Testing prescribed fire as a tool to promote Scots pine <Emphasis Type="Italic">Pinus sylvestris</Emphasis> regeneration

Techniques for encouraging natural tree regeneration are of increasing interest to managers of Scots pine Pinus sylvestris forests. We tested prescribed fire at a management scale, with deer present or excluded, as a means of increasing rates of Scots pine forest expansion on heathland. At a semi-natural pinewood in Scotland, ten experimental blocks were established, within range of pine seed-fall. Each block comprised four, 100 m² plots. Two plots at each site were burnt and two fenced, allowing the effects of burning on pine regeneration to be measured at two levels of deer abundance. We monitored pine seedlings, seed-fall, deer dung and vegetation for 5 years following treatment. Differences in seedling detection rates between treatments were quantified using dummy seedlings, and analyses corrected accordingly. Mean new pine seedling establishment was 9.8 times higher on burnt ground than unburnt ground (confidence intervals 3.2–30). Differences were even more pronounced in a year of high seed-fall, and following fires with low rates of spread. Establishment rates varied strongly between experimental blocks. Exclusion of deer increased establishment rates, but only in the first 2 years after fire. There was evidence that both seedling survival, and cumulative recruitment of older (over 12 months) seedlings, were improved by prescribed burning. Our results support the use of prescribed fire as a tool to promote increased Scots pine seedling establishment. This technique is likely to give strong fine-scale variation in seedling densities, and so would most suit areas where a variable spatial pattern of regeneration is sought, for landscape or naturalness reasons. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

The colonisation of Scots pine (<Emphasis Type="Italic">Pinus sylvestris</Emphasis> L.) by <Emphasis Type="Italic">Tomicus minor</Emphasis> Hartig in southern Poland: modelling and monitoring

Tomicus minor Hartig (Col., Curculionidae, Scolytinae), occurring on Pinus sylvestris L., is a species which demonstrates high reproductive capability on weakened stands, accelerating the process of forest death. In protected areas, T. minor is regarded as a sensitive bioindicator that reacts to decline in the health and vitality of forests. Although there have been many publications concerning T. minor, no precise method has yet been given for estimating its population so as to enable the monitoring of forest vitality and assessment of the role played by T. minor in the forest ecosystem. The aim of the present work is to develop a statistical method for estimating populations of T. minor, requiring minimum work and interference with the forest ecosystem and permitting the computation of estimation errors. Research was carried out in the years 1992–2011 in pine stands aged over 80 years, growing in a variety of habitats and situated at varying distances from sawmill timber yards. Attack density of T. minor was measured on trap logs made from uninfested living trees. The population of T. minor on the trap logs was described using a multiple linear regression model with two explanatory variables. Among the features investigated, the T. minor population was found to depend significantly on the number of egg galleries on the fifth metre of the trap log counted from the thinner end (p < 0.001) and on the diameter of the trap log in bark at the thinner end (p < 0.05). The model explains approximately 85% (R ² = 0.8564) of the variation in the total number of T. minor egg galleries on the trap logs. The numbering of units beginning from the thinner end of the log enabled increased precision in determining the model parameter resulting from the concentration of egg galleries on certain units of the log. In all validated plots, the mean real and model values for the number of T. minor egg galleries on the trap logs are similar (p > 0.5), confirming the high accuracy of the developed model.     

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.     

Evolutionary history and phylogeography of Scots pine (<Emphasis Type="Italic">Pinus sylvestris</Emphasis> L.) in Europe based on molecular markers

In this review we summarized recent historical records and molecular studies on evolutionary history and phylogeography of Scots pine with focus on the European highly fragmented distribution area of the species. Fossilized pollen, plant micro- and macrofossil records provided evidences on the large-scale species’ range shifts and demographic changes during the Quaternary. Populations of Scots pine were documented both in the glacial (incl. full glaciation) and interglacial periods. Recolonization of Europe after the glaciation originated from the (Sub) Mediterranean areas like the Balkan Peninsula but also from around the Eastern Alps and the surroundings of the Danube plain. Fennoscandia and northern European Baltic regions were most probably colonized from two main directions, from Western Europe and from the Russian Plain. Modern history of Scots pine was hardly affected by anthropogenic activities that started to strengthen in the Bronze and Iron Age. Along with the fossil records, molecular genetic tools were used to infer the origin and putative history including migration, differentiation and demography of the species. In this paper we compiled the major publications (30) of molecular genetic studies of the past 20 years derived from distinctly inherited organelle genomes (mitochondrial, chloroplast, nuclear) revealed by different marker systems (mtDNA-cox1, -nad1, -nad3, -nad7, ISSR, cpSSR, nSSR, B-SAP, SNP). It is important to consider that different phylogeographic patterns can be drawn by the analysis of different DNA marker types. Accordingly the use of more than one marker simultaneously outlines the most sophisticated phylogeographical pattern on the genetic lineages and can reveal high differentiation of the European distribution. Combined marker systems and markers derived from coding sequences have also been used to detect species’ phylogeographic patterns, but these were rarely applied to Scots pine. Although new molecular techniques can provide higher resolution data for populations, the reviewed results can shape the direction of further studies.     

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.

     

Dendroclimatological analysis of seeded and thinned Scots pine (<Emphasis Type="Italic">Pinus sylvestris</Emphasis> L.) stands at the coniferous timberline

Radial growth was examined in two Scots pine stands that were seeded during the 1920s and 1930s due to reforestation and afforestation activity on the timberline of northern Finnish Lapland. Tree-rings of seeded pines were calibrated against the instrumental records of local weather and large-scale atmospheric patterns and further compared to pines of natural origin on the timberline. The studied stands were shown to contain common growth variability that differed from the variability of natural pines. Deviating growth of seeded pines was attributed to their only moderate growth dependence on mid-summer (July) temperatures and, likewise, their strong dependence on the autumn climate in the previous year, and linked to the different genetic origin of artificial stands due to southern seeds. Stands were thinned for different densities in 1985 and 1986. The growth response to thinning was markedly better than could be expected according to previous studies. We found that the positive growth reaction conceivably benefited from the ameliorated winter conditions, expressed as warmed (March) temperatures and the prolonged positive phase of the Arctic Oscillation (February). The results emphasize the determining influence of climatic fluctuations on reforestation and afforestation near the distributional limits of tree species.     

Species-specific spatial structure,species coexistence and mortality pattern in natural,uneven-aged Scots pine (<Emphasis Type="Italic">Pinus sylvestris</Emphasis> L.)-dominated forest

Complexity of uneven-aged forests results from the heterogeneity of their structure reflected among others by the spatial pattern of their components. Forest structure is usually modified by various processes operating at different scales and time. Structure and processes are not independent, and both are important drivers of forest dynamics. The impact of natural processes on forest structure manifested in the specific spatial pattern of trees can be quantified by point pattern analysis applied to long-term repeatedly measured stem-mapped plots. Such studies are relatively scarce in the literature although they provide better insight into the mechanisms affecting forest dynamics. Our study is focused on the spatiotemporal analysis of the structure of mixed uneven-aged Scots pine-dominated forest located at the Kampinoski National Park (Poland). Univariate analysis showed that the initial pattern of all live trees was initially random and it shifted toward more uniform with forest aging. Spatial patterns of individual tree species varied from that stated for all forest community. We observed changes in spatial pattern of Scots pine and common oak from random toward more clumped (pine) or uniform (oak) pattern. In case of black alder and common birch, the initial aggregated pattern was maintained over the examined 14-year period of the forest succession. Bivariate analysis showed that the most common interspecific association between pairs of tree species was spatial segregation (pine vs. alder, alder vs. birch and oak vs. birch) followed by spatial independence (pine vs. oak and oak vs. alder). The positive association was stated only for pine and birch and only for certain spatial scales (> 5 m). Simultaneously, at small distances they showed reciprocal repulsion. Changes in spatial relationships between tree species were negligible over 14-year period of forest succession. Our results confirmed the density-dependent mortality process in the uneven-aged Scots pine-dominated forest over 14-year period of forest development. Our study showed that spatial interactions between individuals along with species-specific ecological requirements should be incorporated into realistic models of forest development, helping to manage the forest ecosystems toward their greater structural complexity.     

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.     

Plastic deformation in small clear pieces of Scots pine (<Emphasis Type="Italic">Pinus sylvestris</Emphasis>) during densification with the CaLignum process

Specimens made of clear wood from Scots pine (Pinus sylvestris L.) were compressed semi-isostatically at 25°C in a Quintus press. Pressure ranged from 0 to 140MPa and the maximum decrease in the crosscut area was about 60%. Quarter-sawn and plain-sawn specimens were densified with the inside face (pith side) up or down. A laser-made dot grid on the crosscut area of the uncompressed specimen was used to calculate plastic strains by image analysis of the displacement of dots after compression. Multivariate models were developed to determine the causes of deformation. The lower face was restrained by the press table and remained flat whereas sides attached to the rubber diaphragm became more irregularly shaped when compressed. Most of the total compression occurred below 50MPa and was determined exclusively by pressure. Above 50MPa, wood density was more important and compression was lower in the interior of specimens and in heartwood. Plastic compressive strain occurred predominately in the radial direction and toward the rigid press table. Strains were dependent on the sawing pattern and orientation. The growth rings of quarter-sawn specimens oriented with the outer face (bark side) down tended to buckle.     

Root architecture might account for contrasting establishment success of <Emphasis Type="Italic">Pseudotsuga menziesii</Emphasis> var. <Emphasis Type="Italic">menziesii</Emphasis> and <Emphasis Type="Italic">Pinus sylvestris</Emphasis> in Central Europe under dry conditions

Key message

Pinus sylvestris seedlings quickly expand their roots to deeper soil layers while Pseudotsuga menziesii concentrates its root system in the topsoil, thereby running the risk of desiccation during long dry spells, as indicated by lower survival after simulated summer drought.

Context

Pseudotsuga menziesii (Douglas-fir) is regarded as a promising species to maintain the productivity of Central European lowland forests given the projected increase of long dry spells.

Aims

Will the species be able to regenerate from seed and spread outside plantations in a drier temperate Europe?

Methods

We measured the relative growth rate, biomass allocation, root architecture, and phenotypic plasticity of Pseudotsuga menziesii seedlings sown in a common garden and grown under current precipitation and prolonged drought, respectively. The species’ competitive ability with respect to Pinus sylvestris L., the most drought-tolerant native conifer in Central Europe, was assessed during three growing seasons.

Results

Pinus sylvestris seedlings had higher relative growth rates than did Pseudotsuga menziesii seedlings, first in terms of aboveground biomass and later in terms of shoot height. This resulted in heavier and taller seedlings after three growing seasons under both moist and dry conditions. Shorter vertical roots corresponded with lower survival of Pseudotsuga menziesii seedlings under dry conditions.

Conclusion

Fast root proliferation allows Pinus sylvestris seedlings to reach deeper water pools that are less rapidly depleted during transient drought. By contrast, the shallow root system might put Pseudotsuga menziesii seedlings at the risk of desiccation during prolonged dry spells.

     

Genetic variability of Italian southern Scots pine (<Emphasis Type="Italic">Pinus sylvestris</Emphasis> L.) populations: the rear edge of the range

Understanding the genetic structure and diversity of edge populations can shed light on the role of peripheral populations and their relevance for conservation strategies. In this study, three fragmented and isolated Apennine populations of Scots pine (Pinus sylvestris L.) belonging to the rear edge of the species’ distribution were analyzed using both nuclear (nSSR) and chloroplast (cpSSR) microsatellites and were compared with an Alpine population belonging to the species’ main range. Although small population size and considerable isolation have probably reduced the genetic variability of Apennine populations, these fragmented populations maintain a high level of within-population genetic diversity. A significant among-population differentiation was found using both nSSR (F _ST = 0.08) and cpSSR markers (ρ = 0.14). Analysis of molecular variance (AMOVA) on the nSSRs attributed all variabilities to the among Apennine populations component supporting the theoretical predictions regarding fragmentation effects on genetic structure. On the other hand, AMOVA on the cpSSRs attributed all variances to the between-region component and no differentiation was found within region, among the Apennine populations. This result suggests the importance of pollen gene flow in homogenizing populations on this geographical scale. Our results confirm the genetic distinctiveness of Apennine populations and their possible derivation from different glacial refugia than those of the Alps. Considering their peculiarity and the high level of intrapopulation genetic diversity that they still retain, fragmented Apennine populations should be considered of high priority for conservation.     

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).     

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.     

Modelling stemflow production by juvenile lodgepole pine (<Emphasis Type="Italic">Pinus contorta</Emphasis> var. <Emphasis Type="Italic">latifolia</Emphasis>) trees

Stemflow is a focused point source input of precipitation and nutrients at the base of a tree or plant and can have a significant impact on site hydrology. To date, no known studies have modelled stemflow production for juvenile lodgepole pine (Pinus contorta var. latifolia). Meteorological conditions, tree characteristics, and stemflow were sampled for two juvenile lodgepole pine stands over the course of the 2009 growing season. Step-wise multiple regression was used to assess which meteorological and tree architecture variables influenced stemflow production for each research plot. Once predictor variables were identified, models were produced for each stand and a generic model was produced that applied to both plots. A model employing precipitation depth and crown projection area successfully explained 71.3% of the variation in stemflow production from sampled trees. Stemflow was found to represent 1.8% of the study period rainfall and, although not a large component of the plot-scale canopy water balance, it is an order of magnitude greater than the fractioning of stemflow from mature lodgepole and lodgepole pine dominated forest. Additionally, stemflow funnelling ratios were found to average 22.2 and 24.3 from the two sample plots over the study period with a single tree, single event maximum of 111.7 recorded for a tree with a 3.3 cm bole diameter and a rain depth of 17.4 mm.     

Effects of spacing and genetic entry on radial growth and ring density development in Scots pine (<Emphasis Type="Italic">Pinus sylvestris</Emphasis> L.)

• Introduction  

The objective of this work was to study how spacing and genetic entry and/or origin group affect radial growth (ring width and cumulative diameter development) and ring density from pith to bark in Scots pine (Pinus sylvestris L.) and in what sense their annual variability could be explained by the climatic variables. The spacing trial was located in central Finland, with current stand density range of 2,000–4,000 trees per hectare. All the six genetic entries had Kanerva pine (plus tree S1101) as a father tree, whereas the mother tree represented Finnish plus trees from southern, central and northern Finland.     

Species and functional diversity of ectomycorrhizal fungal communities on Scots pine (<Emphasis Type="Italic">Pinus sylvestris</Emphasis> L.) trees on three different sites

• Introduction   

In the present study, we describe the aboveground (epigeous sporocarps) and belowground (ectomycorrhizal root tips and their exploration types) composition of ectomycorrhizal (ECM) fungal species on three 20-year-old Scots pine sites.     

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).     

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.