the conclusive method for estimating the species proportion as a fraction of the stand area relates the observed density (tree number or basal area) to its potential. This stand-level estimation makes use of the potential from comparable neighboring monospecific stands or from maximum density lines derived from other data, e.g. forest inventories or permanent observations plots. At tree level, the fraction of the stand area occupied by a species can be derived from the proportions of their crown projection area or of their leaf area. The estimates of the potentials obtained from neighboring monospecific stands, especially in older stands, were poorer than those from the maximum density line depending on the Martonne aridity index. Therefore, the stand-level method in combination with the Martonne aridity index for potential densities can be highly recommended. The species’ proportions estimated with this method are best approximated by the proportions of the species’ leaf areas. In forest practice, the most commonly applied method is an ocular estimation of the proportions by crown projection area. Even though the proportions of pine were calculated here by measuring crown projection areas in the field, we found this method to underestimate the proportion by 25% compared to the stand-level approach.     

Regional mixed-effects height–diameter models for loblolly pine (<Emphasis Type="Italic">Pinus taeda</Emphasis> L.) plantations

A height–diameter mixed-effects model was developed for loblolly pine (Pinus taeda L.) plantations in the southeastern US. Data were obtained from a region-wide thinning study established by the Loblolly Pine Growth and Yield Research Cooperative at Virginia Tech. The height–diameter model was based on an allometric function, which was linearized to include both fixed- and random-effects parameters. A test of regional-specific fixed-effects parameters indicated that separate equations were needed to estimate total tree heights in the Piedmont and Coastal Plain physiographic regions. The effect of sample size on the ability to estimate random-effects parameters in a new plot was analyzed. For both regions, an increase in the number of sample trees decreased the bias when the equation was applied to independent data. This investigation showed that the use of a calibrated response using one sample tree per plot makes the inclusion of additional predictor variables (e.g., stand density) unnecessary. A numerical example demonstrates the methodology used to predict random effects parameters, and thus, to estimate plot specific height–diameter relationships.     

Spatial pattern of beech (<Emphasis Type="Italic">Fagus sylvatica</Emphasis> L.) and oak (<Emphasis Type="Italic">Quercus pubescens</Emphasis> Mill.) seedlings in natural pine (<Emphasis Type="Italic">Pinus sylvestris</Emphasis> L.) woodlands

Spatial pattern of recruitment is an important factor influencing population dynamics of plant communities. The spatial pattern is determined by seed dispersal and by the spatial variability of germination and initial survival. In the process of forest expansion following farmland abandonment, mid- and late-successional species are often dispersed in pioneer forests by birds. This could result in an aggregated spatial pattern of seeds that could influence the dynamics of these species in mixed pioneer forests. In the sub-Mediterranean area, mid- and late-successional species such as Quercus pubescens (downy oak) and Fagus sylvatica (European beech) are expected to replace pioneer Pinus species. Using a point sampling method we demonstrated that beech and oak seedlings (height <2 m) have a clumped distribution in the understorey of pine. This could result from an aggregated dispersal by jays (dispersal effect) or from preferential recruitment in particular habitats (habitat effect). To test these hypotheses we proposed a statistical analysis of spatial patterns of regeneration of beech and oak. Ground cover variables (i.e. cover by rock outcrops, herbs, box shrubs, mosses or pine) did not differ significantly around seedlings as compared with random sample plots. Likewise, clumped seedlings had growth similar to isolated seedlings, thus refuting the hypothesis of preferential location in the most favourable microsites. Aggregated dispersal seems to be involved in the process of regeneration. Since beech and oak seedlings have contrasting ecological demands, we discuss the implication of this pattern for the replacement dynamics of pine by these species.     

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.     

<Emphasis Type="Italic">Quercus suber</Emphasis> forest and <Emphasis Type="Italic">Pinus</Emphasis> plantations show different post-fire resilience in Mediterranean north-western Africa

Key message

In the African rim of the Western Mediterranean Basin, cork oak forests and pine plantations coexist. Under similar fire regimes, cork oak forest is more resilient in terms of habitat structure (canopy, understory, and complexity of vegetation strata) than pine plantation. By contrast, both woodland types show similar resilience in plant species composition. Resilience in habitat structure varies between the two woodland types because of the resprouting and seeding strategies of cork oak and pine species, respectively. These differences can be relevant for the conservation of biodiversity of forested ecosystems in a future scenario of increased fire frequency and scale in the Mediterranean basin.

Context

Wildfires have major impacts on ecosystems globally. In fire-prone regions, plant species have developed adaptive traits (resprouting and seeding) to survive and persist due to long evolutionary coexistence with fire. In the African rim of the Western Mediterranean Basin, cork oak forest and pine plantation are the most frequently burnt woodlands. Both species have different strategies to respond fire: cork oak is a resprouter while pines are mostly seeders.

Aims

We have examined the hypothesis that pine plantations are less resilient in habitat structure (canopy, understory, diversity of vegetation strata) and plant composition than cork oak woodlands.

Methods

The habitat structure and plant species composition were measured in 30 burnt and 30 unburnt 700-m transects at 12 burnt sites from north-western Africa, where the two forest types can coexist. Habitat structure and plant species composition were compared between burnt and unburnt transects from cork oak and pine plantation woodlands with generalized linear mixed models and general linear models.

Results

The results showed significant interaction effect of fire and forest type, since cork oak forest was more resilient to fire than was pine plantation in habitat structure. By contrast, both forest types were resilient to fire in the composition of the plant communities, i.e., plant composition prior to fire did not change afterwards.

Conclusion

The higher structural resilience of cork oak forest compared to pine plantation is related to the resprouting and seeding strategies, respectively, of the dominant tree species. Differences in the responses to fire need to be considered in conservation planning for the maintenance of the Mediterranean biodiversity in a future scenario of changes in fire regime.

     

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.

     

Factors determining enzyme activities in soils under <Emphasis Type="Italic">Pinus halepensis</Emphasis> and <Emphasis Type="Italic">Pinus sylvestris</Emphasis> plantations in Spain: a basis for establishing sustainable forest management strategies

Key message

Water availability and soil pH seem to be major constraints for enzyme activities in calcareous soils under Pinus halepensis and acidic soils under Pinus sylvestris plantations respectively. Proposals for improving enzyme activities may include the promotion of broadleaf species to increase soil pH and the modulation of stand density or the implementation of soil preparation techniques to facilitate water infiltration.

Context

Soil enzymes play a key role in nutrient turnover in forest ecosystems, as they are responsible for the transformation of organic matter into available nutrients for plants. Enzyme activities are commonly influenced by temperature, humidity, nutrient availability, pH, and organic matter content.

Aims

To assess the differences between enzyme activities in calcareous soils below Pinus halepensis and acidic soils below Pinus sylvestris plantations in Spain and to trace those differences back to edapho-climatic parameters to answer the questions: Which environmental factors drive enzyme activities in these soils? How can forest management improve them?

Methods

The differences in climatic, soil physical, chemical, and biochemical parameters and the correlations between these parameters and enzyme activities in soils were assessed.

Results

Low pH and high level of phenols in acidic soils under Pinus sylvestris and water deficit in calcareous soils under Pinus halepensis plantations appeared to be the most limiting factors for enzyme activities.

Conclusion

Options such as the promotion of native broadleaf species in the Pinus sylvestris stands and the modulation of Pinus halepensis stand density or the implementation of soil preparation techniques may improve enzyme activities and, therefore, nutrient availability.

     

Vermicompost enhances germination of the maritime pine (<Emphasis Type="Italic">Pinus pinaster</Emphasis> Ait.)

We investigated the effect of vermicompost on the germination and early development of six different progenies of the maritime pine (Pinus pinaster Ait.). We compared the effects of incorporating solid vermicompost into the potting media to those of vermicompost water extract. The incorporation of vermicompost in the growing media of maritime pine increased germination by 16%, and particularly, addition of vermicompost water extract produced the best results. Plants germinated with vermicompost showed higher N content as compared to control plants, and this could have determined the faster maturation of the treated seedlings. Since the best effects on pine germination were observed after application of vermicompost water extract, other mechanisms, rather than the physical amelioration of the substrate, such as the presence of water soluble nutrients and organic compounds (i.e. humic acids and plant growth regulating substances) in the vermicompost, might be involved in the promotion of germination. We also report variation in the response of the different pine progenies to vermicompost application thereby confirming the necessity of taking into account genetic variability in order to study the potential of vermicompost and other biologically-active organic materials as a potting amendments.     

Insecticidal properties of <Emphasis Type="Italic">Thymus persicus</Emphasis> essential oil against <Emphasis Type="Italic">Tribolium castaneum</Emphasis> and <Emphasis Type="Italic">Sitophilus oryzae</Emphasis>

Red flour beetle Tribolium castaneum (Herbst) and rice weevil Sitophilus oryzae (L.) are considered to be the major insect pests in storage. Essential oils from aromatic plants are recognized as proper alternatives to fumigants. Thymus persicus (Ronniger ex Rech. f.) is one of these plants that have medicinal properties and is indigenous to Iran. The essential oil was obtained from aerial parts of the plant and analyzed by GC and GC–MS. Carvacrol (44.69%) and thymol (11.05%) were the major constituents of the oil extracted. In this experiment, fumigant toxicity of the essential oil was studied against T. castaneum, S. oryzae at 27 ± 1°C and 60 ± 5% RH in dark condition. The adult insects were exposed to the concentrations of 51.9, 111.1, 207.4 and 370.4 μl/l air to estimate median lethal time (LT₅₀) values. The fumigant toxicity was increased in response to increased essential oil concentrations. The LT₅₀ values at the lowest and the highest concentrations tested were ranged from 28.09 to 13.47 h for T. castaneum, and 3.86 to 2.30 h for S. oryzae. It was found that S. oryzae adults were much more susceptible to the oil than T. castaneum. After 24 h of exposure, the LC₅₀ values (95% fiducial limit) for T. castaneum and S. oryzae were estimated to be 236.9 (186.27–292.81) and 3.34 (2.62–4.28) μl/l air, respectively. These results suggest that T. persicus essential oil merits further study as potential fumigant for the management of these stored-product insects.     

Magnitude and efficiency of genetic diversity captured from seed stands of <Emphasis Type="Italic">Pinus nigra</Emphasis> (Arnold) subsp. <Emphasis Type="Italic">pallasiana</Emphasis> in established seed orchards and plantations

To test how efficiently plantations and seed orchards captured genetic diversity from natural Anatolian black pine (Pinus nigra Arnold subspecies pallasiana Holmboe) seed stands, seed sources were chosen from 3 different categories (seed stands (SS), seed orchards (SO) and plantations (P)) comprising 4 different breeding zones of the species in Turkey. Twenty-five trees (mother trees) were selected from each SS, SO and P seed sources and were screened with 11 Random Amplified Polymorphic DNA (RAPD) markers. Estimated genetic diversity parameters were found to be generally high in all Anatolian black pine seed sources and the majority of genetic diversity is contained within seed sources (94%). No significant difference in genetic diversity parameters (numbers of effective alleles, % of polymorphic loci and heterozygosity) among seed source categories was found, except for a slight increase in observed heterozygosities in seed orchards. For all seed source categories, observed heterozygosity values were higher (H_o = 0.49 for SS, 0.55 for SO and 0.49 for P) than expected ones (H_e = 0.40 for SS, 0.39 for SO and 0.38 for P) indicating the excess of heterozygotes. In general, genetic diversity in seed stands has been transferred successfully into seed orchards and plantations. However, the use of seeds from seed orchards can increase the amount of genetic diversity in plantations further. The study also demonstrated that number of plus-tree clones (25–38) used in the establishment of seed orchards was adequate to capture the high level of diversity from natural stands.     

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.     

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.