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Importance and potential of Scots pine (Pinus sylvestris L.) in 21st century

Importance and potential of Scots pine (Pinus sylvestris L.) in 21st century We are currently witnessing significant global changes in climate conditions. We cannot change the natural condi- tions, but with regard to sustainable landscape management, we can increase our knowledge of tree species and adapt forest management to them. Surprisingly, one of the most affected tree species in Central Europe today is Scots pine (Pinus sylvestris L.). The following literature review summarizes over 200 studies from 1952–2022 regarding Scots pine across its entire range while addressing various topics in the ecology and management of this taxon. It is a tree species with a large natural range, nearly covering the entire Eurasian area. In the Czech Republic, it is the second most important tree species in terms of industrial wood production. Scots pine is characterized not only by a signifi- cant genetic variability of its populations but also by its wide ecological plasticity. Typically, it grows on sandy soils, poor habitats, and stony scree–but also in peat bogs. The wide habitat valence justifies the economic significance 3 −1 −1 of this species, both in terms of its high production potential (mean annual increment of up to 10.8 m ha yr ) but also its wide range of use. However, in the light of climate variations, the practices of Scots pine silviculture are also gradually transforming from the traditional reforestation by clear-cutting to a more natural system–shelterwood felling. In view of climate change, its range of distribution is changing, as with other species, but Scots pine remains a very resistant tree species, depending on the habitat. Key words: silviculture; ecology; threats; wood production; European forests Editor: Igor Štefančík (Bílek et al. 2016; Hebda et al. 2017; Łabiszak et al. 2017; 1. Introduction Vacek et al. 2022). Different ecotypes of Scots pine are Scots pine (Pinus sylvestris L.) is one of the most ecologi- based on populations that have survived from the Late cally and economically important tree species in Europe Glacial period in the form of isolated refugia in Central (Krakau et al. 2013; O’Reilly-Wapstra et al. 2014; Sevik Europe (Jankovská & Pokorný 2008; Mikeska et al. 2008; & Topacoglu 2015; Wójkiewicz et al. 2016). It is a taxon Tóth et al. 2017). Presumably, some of the refugia of with a wide climatic and edaphic range (Kelly & Connolly autochthonous Scots pine may have provided the basis 2000; Úradníček et al. 2001; Durrant et al. 2016; Vacek et for the developmental lineages of the different ecotypes al. 2016). Due to the fact that it does not tolerate consider- occurring in Central Europe. The oldest pine forests in able shading–among other things–it tends to be displaced the Czech Republic grow on rock outcrops, in rock forma- from rich habitats by competing tree species (Průša 2001; tions, on serpentinite, and mineral-poor and dry sands. Mikeska et al. 2008). However, extensive forest stands of Regarding development, pine forests on peat bogs are Scots pine are typically found on dry and poor sandy soils, younger (Plíva 1971; Mikeska et al. 2008; Poleno et al. in areas with sandstone subsoils, and on extreme sites 2009). The natural habitat for Scots pine ranged from the with limited soil depth, as well as on peatland (Kučera oligotrophic to herb-rich pine forests (Øyen et al. 2006; 1999; Vacek et al. 2017, 2021a; Şofletea et al. 2020). Mikeska et al. 2008; Vacek et al. 2022). In Europe, Scots pine occurs at elevations ranging Historically, the oldest acrofossil evidence and from lowlands to mountains, and in different ecotypes sedimentary records showed the presence of Scots pine *Corresponding author. Jakub Brichta, e-mail: brichtaj@fld.czu.cz © 2022 Authors. This is an open access article under the CC BY 4.0 license. J. Brichta et al. / Cent. Eur. For. J. 69 (2023) 3–20 st between 70,000 BP in Carpathian Basin to 20,000 BP in However, what is the role of Scots pine in the 21 cen- Hungarian plain (Magyari 2011; Tóth et al. 2017). How- tury, when climate change is altering the range of forest ever, current autochthonous pine and pine-oak forests tree species and their growth optimum (Falk & Hemp - were formed by the evolution of vegetation mostly during elmann 2013; Dyderski et al. 2018; Klopčič et al. 2022)? the 10,000 years of the Postglacial era. Scots pine and sil- This literature review, based on over 200 studies, aims ver birch (Betula pendula Roth.) were the most abundant to assess the dynamics, opportunities, and risks of Scots tree taxa in the Early Postglacial. The present occurrence pine from the perspective of climate change in European of Scots pine can, therefore, be considered as a remnant forests with a focus on the Czech Republic. Specifically, of its original range and a relic of the Early Postglacial the review focuses on (i) morphological description of the phases, especially on such ecotopes that were not suit- species, (ii) taxonomic classification, (iii) natural range able for other tree species due to edaphic reasons (Stasz- and distribution, (iv) habitat and ecological preferences, kiewicz 1968; Jankovská & Pokorný 2008). Although (v) silviculture and production, (vi) importance and uses, these azonal communities have gradually evolved since and (vii) threats and diseases, all within the context of the Preboreal, they have only been preserved in extreme climate change. habitats with limited competition from other tree species. The first stands of pine and birch appeared in Central Europe in the older Holocene–in the Preboreal (Husová 1999; Mikeska et al. 2008). In the Boreal period, light 2. Morphological description of the species pine forests with an admixture of common hazel (Cory- Scots pine is characterized by a highly variable habitus, lus avellana L.) appeared in the mentioned area. In the growing to an average height of 26 m but reaching up Atlantic period of climatic optimum, mixed deciduous to 40 m in optimum conditions, while in extreme habi- forests predominated by oak (Quercus spp.) developed, but at the same time, in colder and more humid areas, tats, we find dwarf pines or shrubby pines (Mikeska et al. 2008; Praciak et al. 2013). Scots pine can live up to Norway spruce (Picea abies [L.] Karst.) also spread rap- 300 years (exceptionally, up to 750 years) (Pokorný 1963; idly. Approximately 6200 to 4000 years ago, European Koblížek 2006; Musil & Hamerník 2007; Wallenius et beech (Fagus sylvatica L.) and silver fir (Abies alba Mill.) al. 2010). Many different ecotypes of Scots pine can be began to spread, especially in the mid-elevations (Bolte et al. 2007; Mikeska et al. 2008). All of these tree spe- distinguished, including different characteristics of tree cies then gradually forced pine into habitats where they size or crown shape (Svoboda 1953; Businský 1999). could not compete with it in terms of autoecology. In The root system of the pine is massive, consisting, the Epi-atlantic, a natural zonation of climax vegetation in most cases, of a taproot with richly branched lateral roots (Poleno et al. 2009). Pine roots are very plastic with was established, and pine thus retreated to habitats of an extreme and azonal nature (poor sands, rocks, peat respect to habitat conditions, but they also react to, for bogs). In extreme habitats that were unapproachable to example, the tilting of the tree (Čermák et al. 2008). It is human intervention, communities similar to the current the significant plasticity and resistance of the pine root ones eventually formed (Husová 1999; Poleno et al. 2007; system that makes the pine a valuable stabilizing tree in Mikeska et al. 2008). some habitats (Kacálek et al. 2017). In the northern and Fig. 1. Habitus of the tree, branch, needle, cone, and seed of Scots pine (Pinus sylvestris L.). 4 J. Brichta et al. / Cent. Eur. For. J. 69 (2023) 3–20 northeastern parts of the European range, the Scots pine al. 2001; Koblížek 2006; Musil & Hamerník 2007; Poleno crown is slender with fine branching, whereas arched to et al. 2009; Praciak et al. 2013). The number of pure seeds umbrella-shaped crowns and thick branches are typical in 1 kg is 74–245 thousand. The average weight of 1,000 for pines in Central and Southern Europe. The trunk of seeds is 6.3 g. Well-stored seeds can remain viable for up the tree is most often straight, but on extreme sites, it is to 15 years (Musil & Hamerník 2007; Poleno et al. 2009). often crooked and twisted (Pokorný 1963; Koblížek 2006; Musil & Hamerník 2007; Praciak et al. 2013). At the base of the trunk, the pine is covered with a thick, cracked, grey-brown bark. In the upper parts of the tree, the bark 3. Taxonomic classification turns orange or rusty red, and it peels off in scaly patches (Úradníček et al. 2001; Musil & Hamerník 2007). Scots According to Řepka & Koblížek (2007), Scots pine pine has greenish-brown and glabrous annuals, and (Pinus sylvestris L.) is classified into the following taxo- older twigs tend to be grey-brown (Fig. 1). The needles nomic categories: Domain: Eukarya; Kingdom: Plantae; of Scots pine are stiff and pointed, with a slight longitu- Subkingdom: Viridiplantinae; Developmental lineage: dinal twist, 1–8 cm long, up to 2 mm wide, dark green or Streptophytae; Developmental branch: Cormophytae; bluish grey-green in color, and generally growing in pairs Developmental stage: Gymnospermae; Division: Pyno- on brachyblasts, with a life span of 2–3 years. The buds phyta; Family: Pinaceae; Genus: Pinus; Species: Pinus syl- at the ends of the annuals are elongated-ovate, pointed, vestris, however, splits into numerous lower taxa. Given without resin, or weakly resinous, and covered with rusty- the vast Eurasian range of Scots pine, an intraspecific colored, membranous scales (Pokorný 1963; Úradníček taxonomic system is very difficult to set (Kindel 1995). et al. 2001; Koblížek 2006; Praciak et al. 2013; Krakau Nevertheless, Scots pine can be divided into the follow- et al. 2013). ing four varieties: Pinus sylvestris var. sylvestris; Pinus Scots pine is a monoecious tree species; male and sylvestris var. lapponica Hartm., 1849; Pinus sylvestris female strobili differ in Scots pine. While male strobili var. mongolica Litvinov, 1905, and Pinus sylvestris var. are ovoid, 4–8 mm long, usually yellow (rarely red), hamata Steven, 1838. female strobili are 5–6 mm long, usually pink. The male The variation within the taxon Pinus sylvestris is strobili grow at the base of the elongating shoot instead indeed extremely large. Over 140 subspecies, varieties, of needles, most often in the lower part of the crown, and forms have been studied. In terms of distribution, while the female strobili grow at the end of the last year’s approximately 22 geographical varieties are categorized. branches in the upper part of the crown (Úradníček et al. Svoboda (1953) divides the species into three basic cli- 2001; Musil & Hamerník 2007). The cones are usually matypes: northern, steppe, and mountain pine. Łabiszak single or in groups of 2–3, pedunculate or nearly sessile, et al. (2017) demonstrated the distinct character of fol- ovoid-conical, rounded at the base, often asymmetrical, lowing groups: mountain, lowland and coastal popula- nonglossy, grey-brown, 2.5–7 × 2–3.5 cm. Seed scale tions. Businský (1999) divides Scots pine into varieties shields are rhombic, more developed on the illuminated according to geography and morphological features: side, and flat to pyramidal. The cone umbo is small, flat, P. sylvestris var. sylvestris (including the former var. or short-tipped, light brown, shiny, and without black sibirica), lapponica, hamata, and mongolica. In addi- edging. Seeds are ovoid, 3–4 mm long, whitish, brown or tion to the subdivisions mentioned above, a number of grey to black, with 3–4 times longer brownish to reddish forms have also been detailed in terms of their economic brown wings, and pincer-like at the base (Koblížek 2006; utility–according to the quality of the timber and habi- Musil & Hamerník 2007; Praciak et al. 2013). tus of the pine (in the Czech Republic, “Třeboň pine” or Pines flower in spring and early summer (April– “Týniště pine”). Other forms are based on the variability June) for the first time, at around the age of 15 years. In of needles, bark, and cones (according to the shape of the a closed stand, they do not flower until the age of 30–40 shield, f. plana, f. gibba, f. reflexa, and according to size, (Úradníček et al. 2001; Musil & Hamerník 2007). The f. macrocarpa, f. microcarpa). In nature, Scots pine forms flower primordia of male and female strobili are formed in spontaneous hybrids with Pinus mugo, Pinus uncinata the summer of the previous year (Johnson & More 2006). and Pinus uncinata subsp. uliginosa (Businský 1999; More than 12 months after pollination, the germinating Musil & Hamerník 2007; Poleno et al. 2009; Sobie rajska pollen resumes its growth and fertilizes the egg. Shortly et al. 2020). after, in June (year 2), the entire formation rapidly enlarges and reaches the final cone size in the summer. In early October, the cones ripen. In favorable weather, a small number of seeds emerge during October–Decem- 4. Natural range and distribution ber, but the main period of cone opening is in the spring Scots pine has the largest range of all described pines. The of year 3. The empty cones fall off during the summer of rd distribution range of Scots pine mainly includes the tem- the 3 year after pollination. A seed year in pine occurs on rd th average every 3 to 6 year (Pokorný 1963; Úradníček et perate and cooler belts of much of Europe and Asia (this 5 J. Brichta et al. / Cent. Eur. For. J. 69 (2023) 3–20 Šumava Mountains (the highest occurrence of Scots pine area is otherwise known as Eurasia), and in Europe from northern Portugal and Scotland to the Far East between in the Czech Republic is on the scree near Plešné Lake 37°–70.5° N latitude. The center of its range is Siberia. In at an altitude of 1,070 m). Scots pine is also found on Europe, Scots pine is rarely found in the Mediterranean. sands and peaty soils, the edges of peat bogs in the Třeboň The northernmost occurrence in Europe is in Lapland region, on sandstone cliffs and rock formations in North- (Fig. 2; Úradníček et al. 2001; Musil & Hamerník 2007; ern and Northeastern Bohemia, and on rocky and steep Durrant et al. 2016). slopes of river valleys. It also grows on the outcrops of the Drahanská vrchovina (Drahany Uplands), on the scree Pinus sylvestris var. sylvestris (Syn. P. sylvestris subsp. sibirica [Ledebour] Businsky) is typical in Europe (north of the Hrubý Jeseník Mountains, and on limestone rocks to ca. 62° N), its range extending to the Far East (to ca. of Southern Moravia. Typically, however, Scots pine is 142° E). Pinus sylvestris var. lapponica (Hartman) occurs also cultivated in many places outside of its natural range, from northern Scandinavia to NW Siberia, approximately which is only 3.4% out of the current total of 16.1% in the north of 62° N latitude. Pinus sylvestris var. hamata (Ste- forests of the Czech Republic (Musil & Hamerník 2007; ven) (Syn. Pinus armena K. Koch; P. kochiana Klotzsch Mikeska et al. 2008; Poleno et al. 2009; MZe 2021). ex K. Koch) is found in the Caucasus region and Trans- The frequency of pine forests in the Czech Republic is caucasia south to S Armenia, W Azerbaijan, and Turkey. shown in Fig. 3. In natural forest areas (NFA), the highest Pinus sylvestris var. mongolica (Litvinov) (Syn. Pinus syl- abundance of pines can be seen in the North Bohemian Sandstone Plateau and the Bohemian Paradise – NFA vestris subsp. kulundensis Sukaczev) occurs in Northern Mongolia, NE China, and SE Siberia (Businský 2008). 18 (36.6%), in the South Bohemian Basin – NFA 15 The Scots pine does not occur naturally on the steppes (19.7%), the West Bohemian Upland – NFA 6 (17.14%), of southern Ukraine, southern Russia, or in the oceanic and the Lusatian Sandstone Upland – NFA 19 (11.78%). lowlands of the British Isles and Denmark. Outside its In other natural forest areas of the Czech Republic, the core Eurasian range, however, Scots pine has also sec - representation of pine is significantly lower. ondarily spread in North America, where it is mainly Pine forests have a unique position in the develop- ment and zonation of vegetation. Regarding the co- cultivated on plantations (Poleno 1990; Úradníček et al. 2001; Musil & Hamerník 2007; Schildler et al. 2010). existence of Scots pine with other tree species, its colo- In Central Europe, deciduous forests are the domi - nization of most of the landscape in the Postglacial times nant communities (Leuschner & Ellenberg 2017), was a critical moment. Later, Scots pine spread to soils while pine forests are restricted to poor habitats (Ahti & and habitats where other tree species could not adapt Oksanen 1990). In the Czech Republic, Germany, and (Horsák & Chytrý 2010). Naturally preserved, Scots pine Poland, native Scots pine currently grows only in islands retained its dominant position only on sandstone bedrock and sandy sediments in general, primarily on Cretaceous in extreme relict habitats (Heinken 2007; Chytrý 2013). In Poland, Scots pine is the main economic tree species, sandstones and sands, serpentinite, and, in extreme con- covering 58% of the forest area (DGLP 2021). In the ditions, also on limestone, peats, and on rocky outcrops Czech Republic, Scots pine occupies 16.1% of the forest of various acid rocks (relict). In particular, fires were area (MZe 2021), with its lowest frequency in the Polabí more frequent on dry sands, an essential natural factor sandy terraces of poor loamy sands, on the serpentinite in the colonization of the landscape by pine. The majority rocks of the Slavkov Forest and the Bohemian-Moravian of the sites mentioned above are located approximately in the climate range of forest vegetation zones 3–4, i.e., Highlands, and on the boulder slopes and scree of the Fig. 2. Scots pine distribution in Eurasia; native range; introduced and naturalised (synanthropic) area (Caudullo et al. 2017). 6 J. Brichta et al. / Cent. Eur. For. J. 69 (2023) 3–20 Three groups of relict pine forests can be characterized 300–700 m above sea level. Contrarily, some inversion sites with spruce, or higher, precipitation-deficient areas in Central Europe: on sediments can be assessed as climatically “higher” • Continental Eastern European to South Siberian pine (Mikeska et al. 2008; Vacek et al. 2022). forests growing in contact with sub-xerophilous oak forests on gravelly terraces of larger rivers (class Pul- satillo-Pinetea sylvestris, alliance Pulsatillo-Pinion), their marginal distribution in the Alps is linked to the rain shadow of the inner Alpine valleys. • Basophilous (flowery) pines on marl, limestone, and dolomite rocks and on serpentinites (class Erico- Pinetea, alliance Erico-Pinion), whose distribution extends from the Balkans through the limestone foot- hills of the Alps to Central Europe. • Oligotrophic pine forests belonging to the boreal coniferous forests (class Vaccinio-Piceetea), within which they form a group including primary relict pine forests of siliceous rocks, sandy soils, and peat bog pine forests (alliance Dicrano-Pinion). While the r fi st Fig. 3. Representation of natural pine forests (forest altitudi- two groups are only marginally found in the Czech nal zone 0 – pine forest) in the Czech Republic (GIS-ÚHÚL Republic, oligotrophic pine forests are relatively com- Brandýs n. L., modified according to Mikeska et al. 2008). mon in the Czech Republic (Kučera 1999; Mikeska et al. 2008). Within the European Forest Types (EFT) (Marchetti 2007), all pine forests are designated by the units: 1.2 5. Habitat and ecological preferences Pine and pine-birch boreal forest; 2.2 Sub-boreal Scots pine forest; 2.4 Sub-boreal black pine forest; 2.5 Mixed Scots pine is a typical pioneer tree species (Linder et pine-birch forest (Scots pine); 2.6 Mixed pine-oak for - al. 1997; Durrant et al. 2016). It is a distinctly light- est (pine-oak forest – Scots pine and common oak); 3.1 demanding tree species that barely tolerates shading Subalpine larch-Swiss pine-dwarf pine forest (European but can adapt to a wide range of conditions in terms of larch, Swiss pine, and dwarf pine); 3.3 Alpine pine for- soil and climate requirements (Plíva 1971; Mikeska et est (Scots pine and black pine); 10.1 Thermophilous al. 2008; Poleno et al. 2009). The soils of pine forests on Mediterranean pine forest; 10.2 Black pine forest of the which pine appears are predominantly sandy to gravelly, Mediterranean and Anatolia region; 10.3 Canary Island permeable, arid, and acidic types of arenaceous podzol pine forest; 10.4 Scots pine forest of the Mediterranean or arenaceous cambisol. On extreme geological sub- and Anatolia region; 10.5 Mountain Mediterranean pine strates, it is lithic leptosol, podzolic ranker, and arena- forest; 11.1 Coniferous and mixed peat forest; 11.3 Birch ceous regosol. On sites influenced by water, it is mainly peat forest. stagnic cambisol, stagnic podzol, and gleyic podzol. On In Europe, in the poorest habitats, Scots pine forms transitional peatlands with groundwater, fibric histosol, monocultures. In slightly richer habitats or boggy and occasionally gleyic histosol and gleyic podzol occur. Only upland areas, it grows together with oaks (Quercus pet- in sporadically occurring basophilous pine forests do we raea, Quercus robur), European beech (Fagus sylvatica), find haplic rendzic leptosol, cambic r. l., detrital r. l., or silver birch (Betula pendula), Norway spruce (Picea modal leptosol (Vacek et al. 2022). abies), European larch (Larix decidua), silver fir (Abies Scots pine adapts to a wide climatic range, with a veg- alba), and other pines (primarily Pinus nigra, Pinus etation period of 90–200 days, an annual precipitation of uncinata) (Mason & Alìa 2000; Kelly & Connolly 2000; 200–1,780 mm, and a common mean annual tempera- Úradníček et al. 2001; Musil & Hamerník 2007). ture of 5–9 °C. It tolerates frost and the occasional lack In the Czech Republic, Scots pine is found mainly in of precipitation, as well as poor soil of extreme – rocky, alliance associations Erico-Pinion, Dicrano-Pinion, and sandy, and peaty – habitats, where it is unrivaled in mono- Vaccinion, in rock alliance associations Alysso-Festucion cultures (Richardson 1998; Mikeska et al. 2008). Its deep pallentis, Asplenion serpentini, Seslerio-Festucion glaucae root system and thick bark make Scots pine resistant to (Chytrý et al. 2001). The accompanying tree species of fires and able to regenerate on the mineral soil of burn lowland and wooded-hill variants of Scots pine are mainly sites. Its ecological optimum is far from the physiological Quercus petraea, Tilia cordata, Carpinus betulus, Acer one. Scots pine does not occur naturally in nutrient-rich campestre, and Betula pendula. Within the upland vari- habitats but is often cultivated there locally (Úradníček et ant, Scots pine grows with Picea abies, Abies alba, Fagus al. 2001; Musil & Hamerník 2007; Koblížek 2006; Poleno sylvatica, Larix decidua, and Betula pendula (Mikeska et et al. 2009). al. 2008; Poleno et al. 2009). 7 J. Brichta et al. / Cent. Eur. For. J. 69 (2023) 3–20 In terms of habitat, three basic variants of Scots pine can ditions of natural regeneration can also be impaired by be identified: competition from herbaceous vegetation (Lucas-Borja • Lowland – pioneer: grows primarily on sandy soils, et al. 2011; Mirschel et al. 2011; Prévosto et al. 2012; in monocultures with a minimal or no admixture Hyppönen et al. 2013). If we proceed with the clear-cut, of other tree species, regenerates on mineral soil the use of seed trees is recommended, not only as part in clearings and open areas. It grows quickly when of supporting natural regeneration, but also to increase young, bears fruit early and does not tolerate competi- the radial growth of the remaining mature individuals tion from other species; (Brichta et al. 2019). • Wooded-hill – pioneer: grows mainly on sandy soils, However, one of the most important factors for rocky ecotopes, and peat soils, mostly in monocul- the success of natural regeneration of Scots pine is the tures with little admixture of other tree species, regen- weather conditions, i.e., temperature and precipitation in erates on mineral soil in clear-cuts and open areas; close relation to light during seed germination and initial • Montane – climax: grows mostly in mixtures (with seedling growth (Oleskog & Sahlén 2000b; Puhlick et al. spruce, fir, and beech) at higher altitudes (700–1,000 2012). Pine seeds are able to germinate at 6 °C, however, m) but also descends to lower altitudes. It regenerates the optimum temperature is up to 20–25 °C with a seed under the canopy and does not tolerate open habitats moisture content of approximately 35% (Oleskog & (clear-cuts). It sometimes dominates its competitors Sahlén 2000). Another factor limiting seed germination in height and has a large wood production (Mikeska can be a thick layer of surface humus, which prevents et al. 2008; Poleno et al. 2009). roots from penetrating the mineral soil layer (Hille & Under conditions of global climate change, Scots pine is den Ouden 2004; Oleskog & Sahlén 2000a). Scots pine increasing its range at higher altitudes and in northern germinates optimally only on mineral soil, and therefore locations, and, conversely, declining due to dieback in the soil scarification is usually used in pine regeneration southern part of its European range (Benito Garzón et (Örlander et al. 1996; Remeš et al. 2015; Aleksandrowicz- al. 2008; Reich & Oleksyn 2008; Matìas & Jump 2012). Trzcińska et al. 2017; Saursaunet et al. 2018; Ilintsev et al. 2021). The germination of Scots pine seeds can also be supported artificially - by cold stratification (Houšková et al. 2021) or low-intensive coherent seed irradiation (Novikov et al. 2021). Some sources describe wood ash 6. Silviculture and production fertilization for improving the soil environment (Remeš In silviculture and production, forest structure is a crucial et al. 2016; Petrovský et al. 2018). On dry sites, soil prepa- element, quantified primarily by stand density, canopy, ration also improves the water supply in the root zone vertical canopy structure, stand basal area, horizontal because transpiring herbaceous vegetation is removed by tree distribution, heterogeneity in the spatial arrange- these interventions (Fleming et al. 1994). Moreover, bare ment of trees, the volume of deadwood, or categoriza- mineral soil has less variability in water availability than a tion of individuals into tree classes (Pommering 2002; humus layer (Oleskog & Sahlén 2000a). Soil preparation Puettmann et al. 2008; Silver et al. 2013). Stand struc- increases soil temperature (Nilsson & Örlander 1999; ture noticeably influences most variables in the forest Bedford & Sutton 2000), accelerates humus decomposi- ecosystem, but in the context of forest regeneration and tion, and increases mineral availability (Lunmark-Thelin the silviculture of different tree species, it also influences & Johansson 1997; Nilsson, Örlander 1999; Nilsson et al. the existence and establishment of natural regeneration, 2006), thus increasing the probability and rate of seedling especially for shade-tolerant tree species (Jaworski 2000; growth (Karlsson & Örlander 2000; Mattsson & Berg- Poleno et al. 2009). The aforementioned forest structure sten 2003; Nordborg & Nilsson 2003) and reducing soil issue is crucial for Scots pine silviculture, which requires bulk density (MacKenzie et al. 2005). a high intensity of light for its successful growth in juve- Scots pine produces seeds annually, but moderate to nile stages (Vacek et al. 2016); Oleskog & Sahlén (2000) heavy seed years typically occur every 3–6 years (Poleno reported about 30% of free space light. Numerous stud- et al. 2009; Przybylski et al. 2021). Pine seeds are dis- ies, e.g., Urbieta et al. (2011), Carnicer et al. (2014), and persed primarily by wind, with effective seed dispersal Martin-Alcón et al. (2015) show that as light availability occurring up to a maximum distance of 30–100 m from decreases, the quantity and quality of natural regenera- the parent tree (Farmer 1997; Adams 1992; Mikeska et al. tion decreases in pine stands with higher canopy den- 2008). However, sufficient soil moisture is required for sity. In contrast, Pardos (2017), Schönfelder et al. (2017, seed germination and seedling establishment (MacKen- 2018), and Lundqvist et el. (2019) report that lower light zie et al. 2005). Seedling numbers in Scots pine stands −2 intensity compared to clear-cuts can lead to higher qual- have been reported to range from 0.5–2.3 pcs m (Nils- ity in natural regeneration in Scots pine. Brichta et al. son et al. 2002; Karlsson & Nilsson 2005; Erefur et al. (2020) also mention in their study that partial cover of 2008; Marcos et al. 2007; Marozas et al. 2007; Beghin et the parent stand in turn may have a positive effect on the al. 2010; Mirschel et al. 2011; Jäärats et al. 2012) with a −2 abundance of natural regeneration. Of course, light con- maximum of 10 pcs m (Mirschel et al. 2011). Under- 8 J. Brichta et al. / Cent. Eur. For. J. 69 (2023) 3–20 story recovery typically starts at 10% relative radiation but self-thinning is a natural process in pine stands. On –1 (Ulbrichová et al. 2018). the other hand, higher numbers of individuals ha are In the first years of seedling development, due to recommended in the age of up to about 10 years of growth unfavorable abiotic and biotic factors, seedlings undergo in order to achieve a better morphological quality of the considerable self-thinning. (Aleksandrowicz-Trzcińska trees (Houšková & Mauer 2014). Even before reaching a et al. 2018). The ecotone effect also influences seedling height of 5 m, however, it is necessary to thin out the stand growth, with a higher density of seedlings tending to in order to increase its stability against the action of wet occur at the edge of the stand compared to the interior snow (Novák et al. 2013). The height to diameter ratio is (Vacek et al. 2017b). A risk for newly established pine then most affected by thinning in young pine stands, as stands can be, for example, grubbing by the pine weevil the stand’s age increases, the growth response to thin- (Hylobius abietis L.) (Kovalchuk et al. 2015; Lundborg ning also decreases (Dušek et al. 2011). But it is also pos- et al. 2016) or the formation of proleptic shoots, which, sible to maximize production or reduce silviculture costs however, are removed in sparse cultures and growths by thinning the stand (Sloup & Lehnerová 2016). In most by cutting or simple selection regeneration individuals habitats, the healthy development of pine stands requires (Slodičák & Novák 2007). Although the natural regen- an understory of shade or semi-shade tree species. These eration of Scots pine is generally dominated by small- are usually self-seeding trees, which are intentionally left scale clear-cuts and border cutting areas, shelterwood in the stand during the process of pine stand tending. methods of natural regeneration are increasingly used Later, at about 50 years of age, a combined thinning is in the context of global climate change (Bílek et al. 2016; carried out to encourage the development of as many qual- Brichta et al. 2020). Shelterwood natural regeneration ity individuals as possible depending on the production methods are now common, for example, in Scandi- capacity of the site (about 150–300 target trees). In addi- navia (Hyppönen et al. 2013; Lundqvist et al. 2019), tion to trees suppressing the crowns of the target trees, Germany (Spathelf et al. 2015; Drössler et al. 2017), we remove damaged, diseased, and severely malformed Poland (Bielak et al. 2014; Aleksandrowicz-Trzcińska trees through stand-improving tending operations. We et al. 2017, 2018), and also in some areas in the Czech encourage the presence of soil-improving and admixed Republic (Bílek et al. 2017, 2018; Brichta et al. 2020). tree species to increase the species diversity of pine stands This way of natural regeneration is more favorable with (Poleno et al. 2009; Vacek et al. 2022). Considering the respect to the nature of microhabitats under advancing mostly very poor pine habitats, it is recommended to leave global climate change (Montero et al. 2001; Matías & the residual biomass after thinning in the stand (Novák Jump 2012; Aleksandrowicz-Trzcińska et al. 2014, 2017, et al. 2017), and this despite the consideration of soil pH 2018; Vítámvás et al. 2019; Brichta et al. 2020). deterioration (Peřina & Vintrová 1958). Thus, within the diverse conditions of pine manage- Pine stands are generally restored by border cutting ment, natural regeneration can be achieved by a clear- and small- and large-area clear-cutting, but shelterwood cutting system with different sizes and orientations of cutting is becoming a common practice too. However, cutting. These also include border cutting, patch cutting, the yield from pine stands during regeneration cannot large- and small-area shelterwood cutting, transitioning be precisely totaled. Considering the significant genetic to group or individual selections (Poleno et al. 2009). variability of Scots pine (Kosinska et al. 2007; Businský Within ecologically oriented management, two basic 2008) but also its wide ecological amplitude and habi- silvicultural approaches can be implemented. The first tat range (Plíva 1971; Mikeska et al. 2008; Poleno et al. is to aim for the areal initiation of natural regeneration 2009), the production indices of Scots pine stands vary under the parent stand; the second is small-area group widely (Table 1). Depending on the parameters men- regeneration with a transition to selection principles. In tioned above, as well as on the type and intensity of man- both cases, the start of regeneration must be preceded by agement, we can conclude that Scots pine stand stock the determination of a suitable time frame for the silvi- volumes in Europe are indeed quite variable. While the cultural development of the stand. Determining the mini- studies by Starr et al. (2005) or Makkonen & Helmisaari mum stand age for the start of regeneration depends on (1999) describe roughly 140-year-old pine stands with 3 −1 ha from the lowlands of the specific conditions of the stand, taking into account a stand volume up to 100 m its age, quality, expected production, the presence of Finland, the work by Gallo et al. (2020) reports stand vol- 3 −1 spontaneous regeneration, habitat conditions, and the ume up to 441 m ha in montane pine stands in Spain. nature of the vegetation. The parent stand must not incur Substantially high stocks are in the lowland areas in production losses by premature harvesting, especially of Poland, where the stock of Scots pine stands over 130 3 –1 the best quality trees (Poleno et al. 2009; Bílek et al. 2016, years reaches up to 740 m ha (Bielak et al. 2014). 2018; Vacek et al. 2022). Particularly in Poland, Scots pine is a common tree spe- The first cleaning and thinning from above are car- cies and is even considered the primary economic tree species (DGLP 2021). The highest mean annual incre- ried out by negative selection, i.e., by removing domi- nant, malformed, or damaged individuals at the crown ment (MAI) in the Czech Republic is reported by Vacek 3 –1 –1 and dominant level. Thinning from below is not desirable, et al. (2021a) – 10.87 m ha yr . The high production 9 J. Brichta et al. / Cent. Eur. For. J. 69 (2023) 3–20 Table 1. Overview of available publications related to Scots pine (Pinus sylvestris L.) production parameters. Altitude Age DBH Height Basal area Volume MAI Density Study Country 2 −1 3 −1 3 −1 –1 −1 [m a.s.l.] [year] [cm] [m] [m ha ] [m ha ] [m ha yr ] [trees ha ] Vacek et al. (2016) Czech Republic 245–267 70–130 26–31 21–24 33–40 320–434 3.08–5.50 508–660 Bílek et al. (2016) Czech Republic 270–600 129–191 25–42 14–25 25–47 177–456 0.93–2.39 476–1,072 Bílek et al. (2016) Poland 470 191 42 19 19 159 0.83 200 Vacek et al. (2017) Czech Republic 575–630 123–130 24–36 15–23 12–33 91–267 0.70–2.05 172–512 Gallo et al. (2020) Czech Republic 600–590 142–145 25–27 16–19 27–28 240–245 1.69 488–552 Gallo et al. (2020) Spain 1710 140 21–46 9–20 33–46 231–441 1.65–3.15 276–996 Vacek et al. (2021a) Czech Republic 250–495 40–46 15–22 17–23 38–47 318–500 7.07–10.87 1,355–2,822 Vacek et al. (2021c) Czech Republic 430 48 19 17.5 45 364 7.92 1,700 Starr et al. (2005) Finland 35–280 35–200 12–38 10–25 11–29 48–315 1.37–1.58 374–2,660 Montero et al. (2001) Spain 1700 41–66 17–31 15–21 49 364–478 7.24–8.89 635–2,104 Makkonen & Helmisaari (1999) Finland 144 37 8 8 15 70 1.89 2,660 Van Oijen et al. (2013) Austria 495 60 27 18 — — — 790 Van Oijen et al. (2013) Belgium 50 66 28 20 — — — 380 Van Oijen et al. (2013) Estonia 40 73 26 26 32 374 5.12 — Vanninen & Mäkelä (2000) Finland 150 16–71 4–23 4–22 19–23 — — 595–18,727 Pretzsch et al. (2015) Europe 20–1,290 69 28 22 41 413 11.3 970 Bielak et al. (2014) Poland 79–151 124–132 39–47 30–36 — 319–740 2.57–5.60 177–324 del Río et al. (2008) Spain 1,200–1,750 41–50 14–20 7–13 35–49 159–321 3.89–6.42 1,415–5,495 Beker et al. (2021) Poland 100 25–95 13–31 15–28 27–40 256–396 4.17–10.24 402–2,590 Notes: DBH – diameter at breast height, MAI – mean annual increment potential is also illustrated by the work of Vacek et al. to the wood density of spruce, which is approximately −3 (2021c), where Scots pine achieved the highest incre- 0.410 g·cm (Repola 2006; Saranpää 2003), the wood −3 ment and stand volume of all 12 coniferous tree species density of pine can be up to 0.100 g·cm higher. studied on reclamation dumps following coal mining. On The distinctly differentiated summer rings on pine the other hand, Lovynska et al. (2019) describes that, wood are also complemented by its natural luster; in the case of pine, it is due to its high resin content, which for example, in the conditions of the Northern Steppe of makes the wood very durable, especially in water and Ukraine, Scots pine shows a lower volume production humid environments, which is why it is mainly used for than the acacia tree (Robinia pseudoacacia L.). water structures, pumps, mine timber, sleepers, masts, poles, and fencing; the wood might require impregna- tion for increased durability (Milner 1992; Reynolds & Bates 2009; Farjon 2010; McLean 2019). Pine timber is 7. Importance and use also used to manufacture of timber structures, particu- Scots pine is one of the most important economic tree larly composite timber, in timber construction, lumber, species not only in Central Europe but also in Eurasia furniture, and paneling (Davies et al. 2002; Kuklík 2005; (Praciak et al. 2013; Sevik & Topacoglu 2015; Lundqvist Hairstans 2018), as it has similar durability to larch tim- et al. 2019). Due to its dynamic ecological plasticity and ber (British Standards Institute 1994). Lower-quality ability to occupy hostile habitats, pine plays a crucial role wood is used for fiber and fuel (McLean 2019). in both forestry and, subsequently, in the timber industry. The distillation of the wood was used to prepare tar Pinewood has an orange-brown heartwood and a broader and, subsequently, black pitch, lamp oil, and essential yellow sapwood. The annual rings are very distinctive, oils. Burning the heavily resinous wood of stumps and hence, there is a considerable difference in density and roots yielded soot, which was utilized to make domestic hardness between spring and summer wood (Pokorný ink and printing ink. By scarring the trunks or peeling the 1963). The density of the wood substance reaches val- bark, the resin was extracted (Neumann 2015). In many −3 ues between 0.412 and 0.541 g cm (Table 2). It should countries, including the Czech Republic, the traditional be added, however, that wood density values for pine methods of slitting and debarking live Scots pine trees have a wide variance; this is due not only to the transi- and capturing the resin that oozes out (so-called pitch- tion between spring and summer woods but also to the ing) are no longer allowed. Resin was widely used for extreme genetic variability of the species. The differences sealing and impregnating ships but also as a medicine in wood density of Scots pine individuals may also be due or natural glue. It is also a source of natural turpentine, to its silviculture and different degrees of stand canopy. which, together with its distillation residue (colophony), While individuals with a well-lit crown exhibit lower is the starting material for several other products such as wood density, trees sheltered by the parent stand possess varnishes, paint thinning solvents, insecticides, rubbers, higher wood density (Schönfelder et al. 2017). Compared printing inks, etc. (Schreiner et al. 2018; Praciak et al. Table 2. Overview of the available publications related to Scots pine (Pinus sylvestris L.) wood density. Schönfelder et al. Wagenführ Novák Lexa et al. Repola Saranpää Auty et al. Fundova et al. Study (2017) (2002) (1970) (1952) (2006) (2003) (2014) (2018) −3 Density [g cm ] 0.488–0.541 0.510 0.470 0.510 0.412 0.460 0.423 0.430 10 J. Brichta et al. / Cent. Eur. For. J. 69 (2023) 3–20 2013; Gardner 2013; McLean 2019). Black pine wood ity against insect pests (Allet et al. 2015; Haberstroh et impregnated with resin was also used to make torches al. 2022). Although Scots pine is considered a resistant in the Balkans (Musil & Hamerník 2007). tree species to precipitation deficiency, pine stands across Pine essential oil contains a variety of terpenes. These Europe have still been enormously damaged by recur- substances contained in essential oils and other products rent drought in recent years (Merlin et al. 2015; Vacek are known for their pleasant aroma, which helps to calm et al. 2017; Buras et al. 2018), when, in particular, pre- the nervous system, relieve stress, release anxiety and cipitation is the main factor affecting pine growth pro- tension, and refresh the mind. They are also a component cesses (Vacek et al. 2019). It can be argued that Scots of perfumes, aromatic soaps, massage oils, air freshen- pine is now one of the most threatened tree species in ers, and similar products (Podlech 2002; Schreiner et al. Europe (Gao et al. 2017; Buras et al. 2018; Etzold et al. 2018). Pine bark contains antioxidants, flavonoids, tan- 2019). As a rule, stands with a homogeneous structure nins, and a variety of vitamins and has been consumed or stands with an unsuitable pine ecotype are the most by, for example, Native Americans for centuries. It was affected (Bottero & Vacchiano 2015; van Halder et al. used as a remedy against scurvy by Russian Cossacks in 2019). Paradoxically, the cause of pine dieback may be Siberia and the Far East (Aleksandrov 1969). Tradition- its taproot system (Lokvenc et al. 1985), which does not ally, the inner bark of Scots pine was used by the Sámi adapt to absorb available precipitation from surface soil as a source of food and packaging material in Lapland layers as the water table recedes. Not only is the amount th until the late 19 century (Zackrisson et al. 2000). Com- of available water gradually becoming depleted, but its mercially, the shredded bark of Scots pine is considered nutrients (S, P) are currently lacking in pine stands as a valuable by-product in horticulture (Moore 2011). The well (Prietzel et al. 2020). The solution to the decline bark is also used to make insulation products for build- appears to be the silviculture of structurally differentiated ings (Pásztory & Ronyecz 2013). pine stands (del Río Gaztelurrutia et al. 2017; Brichta et In the past, the maceration of fresh needles was used al. 2020), as well as mixed pine stands (Czerepko 2004; to prepare a tissue called sosnovka or “forest wool”, Pretzsch et al. 2013; Zeller et al. 2017; Vacek et al. 2019). which was used to make carpets, blankets, or as a stuffing In some places, pine is already spontaneously shifting its material. The essential oil contained in pine has medici- range into communities of deciduous trees (Haberstroh nal uses. Extracted from the resin, needles, and buds, it et al. 2022). However, it is still a relatively resistant tree has antiseptic properties. It is used to relieve respiratory species to the effects of climate change, considering the and lung diseases, and rheumatic disorders, as a sedative, habitat. For example, Vacek et al. (2021c) reported that and also in aromatherapy (Ciesla 1998). Scots pine was the most resistant of the 12 tree species In extreme habitats, Scots pine acts as an anti-ero- studied concerning the effects of climate extremes in the sion and reclamation tree species (Vacek et al. 2021a, Czech Republic. c). However, besides the soil-protective function, pine Common insect pests of pine trees include the nun also performs other ecological functions; several fungal moth, pine tree lappet, common pine shoot beetle, bark species form mycorrhizal, parasitic, and saproparasitic beetles (genus Dendroctonus), or tortrix. Trees can also associations with pine trees. About 120 fungal species be attacked by plant parasites and semi-parasites such as have been observed in ecto- and endotrophic symbiosis mistletoe and related species (Mutlu et al. 2016). Trees with pine roots. For consumption, boletes, Bay boletes, weakened by pests or by various abiotic stresses (e.g., brittlegills, and blewits are collected (Klán 1989; Carlile drought) are susceptible to damage by fungal pathogens, & Watkinson eds.1994; Gryndler et al. 2004; Antl 2014). the spread of which may be enhanced in monospecific The collection of bilberries and cranberries, as well as commercial plantations. For example, Sphaeropsis sap- other forest fruits are also abundant in pine forests (Šišák inea and Cenangium ferruginosum cause withering and 2006). dieback of pine trees, while Mycosphaerella pini, Lopho- dermium seditiosum, and related species cause needle cast. Various species of rust and cenangium are also damaging. Cronartium asclepiadeum infests primarily 8. Threats and diseases Scots pine. Pine twisting rust (Melampsora pinitorqua) is a dioecious rust that causes typical twisting of shoots, Currently, the most discussed threat to Scots pine stands especially in Scots pine (Fjellborg 2009). Naemacyclus is undoubtedly drought and the associated decline in needle cast, which causes browning and needle dieback, groundwater levels due to climate change (Vacek et al. is caused by the fungus Cyclaneusma minus. Among the 2016; Gao et al. 2017; Buras et al. 2018). It is climatic wood-destroying fungi are fire sponge, Onnia triquetra, stress periods that negatively affect the photosynthetic crisped sparassis, honey fungus, or velvet-top fungus activity of Scots pine (Flexas & Medrano 2002; Reddy (Businský & Velebil 2011; Pešková & Čížková 2015). et al. 2004). Increasing air temperatures, along with The sawfly species Diprion pini and Neodiprion sertifer low water availability, are responsible for a range of can cause severe defoliation, making the tree susceptible other diseases, as well as reduced tree defense capac- to attack by other pests (Virtanen et al. 1996; Langström 11 J. Brichta et al. / Cent. Eur. For. J. 69 (2023) 3–20 et al. 2001). Scots pine is also attacked by Ips acuminatus, increasing needs of society, a study of the non-productive functions of pine stands is also essential. Pityogenes chalcographus, Tomicus piniperda, Tomicus minor, Phaenops cyanea, and Ips typographus, which can also be a vector of various fungal pathogens, such as Armillaria ostoyae (Kirschner et al. 2001; Jankowiak Acknowledgement & Hilszczański 2011; Giordano et al. 2013; de Rigo et al. 2016). The most important pest of pine seedlings cannot This research was funded by: 1. the Czech University of Life Sci- be neglected, namely the pine weevil (Hylobius abietis ences Prague, Faculty of Forestry and Wood Sciences (No. IGA L.) (Modlinger 2015; Kovalchuk et al. 2015; Lundborg A_21/27); 2. the LIFE Climate Action sub-programme of the et al. 2016). New seedlings and individuals of natural European Union – project CLIMAFORCEELIFE (LIFE19 CCA/ regeneration can also be attacked by Armillaria mellea SK/001276). We would like to thank both Richard Lee Manore, (Nárovcová 2010). For these reasons, fungal pathogens a native speaker, and Jitka Šišáková, an expert in the field, for checking English. We also want to thank Josef Macek for the and nnot only bark insects in pine stands need to be given graphic design of figures. increased attention, and remediation measures should be implemented quickly in case of their occurrence (Zahrad- ník & Zahraníková 2014). 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Importance and potential of Scots pine (Pinus sylvestris L.) in 21st century

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de Gruyter
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© 2023 Jakub Brichta et al., published by Sciendo
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0323-1046
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10.2478/forj-2022-0020
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Abstract

We are currently witnessing significant global changes in climate conditions. We cannot change the natural condi- tions, but with regard to sustainable landscape management, we can increase our knowledge of tree species and adapt forest management to them. Surprisingly, one of the most affected tree species in Central Europe today is Scots pine (Pinus sylvestris L.). The following literature review summarizes over 200 studies from 1952–2022 regarding Scots pine across its entire range while addressing various topics in the ecology and management of this taxon. It is a tree species with a large natural range, nearly covering the entire Eurasian area. In the Czech Republic, it is the second most important tree species in terms of industrial wood production. Scots pine is characterized not only by a signifi- cant genetic variability of its populations but also by its wide ecological plasticity. Typically, it grows on sandy soils, poor habitats, and stony scree–but also in peat bogs. The wide habitat valence justifies the economic significance 3 −1 −1 of this species, both in terms of its high production potential (mean annual increment of up to 10.8 m ha yr ) but also its wide range of use. However, in the light of climate variations, the practices of Scots pine silviculture are also gradually transforming from the traditional reforestation by clear-cutting to a more natural system–shelterwood felling. In view of climate change, its range of distribution is changing, as with other species, but Scots pine remains a very resistant tree species, depending on the habitat. Key words: silviculture; ecology; threats; wood production; European forests Editor: Igor Štefančík (Bílek et al. 2016; Hebda et al. 2017; Łabiszak et al. 2017; 1. Introduction Vacek et al. 2022). Different ecotypes of Scots pine are Scots pine (Pinus sylvestris L.) is one of the most ecologi- based on populations that have survived from the Late cally and economically important tree species in Europe Glacial period in the form of isolated refugia in Central (Krakau et al. 2013; O’Reilly-Wapstra et al. 2014; Sevik Europe (Jankovská & Pokorný 2008; Mikeska et al. 2008; & Topacoglu 2015; Wójkiewicz et al. 2016). It is a taxon Tóth et al. 2017). Presumably, some of the refugia of with a wide climatic and edaphic range (Kelly & Connolly autochthonous Scots pine may have provided the basis 2000; Úradníček et al. 2001; Durrant et al. 2016; Vacek et for the developmental lineages of the different ecotypes al. 2016). Due to the fact that it does not tolerate consider- occurring in Central Europe. The oldest pine forests in able shading–among other things–it tends to be displaced the Czech Republic grow on rock outcrops, in rock forma- from rich habitats by competing tree species (Průša 2001; tions, on serpentinite, and mineral-poor and dry sands. Mikeska et al. 2008). However, extensive forest stands of Regarding development, pine forests on peat bogs are Scots pine are typically found on dry and poor sandy soils, younger (Plíva 1971; Mikeska et al. 2008; Poleno et al. in areas with sandstone subsoils, and on extreme sites 2009). The natural habitat for Scots pine ranged from the with limited soil depth, as well as on peatland (Kučera oligotrophic to herb-rich pine forests (Øyen et al. 2006; 1999; Vacek et al. 2017, 2021a; Şofletea et al. 2020). Mikeska et al. 2008; Vacek et al. 2022). In Europe, Scots pine occurs at elevations ranging Historically, the oldest acrofossil evidence and from lowlands to mountains, and in different ecotypes sedimentary records showed the presence of Scots pine *Corresponding author. Jakub Brichta, e-mail: brichtaj@fld.czu.cz © 2022 Authors. This is an open access article under the CC BY 4.0 license. J. Brichta et al. / Cent. Eur. For. J. 69 (2023) 3–20 st between 70,000 BP in Carpathian Basin to 20,000 BP in However, what is the role of Scots pine in the 21 cen- Hungarian plain (Magyari 2011; Tóth et al. 2017). How- tury, when climate change is altering the range of forest ever, current autochthonous pine and pine-oak forests tree species and their growth optimum (Falk & Hemp - were formed by the evolution of vegetation mostly during elmann 2013; Dyderski et al. 2018; Klopčič et al. 2022)? the 10,000 years of the Postglacial era. Scots pine and sil- This literature review, based on over 200 studies, aims ver birch (Betula pendula Roth.) were the most abundant to assess the dynamics, opportunities, and risks of Scots tree taxa in the Early Postglacial. The present occurrence pine from the perspective of climate change in European of Scots pine can, therefore, be considered as a remnant forests with a focus on the Czech Republic. Specifically, of its original range and a relic of the Early Postglacial the review focuses on (i) morphological description of the phases, especially on such ecotopes that were not suit- species, (ii) taxonomic classification, (iii) natural range able for other tree species due to edaphic reasons (Stasz- and distribution, (iv) habitat and ecological preferences, kiewicz 1968; Jankovská & Pokorný 2008). Although (v) silviculture and production, (vi) importance and uses, these azonal communities have gradually evolved since and (vii) threats and diseases, all within the context of the Preboreal, they have only been preserved in extreme climate change. habitats with limited competition from other tree species. The first stands of pine and birch appeared in Central Europe in the older Holocene–in the Preboreal (Husová 1999; Mikeska et al. 2008). In the Boreal period, light 2. Morphological description of the species pine forests with an admixture of common hazel (Cory- Scots pine is characterized by a highly variable habitus, lus avellana L.) appeared in the mentioned area. In the growing to an average height of 26 m but reaching up Atlantic period of climatic optimum, mixed deciduous to 40 m in optimum conditions, while in extreme habi- forests predominated by oak (Quercus spp.) developed, but at the same time, in colder and more humid areas, tats, we find dwarf pines or shrubby pines (Mikeska et al. 2008; Praciak et al. 2013). Scots pine can live up to Norway spruce (Picea abies [L.] Karst.) also spread rap- 300 years (exceptionally, up to 750 years) (Pokorný 1963; idly. Approximately 6200 to 4000 years ago, European Koblížek 2006; Musil & Hamerník 2007; Wallenius et beech (Fagus sylvatica L.) and silver fir (Abies alba Mill.) al. 2010). Many different ecotypes of Scots pine can be began to spread, especially in the mid-elevations (Bolte et al. 2007; Mikeska et al. 2008). All of these tree spe- distinguished, including different characteristics of tree cies then gradually forced pine into habitats where they size or crown shape (Svoboda 1953; Businský 1999). could not compete with it in terms of autoecology. In The root system of the pine is massive, consisting, the Epi-atlantic, a natural zonation of climax vegetation in most cases, of a taproot with richly branched lateral roots (Poleno et al. 2009). Pine roots are very plastic with was established, and pine thus retreated to habitats of an extreme and azonal nature (poor sands, rocks, peat respect to habitat conditions, but they also react to, for bogs). In extreme habitats that were unapproachable to example, the tilting of the tree (Čermák et al. 2008). It is human intervention, communities similar to the current the significant plasticity and resistance of the pine root ones eventually formed (Husová 1999; Poleno et al. 2007; system that makes the pine a valuable stabilizing tree in Mikeska et al. 2008). some habitats (Kacálek et al. 2017). In the northern and Fig. 1. Habitus of the tree, branch, needle, cone, and seed of Scots pine (Pinus sylvestris L.). 4 J. Brichta et al. / Cent. Eur. For. J. 69 (2023) 3–20 northeastern parts of the European range, the Scots pine al. 2001; Koblížek 2006; Musil & Hamerník 2007; Poleno crown is slender with fine branching, whereas arched to et al. 2009; Praciak et al. 2013). The number of pure seeds umbrella-shaped crowns and thick branches are typical in 1 kg is 74–245 thousand. The average weight of 1,000 for pines in Central and Southern Europe. The trunk of seeds is 6.3 g. Well-stored seeds can remain viable for up the tree is most often straight, but on extreme sites, it is to 15 years (Musil & Hamerník 2007; Poleno et al. 2009). often crooked and twisted (Pokorný 1963; Koblížek 2006; Musil & Hamerník 2007; Praciak et al. 2013). At the base of the trunk, the pine is covered with a thick, cracked, grey-brown bark. In the upper parts of the tree, the bark 3. Taxonomic classification turns orange or rusty red, and it peels off in scaly patches (Úradníček et al. 2001; Musil & Hamerník 2007). Scots According to Řepka & Koblížek (2007), Scots pine pine has greenish-brown and glabrous annuals, and (Pinus sylvestris L.) is classified into the following taxo- older twigs tend to be grey-brown (Fig. 1). The needles nomic categories: Domain: Eukarya; Kingdom: Plantae; of Scots pine are stiff and pointed, with a slight longitu- Subkingdom: Viridiplantinae; Developmental lineage: dinal twist, 1–8 cm long, up to 2 mm wide, dark green or Streptophytae; Developmental branch: Cormophytae; bluish grey-green in color, and generally growing in pairs Developmental stage: Gymnospermae; Division: Pyno- on brachyblasts, with a life span of 2–3 years. The buds phyta; Family: Pinaceae; Genus: Pinus; Species: Pinus syl- at the ends of the annuals are elongated-ovate, pointed, vestris, however, splits into numerous lower taxa. Given without resin, or weakly resinous, and covered with rusty- the vast Eurasian range of Scots pine, an intraspecific colored, membranous scales (Pokorný 1963; Úradníček taxonomic system is very difficult to set (Kindel 1995). et al. 2001; Koblížek 2006; Praciak et al. 2013; Krakau Nevertheless, Scots pine can be divided into the follow- et al. 2013). ing four varieties: Pinus sylvestris var. sylvestris; Pinus Scots pine is a monoecious tree species; male and sylvestris var. lapponica Hartm., 1849; Pinus sylvestris female strobili differ in Scots pine. While male strobili var. mongolica Litvinov, 1905, and Pinus sylvestris var. are ovoid, 4–8 mm long, usually yellow (rarely red), hamata Steven, 1838. female strobili are 5–6 mm long, usually pink. The male The variation within the taxon Pinus sylvestris is strobili grow at the base of the elongating shoot instead indeed extremely large. Over 140 subspecies, varieties, of needles, most often in the lower part of the crown, and forms have been studied. In terms of distribution, while the female strobili grow at the end of the last year’s approximately 22 geographical varieties are categorized. branches in the upper part of the crown (Úradníček et al. Svoboda (1953) divides the species into three basic cli- 2001; Musil & Hamerník 2007). The cones are usually matypes: northern, steppe, and mountain pine. Łabiszak single or in groups of 2–3, pedunculate or nearly sessile, et al. (2017) demonstrated the distinct character of fol- ovoid-conical, rounded at the base, often asymmetrical, lowing groups: mountain, lowland and coastal popula- nonglossy, grey-brown, 2.5–7 × 2–3.5 cm. Seed scale tions. Businský (1999) divides Scots pine into varieties shields are rhombic, more developed on the illuminated according to geography and morphological features: side, and flat to pyramidal. The cone umbo is small, flat, P. sylvestris var. sylvestris (including the former var. or short-tipped, light brown, shiny, and without black sibirica), lapponica, hamata, and mongolica. In addi- edging. Seeds are ovoid, 3–4 mm long, whitish, brown or tion to the subdivisions mentioned above, a number of grey to black, with 3–4 times longer brownish to reddish forms have also been detailed in terms of their economic brown wings, and pincer-like at the base (Koblížek 2006; utility–according to the quality of the timber and habi- Musil & Hamerník 2007; Praciak et al. 2013). tus of the pine (in the Czech Republic, “Třeboň pine” or Pines flower in spring and early summer (April– “Týniště pine”). Other forms are based on the variability June) for the first time, at around the age of 15 years. In of needles, bark, and cones (according to the shape of the a closed stand, they do not flower until the age of 30–40 shield, f. plana, f. gibba, f. reflexa, and according to size, (Úradníček et al. 2001; Musil & Hamerník 2007). The f. macrocarpa, f. microcarpa). In nature, Scots pine forms flower primordia of male and female strobili are formed in spontaneous hybrids with Pinus mugo, Pinus uncinata the summer of the previous year (Johnson & More 2006). and Pinus uncinata subsp. uliginosa (Businský 1999; More than 12 months after pollination, the germinating Musil & Hamerník 2007; Poleno et al. 2009; Sobie rajska pollen resumes its growth and fertilizes the egg. Shortly et al. 2020). after, in June (year 2), the entire formation rapidly enlarges and reaches the final cone size in the summer. In early October, the cones ripen. In favorable weather, a small number of seeds emerge during October–Decem- 4. Natural range and distribution ber, but the main period of cone opening is in the spring Scots pine has the largest range of all described pines. The of year 3. The empty cones fall off during the summer of rd distribution range of Scots pine mainly includes the tem- the 3 year after pollination. A seed year in pine occurs on rd th average every 3 to 6 year (Pokorný 1963; Úradníček et perate and cooler belts of much of Europe and Asia (this 5 J. Brichta et al. / Cent. Eur. For. J. 69 (2023) 3–20 Šumava Mountains (the highest occurrence of Scots pine area is otherwise known as Eurasia), and in Europe from northern Portugal and Scotland to the Far East between in the Czech Republic is on the scree near Plešné Lake 37°–70.5° N latitude. The center of its range is Siberia. In at an altitude of 1,070 m). Scots pine is also found on Europe, Scots pine is rarely found in the Mediterranean. sands and peaty soils, the edges of peat bogs in the Třeboň The northernmost occurrence in Europe is in Lapland region, on sandstone cliffs and rock formations in North- (Fig. 2; Úradníček et al. 2001; Musil & Hamerník 2007; ern and Northeastern Bohemia, and on rocky and steep Durrant et al. 2016). slopes of river valleys. It also grows on the outcrops of the Drahanská vrchovina (Drahany Uplands), on the scree Pinus sylvestris var. sylvestris (Syn. P. sylvestris subsp. sibirica [Ledebour] Businsky) is typical in Europe (north of the Hrubý Jeseník Mountains, and on limestone rocks to ca. 62° N), its range extending to the Far East (to ca. of Southern Moravia. Typically, however, Scots pine is 142° E). Pinus sylvestris var. lapponica (Hartman) occurs also cultivated in many places outside of its natural range, from northern Scandinavia to NW Siberia, approximately which is only 3.4% out of the current total of 16.1% in the north of 62° N latitude. Pinus sylvestris var. hamata (Ste- forests of the Czech Republic (Musil & Hamerník 2007; ven) (Syn. Pinus armena K. Koch; P. kochiana Klotzsch Mikeska et al. 2008; Poleno et al. 2009; MZe 2021). ex K. Koch) is found in the Caucasus region and Trans- The frequency of pine forests in the Czech Republic is caucasia south to S Armenia, W Azerbaijan, and Turkey. shown in Fig. 3. In natural forest areas (NFA), the highest Pinus sylvestris var. mongolica (Litvinov) (Syn. Pinus syl- abundance of pines can be seen in the North Bohemian Sandstone Plateau and the Bohemian Paradise – NFA vestris subsp. kulundensis Sukaczev) occurs in Northern Mongolia, NE China, and SE Siberia (Businský 2008). 18 (36.6%), in the South Bohemian Basin – NFA 15 The Scots pine does not occur naturally on the steppes (19.7%), the West Bohemian Upland – NFA 6 (17.14%), of southern Ukraine, southern Russia, or in the oceanic and the Lusatian Sandstone Upland – NFA 19 (11.78%). lowlands of the British Isles and Denmark. Outside its In other natural forest areas of the Czech Republic, the core Eurasian range, however, Scots pine has also sec - representation of pine is significantly lower. ondarily spread in North America, where it is mainly Pine forests have a unique position in the develop- ment and zonation of vegetation. Regarding the co- cultivated on plantations (Poleno 1990; Úradníček et al. 2001; Musil & Hamerník 2007; Schildler et al. 2010). existence of Scots pine with other tree species, its colo- In Central Europe, deciduous forests are the domi - nization of most of the landscape in the Postglacial times nant communities (Leuschner & Ellenberg 2017), was a critical moment. Later, Scots pine spread to soils while pine forests are restricted to poor habitats (Ahti & and habitats where other tree species could not adapt Oksanen 1990). In the Czech Republic, Germany, and (Horsák & Chytrý 2010). Naturally preserved, Scots pine Poland, native Scots pine currently grows only in islands retained its dominant position only on sandstone bedrock and sandy sediments in general, primarily on Cretaceous in extreme relict habitats (Heinken 2007; Chytrý 2013). In Poland, Scots pine is the main economic tree species, sandstones and sands, serpentinite, and, in extreme con- covering 58% of the forest area (DGLP 2021). In the ditions, also on limestone, peats, and on rocky outcrops Czech Republic, Scots pine occupies 16.1% of the forest of various acid rocks (relict). In particular, fires were area (MZe 2021), with its lowest frequency in the Polabí more frequent on dry sands, an essential natural factor sandy terraces of poor loamy sands, on the serpentinite in the colonization of the landscape by pine. The majority rocks of the Slavkov Forest and the Bohemian-Moravian of the sites mentioned above are located approximately in the climate range of forest vegetation zones 3–4, i.e., Highlands, and on the boulder slopes and scree of the Fig. 2. Scots pine distribution in Eurasia; native range; introduced and naturalised (synanthropic) area (Caudullo et al. 2017). 6 J. Brichta et al. / Cent. Eur. For. J. 69 (2023) 3–20 Three groups of relict pine forests can be characterized 300–700 m above sea level. Contrarily, some inversion sites with spruce, or higher, precipitation-deficient areas in Central Europe: on sediments can be assessed as climatically “higher” • Continental Eastern European to South Siberian pine (Mikeska et al. 2008; Vacek et al. 2022). forests growing in contact with sub-xerophilous oak forests on gravelly terraces of larger rivers (class Pul- satillo-Pinetea sylvestris, alliance Pulsatillo-Pinion), their marginal distribution in the Alps is linked to the rain shadow of the inner Alpine valleys. • Basophilous (flowery) pines on marl, limestone, and dolomite rocks and on serpentinites (class Erico- Pinetea, alliance Erico-Pinion), whose distribution extends from the Balkans through the limestone foot- hills of the Alps to Central Europe. • Oligotrophic pine forests belonging to the boreal coniferous forests (class Vaccinio-Piceetea), within which they form a group including primary relict pine forests of siliceous rocks, sandy soils, and peat bog pine forests (alliance Dicrano-Pinion). While the r fi st Fig. 3. Representation of natural pine forests (forest altitudi- two groups are only marginally found in the Czech nal zone 0 – pine forest) in the Czech Republic (GIS-ÚHÚL Republic, oligotrophic pine forests are relatively com- Brandýs n. L., modified according to Mikeska et al. 2008). mon in the Czech Republic (Kučera 1999; Mikeska et al. 2008). Within the European Forest Types (EFT) (Marchetti 2007), all pine forests are designated by the units: 1.2 5. Habitat and ecological preferences Pine and pine-birch boreal forest; 2.2 Sub-boreal Scots pine forest; 2.4 Sub-boreal black pine forest; 2.5 Mixed Scots pine is a typical pioneer tree species (Linder et pine-birch forest (Scots pine); 2.6 Mixed pine-oak for - al. 1997; Durrant et al. 2016). It is a distinctly light- est (pine-oak forest – Scots pine and common oak); 3.1 demanding tree species that barely tolerates shading Subalpine larch-Swiss pine-dwarf pine forest (European but can adapt to a wide range of conditions in terms of larch, Swiss pine, and dwarf pine); 3.3 Alpine pine for- soil and climate requirements (Plíva 1971; Mikeska et est (Scots pine and black pine); 10.1 Thermophilous al. 2008; Poleno et al. 2009). The soils of pine forests on Mediterranean pine forest; 10.2 Black pine forest of the which pine appears are predominantly sandy to gravelly, Mediterranean and Anatolia region; 10.3 Canary Island permeable, arid, and acidic types of arenaceous podzol pine forest; 10.4 Scots pine forest of the Mediterranean or arenaceous cambisol. On extreme geological sub- and Anatolia region; 10.5 Mountain Mediterranean pine strates, it is lithic leptosol, podzolic ranker, and arena- forest; 11.1 Coniferous and mixed peat forest; 11.3 Birch ceous regosol. On sites influenced by water, it is mainly peat forest. stagnic cambisol, stagnic podzol, and gleyic podzol. On In Europe, in the poorest habitats, Scots pine forms transitional peatlands with groundwater, fibric histosol, monocultures. In slightly richer habitats or boggy and occasionally gleyic histosol and gleyic podzol occur. Only upland areas, it grows together with oaks (Quercus pet- in sporadically occurring basophilous pine forests do we raea, Quercus robur), European beech (Fagus sylvatica), find haplic rendzic leptosol, cambic r. l., detrital r. l., or silver birch (Betula pendula), Norway spruce (Picea modal leptosol (Vacek et al. 2022). abies), European larch (Larix decidua), silver fir (Abies Scots pine adapts to a wide climatic range, with a veg- alba), and other pines (primarily Pinus nigra, Pinus etation period of 90–200 days, an annual precipitation of uncinata) (Mason & Alìa 2000; Kelly & Connolly 2000; 200–1,780 mm, and a common mean annual tempera- Úradníček et al. 2001; Musil & Hamerník 2007). ture of 5–9 °C. It tolerates frost and the occasional lack In the Czech Republic, Scots pine is found mainly in of precipitation, as well as poor soil of extreme – rocky, alliance associations Erico-Pinion, Dicrano-Pinion, and sandy, and peaty – habitats, where it is unrivaled in mono- Vaccinion, in rock alliance associations Alysso-Festucion cultures (Richardson 1998; Mikeska et al. 2008). Its deep pallentis, Asplenion serpentini, Seslerio-Festucion glaucae root system and thick bark make Scots pine resistant to (Chytrý et al. 2001). The accompanying tree species of fires and able to regenerate on the mineral soil of burn lowland and wooded-hill variants of Scots pine are mainly sites. Its ecological optimum is far from the physiological Quercus petraea, Tilia cordata, Carpinus betulus, Acer one. Scots pine does not occur naturally in nutrient-rich campestre, and Betula pendula. Within the upland vari- habitats but is often cultivated there locally (Úradníček et ant, Scots pine grows with Picea abies, Abies alba, Fagus al. 2001; Musil & Hamerník 2007; Koblížek 2006; Poleno sylvatica, Larix decidua, and Betula pendula (Mikeska et et al. 2009). al. 2008; Poleno et al. 2009). 7 J. Brichta et al. / Cent. Eur. For. J. 69 (2023) 3–20 In terms of habitat, three basic variants of Scots pine can ditions of natural regeneration can also be impaired by be identified: competition from herbaceous vegetation (Lucas-Borja • Lowland – pioneer: grows primarily on sandy soils, et al. 2011; Mirschel et al. 2011; Prévosto et al. 2012; in monocultures with a minimal or no admixture Hyppönen et al. 2013). If we proceed with the clear-cut, of other tree species, regenerates on mineral soil the use of seed trees is recommended, not only as part in clearings and open areas. It grows quickly when of supporting natural regeneration, but also to increase young, bears fruit early and does not tolerate competi- the radial growth of the remaining mature individuals tion from other species; (Brichta et al. 2019). • Wooded-hill – pioneer: grows mainly on sandy soils, However, one of the most important factors for rocky ecotopes, and peat soils, mostly in monocul- the success of natural regeneration of Scots pine is the tures with little admixture of other tree species, regen- weather conditions, i.e., temperature and precipitation in erates on mineral soil in clear-cuts and open areas; close relation to light during seed germination and initial • Montane – climax: grows mostly in mixtures (with seedling growth (Oleskog & Sahlén 2000b; Puhlick et al. spruce, fir, and beech) at higher altitudes (700–1,000 2012). Pine seeds are able to germinate at 6 °C, however, m) but also descends to lower altitudes. It regenerates the optimum temperature is up to 20–25 °C with a seed under the canopy and does not tolerate open habitats moisture content of approximately 35% (Oleskog & (clear-cuts). It sometimes dominates its competitors Sahlén 2000). Another factor limiting seed germination in height and has a large wood production (Mikeska can be a thick layer of surface humus, which prevents et al. 2008; Poleno et al. 2009). roots from penetrating the mineral soil layer (Hille & Under conditions of global climate change, Scots pine is den Ouden 2004; Oleskog & Sahlén 2000a). Scots pine increasing its range at higher altitudes and in northern germinates optimally only on mineral soil, and therefore locations, and, conversely, declining due to dieback in the soil scarification is usually used in pine regeneration southern part of its European range (Benito Garzón et (Örlander et al. 1996; Remeš et al. 2015; Aleksandrowicz- al. 2008; Reich & Oleksyn 2008; Matìas & Jump 2012). Trzcińska et al. 2017; Saursaunet et al. 2018; Ilintsev et al. 2021). The germination of Scots pine seeds can also be supported artificially - by cold stratification (Houšková et al. 2021) or low-intensive coherent seed irradiation (Novikov et al. 2021). Some sources describe wood ash 6. Silviculture and production fertilization for improving the soil environment (Remeš In silviculture and production, forest structure is a crucial et al. 2016; Petrovský et al. 2018). On dry sites, soil prepa- element, quantified primarily by stand density, canopy, ration also improves the water supply in the root zone vertical canopy structure, stand basal area, horizontal because transpiring herbaceous vegetation is removed by tree distribution, heterogeneity in the spatial arrange- these interventions (Fleming et al. 1994). Moreover, bare ment of trees, the volume of deadwood, or categoriza- mineral soil has less variability in water availability than a tion of individuals into tree classes (Pommering 2002; humus layer (Oleskog & Sahlén 2000a). Soil preparation Puettmann et al. 2008; Silver et al. 2013). Stand struc- increases soil temperature (Nilsson & Örlander 1999; ture noticeably influences most variables in the forest Bedford & Sutton 2000), accelerates humus decomposi- ecosystem, but in the context of forest regeneration and tion, and increases mineral availability (Lunmark-Thelin the silviculture of different tree species, it also influences & Johansson 1997; Nilsson, Örlander 1999; Nilsson et al. the existence and establishment of natural regeneration, 2006), thus increasing the probability and rate of seedling especially for shade-tolerant tree species (Jaworski 2000; growth (Karlsson & Örlander 2000; Mattsson & Berg- Poleno et al. 2009). The aforementioned forest structure sten 2003; Nordborg & Nilsson 2003) and reducing soil issue is crucial for Scots pine silviculture, which requires bulk density (MacKenzie et al. 2005). a high intensity of light for its successful growth in juve- Scots pine produces seeds annually, but moderate to nile stages (Vacek et al. 2016); Oleskog & Sahlén (2000) heavy seed years typically occur every 3–6 years (Poleno reported about 30% of free space light. Numerous stud- et al. 2009; Przybylski et al. 2021). Pine seeds are dis- ies, e.g., Urbieta et al. (2011), Carnicer et al. (2014), and persed primarily by wind, with effective seed dispersal Martin-Alcón et al. (2015) show that as light availability occurring up to a maximum distance of 30–100 m from decreases, the quantity and quality of natural regenera- the parent tree (Farmer 1997; Adams 1992; Mikeska et al. tion decreases in pine stands with higher canopy den- 2008). However, sufficient soil moisture is required for sity. In contrast, Pardos (2017), Schönfelder et al. (2017, seed germination and seedling establishment (MacKen- 2018), and Lundqvist et el. (2019) report that lower light zie et al. 2005). Seedling numbers in Scots pine stands −2 intensity compared to clear-cuts can lead to higher qual- have been reported to range from 0.5–2.3 pcs m (Nils- ity in natural regeneration in Scots pine. Brichta et al. son et al. 2002; Karlsson & Nilsson 2005; Erefur et al. (2020) also mention in their study that partial cover of 2008; Marcos et al. 2007; Marozas et al. 2007; Beghin et the parent stand in turn may have a positive effect on the al. 2010; Mirschel et al. 2011; Jäärats et al. 2012) with a −2 abundance of natural regeneration. Of course, light con- maximum of 10 pcs m (Mirschel et al. 2011). Under- 8 J. Brichta et al. / Cent. Eur. For. J. 69 (2023) 3–20 story recovery typically starts at 10% relative radiation but self-thinning is a natural process in pine stands. On –1 (Ulbrichová et al. 2018). the other hand, higher numbers of individuals ha are In the first years of seedling development, due to recommended in the age of up to about 10 years of growth unfavorable abiotic and biotic factors, seedlings undergo in order to achieve a better morphological quality of the considerable self-thinning. (Aleksandrowicz-Trzcińska trees (Houšková & Mauer 2014). Even before reaching a et al. 2018). The ecotone effect also influences seedling height of 5 m, however, it is necessary to thin out the stand growth, with a higher density of seedlings tending to in order to increase its stability against the action of wet occur at the edge of the stand compared to the interior snow (Novák et al. 2013). The height to diameter ratio is (Vacek et al. 2017b). A risk for newly established pine then most affected by thinning in young pine stands, as stands can be, for example, grubbing by the pine weevil the stand’s age increases, the growth response to thin- (Hylobius abietis L.) (Kovalchuk et al. 2015; Lundborg ning also decreases (Dušek et al. 2011). But it is also pos- et al. 2016) or the formation of proleptic shoots, which, sible to maximize production or reduce silviculture costs however, are removed in sparse cultures and growths by thinning the stand (Sloup & Lehnerová 2016). In most by cutting or simple selection regeneration individuals habitats, the healthy development of pine stands requires (Slodičák & Novák 2007). Although the natural regen- an understory of shade or semi-shade tree species. These eration of Scots pine is generally dominated by small- are usually self-seeding trees, which are intentionally left scale clear-cuts and border cutting areas, shelterwood in the stand during the process of pine stand tending. methods of natural regeneration are increasingly used Later, at about 50 years of age, a combined thinning is in the context of global climate change (Bílek et al. 2016; carried out to encourage the development of as many qual- Brichta et al. 2020). Shelterwood natural regeneration ity individuals as possible depending on the production methods are now common, for example, in Scandi- capacity of the site (about 150–300 target trees). In addi- navia (Hyppönen et al. 2013; Lundqvist et al. 2019), tion to trees suppressing the crowns of the target trees, Germany (Spathelf et al. 2015; Drössler et al. 2017), we remove damaged, diseased, and severely malformed Poland (Bielak et al. 2014; Aleksandrowicz-Trzcińska trees through stand-improving tending operations. We et al. 2017, 2018), and also in some areas in the Czech encourage the presence of soil-improving and admixed Republic (Bílek et al. 2017, 2018; Brichta et al. 2020). tree species to increase the species diversity of pine stands This way of natural regeneration is more favorable with (Poleno et al. 2009; Vacek et al. 2022). Considering the respect to the nature of microhabitats under advancing mostly very poor pine habitats, it is recommended to leave global climate change (Montero et al. 2001; Matías & the residual biomass after thinning in the stand (Novák Jump 2012; Aleksandrowicz-Trzcińska et al. 2014, 2017, et al. 2017), and this despite the consideration of soil pH 2018; Vítámvás et al. 2019; Brichta et al. 2020). deterioration (Peřina & Vintrová 1958). Thus, within the diverse conditions of pine manage- Pine stands are generally restored by border cutting ment, natural regeneration can be achieved by a clear- and small- and large-area clear-cutting, but shelterwood cutting system with different sizes and orientations of cutting is becoming a common practice too. However, cutting. These also include border cutting, patch cutting, the yield from pine stands during regeneration cannot large- and small-area shelterwood cutting, transitioning be precisely totaled. Considering the significant genetic to group or individual selections (Poleno et al. 2009). variability of Scots pine (Kosinska et al. 2007; Businský Within ecologically oriented management, two basic 2008) but also its wide ecological amplitude and habi- silvicultural approaches can be implemented. The first tat range (Plíva 1971; Mikeska et al. 2008; Poleno et al. is to aim for the areal initiation of natural regeneration 2009), the production indices of Scots pine stands vary under the parent stand; the second is small-area group widely (Table 1). Depending on the parameters men- regeneration with a transition to selection principles. In tioned above, as well as on the type and intensity of man- both cases, the start of regeneration must be preceded by agement, we can conclude that Scots pine stand stock the determination of a suitable time frame for the silvi- volumes in Europe are indeed quite variable. While the cultural development of the stand. Determining the mini- studies by Starr et al. (2005) or Makkonen & Helmisaari mum stand age for the start of regeneration depends on (1999) describe roughly 140-year-old pine stands with 3 −1 ha from the lowlands of the specific conditions of the stand, taking into account a stand volume up to 100 m its age, quality, expected production, the presence of Finland, the work by Gallo et al. (2020) reports stand vol- 3 −1 spontaneous regeneration, habitat conditions, and the ume up to 441 m ha in montane pine stands in Spain. nature of the vegetation. The parent stand must not incur Substantially high stocks are in the lowland areas in production losses by premature harvesting, especially of Poland, where the stock of Scots pine stands over 130 3 –1 the best quality trees (Poleno et al. 2009; Bílek et al. 2016, years reaches up to 740 m ha (Bielak et al. 2014). 2018; Vacek et al. 2022). Particularly in Poland, Scots pine is a common tree spe- The first cleaning and thinning from above are car- cies and is even considered the primary economic tree species (DGLP 2021). The highest mean annual incre- ried out by negative selection, i.e., by removing domi- nant, malformed, or damaged individuals at the crown ment (MAI) in the Czech Republic is reported by Vacek 3 –1 –1 and dominant level. Thinning from below is not desirable, et al. (2021a) – 10.87 m ha yr . The high production 9 J. Brichta et al. / Cent. Eur. For. J. 69 (2023) 3–20 Table 1. Overview of available publications related to Scots pine (Pinus sylvestris L.) production parameters. Altitude Age DBH Height Basal area Volume MAI Density Study Country 2 −1 3 −1 3 −1 –1 −1 [m a.s.l.] [year] [cm] [m] [m ha ] [m ha ] [m ha yr ] [trees ha ] Vacek et al. (2016) Czech Republic 245–267 70–130 26–31 21–24 33–40 320–434 3.08–5.50 508–660 Bílek et al. (2016) Czech Republic 270–600 129–191 25–42 14–25 25–47 177–456 0.93–2.39 476–1,072 Bílek et al. (2016) Poland 470 191 42 19 19 159 0.83 200 Vacek et al. (2017) Czech Republic 575–630 123–130 24–36 15–23 12–33 91–267 0.70–2.05 172–512 Gallo et al. (2020) Czech Republic 600–590 142–145 25–27 16–19 27–28 240–245 1.69 488–552 Gallo et al. (2020) Spain 1710 140 21–46 9–20 33–46 231–441 1.65–3.15 276–996 Vacek et al. (2021a) Czech Republic 250–495 40–46 15–22 17–23 38–47 318–500 7.07–10.87 1,355–2,822 Vacek et al. (2021c) Czech Republic 430 48 19 17.5 45 364 7.92 1,700 Starr et al. (2005) Finland 35–280 35–200 12–38 10–25 11–29 48–315 1.37–1.58 374–2,660 Montero et al. (2001) Spain 1700 41–66 17–31 15–21 49 364–478 7.24–8.89 635–2,104 Makkonen & Helmisaari (1999) Finland 144 37 8 8 15 70 1.89 2,660 Van Oijen et al. (2013) Austria 495 60 27 18 — — — 790 Van Oijen et al. (2013) Belgium 50 66 28 20 — — — 380 Van Oijen et al. (2013) Estonia 40 73 26 26 32 374 5.12 — Vanninen & Mäkelä (2000) Finland 150 16–71 4–23 4–22 19–23 — — 595–18,727 Pretzsch et al. (2015) Europe 20–1,290 69 28 22 41 413 11.3 970 Bielak et al. (2014) Poland 79–151 124–132 39–47 30–36 — 319–740 2.57–5.60 177–324 del Río et al. (2008) Spain 1,200–1,750 41–50 14–20 7–13 35–49 159–321 3.89–6.42 1,415–5,495 Beker et al. (2021) Poland 100 25–95 13–31 15–28 27–40 256–396 4.17–10.24 402–2,590 Notes: DBH – diameter at breast height, MAI – mean annual increment potential is also illustrated by the work of Vacek et al. to the wood density of spruce, which is approximately −3 (2021c), where Scots pine achieved the highest incre- 0.410 g·cm (Repola 2006; Saranpää 2003), the wood −3 ment and stand volume of all 12 coniferous tree species density of pine can be up to 0.100 g·cm higher. studied on reclamation dumps following coal mining. On The distinctly differentiated summer rings on pine the other hand, Lovynska et al. (2019) describes that, wood are also complemented by its natural luster; in the case of pine, it is due to its high resin content, which for example, in the conditions of the Northern Steppe of makes the wood very durable, especially in water and Ukraine, Scots pine shows a lower volume production humid environments, which is why it is mainly used for than the acacia tree (Robinia pseudoacacia L.). water structures, pumps, mine timber, sleepers, masts, poles, and fencing; the wood might require impregna- tion for increased durability (Milner 1992; Reynolds & Bates 2009; Farjon 2010; McLean 2019). Pine timber is 7. Importance and use also used to manufacture of timber structures, particu- Scots pine is one of the most important economic tree larly composite timber, in timber construction, lumber, species not only in Central Europe but also in Eurasia furniture, and paneling (Davies et al. 2002; Kuklík 2005; (Praciak et al. 2013; Sevik & Topacoglu 2015; Lundqvist Hairstans 2018), as it has similar durability to larch tim- et al. 2019). Due to its dynamic ecological plasticity and ber (British Standards Institute 1994). Lower-quality ability to occupy hostile habitats, pine plays a crucial role wood is used for fiber and fuel (McLean 2019). in both forestry and, subsequently, in the timber industry. The distillation of the wood was used to prepare tar Pinewood has an orange-brown heartwood and a broader and, subsequently, black pitch, lamp oil, and essential yellow sapwood. The annual rings are very distinctive, oils. Burning the heavily resinous wood of stumps and hence, there is a considerable difference in density and roots yielded soot, which was utilized to make domestic hardness between spring and summer wood (Pokorný ink and printing ink. By scarring the trunks or peeling the 1963). The density of the wood substance reaches val- bark, the resin was extracted (Neumann 2015). In many −3 ues between 0.412 and 0.541 g cm (Table 2). It should countries, including the Czech Republic, the traditional be added, however, that wood density values for pine methods of slitting and debarking live Scots pine trees have a wide variance; this is due not only to the transi- and capturing the resin that oozes out (so-called pitch- tion between spring and summer woods but also to the ing) are no longer allowed. Resin was widely used for extreme genetic variability of the species. The differences sealing and impregnating ships but also as a medicine in wood density of Scots pine individuals may also be due or natural glue. It is also a source of natural turpentine, to its silviculture and different degrees of stand canopy. which, together with its distillation residue (colophony), While individuals with a well-lit crown exhibit lower is the starting material for several other products such as wood density, trees sheltered by the parent stand possess varnishes, paint thinning solvents, insecticides, rubbers, higher wood density (Schönfelder et al. 2017). Compared printing inks, etc. (Schreiner et al. 2018; Praciak et al. Table 2. Overview of the available publications related to Scots pine (Pinus sylvestris L.) wood density. Schönfelder et al. Wagenführ Novák Lexa et al. Repola Saranpää Auty et al. Fundova et al. Study (2017) (2002) (1970) (1952) (2006) (2003) (2014) (2018) −3 Density [g cm ] 0.488–0.541 0.510 0.470 0.510 0.412 0.460 0.423 0.430 10 J. Brichta et al. / Cent. Eur. For. J. 69 (2023) 3–20 2013; Gardner 2013; McLean 2019). Black pine wood ity against insect pests (Allet et al. 2015; Haberstroh et impregnated with resin was also used to make torches al. 2022). Although Scots pine is considered a resistant in the Balkans (Musil & Hamerník 2007). tree species to precipitation deficiency, pine stands across Pine essential oil contains a variety of terpenes. These Europe have still been enormously damaged by recur- substances contained in essential oils and other products rent drought in recent years (Merlin et al. 2015; Vacek are known for their pleasant aroma, which helps to calm et al. 2017; Buras et al. 2018), when, in particular, pre- the nervous system, relieve stress, release anxiety and cipitation is the main factor affecting pine growth pro- tension, and refresh the mind. They are also a component cesses (Vacek et al. 2019). It can be argued that Scots of perfumes, aromatic soaps, massage oils, air freshen- pine is now one of the most threatened tree species in ers, and similar products (Podlech 2002; Schreiner et al. Europe (Gao et al. 2017; Buras et al. 2018; Etzold et al. 2018). Pine bark contains antioxidants, flavonoids, tan- 2019). As a rule, stands with a homogeneous structure nins, and a variety of vitamins and has been consumed or stands with an unsuitable pine ecotype are the most by, for example, Native Americans for centuries. It was affected (Bottero & Vacchiano 2015; van Halder et al. used as a remedy against scurvy by Russian Cossacks in 2019). Paradoxically, the cause of pine dieback may be Siberia and the Far East (Aleksandrov 1969). Tradition- its taproot system (Lokvenc et al. 1985), which does not ally, the inner bark of Scots pine was used by the Sámi adapt to absorb available precipitation from surface soil as a source of food and packaging material in Lapland layers as the water table recedes. Not only is the amount th until the late 19 century (Zackrisson et al. 2000). Com- of available water gradually becoming depleted, but its mercially, the shredded bark of Scots pine is considered nutrients (S, P) are currently lacking in pine stands as a valuable by-product in horticulture (Moore 2011). The well (Prietzel et al. 2020). The solution to the decline bark is also used to make insulation products for build- appears to be the silviculture of structurally differentiated ings (Pásztory & Ronyecz 2013). pine stands (del Río Gaztelurrutia et al. 2017; Brichta et In the past, the maceration of fresh needles was used al. 2020), as well as mixed pine stands (Czerepko 2004; to prepare a tissue called sosnovka or “forest wool”, Pretzsch et al. 2013; Zeller et al. 2017; Vacek et al. 2019). which was used to make carpets, blankets, or as a stuffing In some places, pine is already spontaneously shifting its material. The essential oil contained in pine has medici- range into communities of deciduous trees (Haberstroh nal uses. Extracted from the resin, needles, and buds, it et al. 2022). However, it is still a relatively resistant tree has antiseptic properties. It is used to relieve respiratory species to the effects of climate change, considering the and lung diseases, and rheumatic disorders, as a sedative, habitat. For example, Vacek et al. (2021c) reported that and also in aromatherapy (Ciesla 1998). Scots pine was the most resistant of the 12 tree species In extreme habitats, Scots pine acts as an anti-ero- studied concerning the effects of climate extremes in the sion and reclamation tree species (Vacek et al. 2021a, Czech Republic. c). However, besides the soil-protective function, pine Common insect pests of pine trees include the nun also performs other ecological functions; several fungal moth, pine tree lappet, common pine shoot beetle, bark species form mycorrhizal, parasitic, and saproparasitic beetles (genus Dendroctonus), or tortrix. Trees can also associations with pine trees. About 120 fungal species be attacked by plant parasites and semi-parasites such as have been observed in ecto- and endotrophic symbiosis mistletoe and related species (Mutlu et al. 2016). Trees with pine roots. For consumption, boletes, Bay boletes, weakened by pests or by various abiotic stresses (e.g., brittlegills, and blewits are collected (Klán 1989; Carlile drought) are susceptible to damage by fungal pathogens, & Watkinson eds.1994; Gryndler et al. 2004; Antl 2014). the spread of which may be enhanced in monospecific The collection of bilberries and cranberries, as well as commercial plantations. For example, Sphaeropsis sap- other forest fruits are also abundant in pine forests (Šišák inea and Cenangium ferruginosum cause withering and 2006). dieback of pine trees, while Mycosphaerella pini, Lopho- dermium seditiosum, and related species cause needle cast. Various species of rust and cenangium are also damaging. Cronartium asclepiadeum infests primarily 8. Threats and diseases Scots pine. Pine twisting rust (Melampsora pinitorqua) is a dioecious rust that causes typical twisting of shoots, Currently, the most discussed threat to Scots pine stands especially in Scots pine (Fjellborg 2009). Naemacyclus is undoubtedly drought and the associated decline in needle cast, which causes browning and needle dieback, groundwater levels due to climate change (Vacek et al. is caused by the fungus Cyclaneusma minus. Among the 2016; Gao et al. 2017; Buras et al. 2018). It is climatic wood-destroying fungi are fire sponge, Onnia triquetra, stress periods that negatively affect the photosynthetic crisped sparassis, honey fungus, or velvet-top fungus activity of Scots pine (Flexas & Medrano 2002; Reddy (Businský & Velebil 2011; Pešková & Čížková 2015). et al. 2004). Increasing air temperatures, along with The sawfly species Diprion pini and Neodiprion sertifer low water availability, are responsible for a range of can cause severe defoliation, making the tree susceptible other diseases, as well as reduced tree defense capac- to attack by other pests (Virtanen et al. 1996; Langström 11 J. Brichta et al. / Cent. Eur. For. J. 69 (2023) 3–20 et al. 2001). Scots pine is also attacked by Ips acuminatus, increasing needs of society, a study of the non-productive functions of pine stands is also essential. Pityogenes chalcographus, Tomicus piniperda, Tomicus minor, Phaenops cyanea, and Ips typographus, which can also be a vector of various fungal pathogens, such as Armillaria ostoyae (Kirschner et al. 2001; Jankowiak Acknowledgement & Hilszczański 2011; Giordano et al. 2013; de Rigo et al. 2016). The most important pest of pine seedlings cannot This research was funded by: 1. the Czech University of Life Sci- be neglected, namely the pine weevil (Hylobius abietis ences Prague, Faculty of Forestry and Wood Sciences (No. IGA L.) (Modlinger 2015; Kovalchuk et al. 2015; Lundborg A_21/27); 2. the LIFE Climate Action sub-programme of the et al. 2016). New seedlings and individuals of natural European Union – project CLIMAFORCEELIFE (LIFE19 CCA/ regeneration can also be attacked by Armillaria mellea SK/001276). We would like to thank both Richard Lee Manore, (Nárovcová 2010). For these reasons, fungal pathogens a native speaker, and Jitka Šišáková, an expert in the field, for checking English. We also want to thank Josef Macek for the and nnot only bark insects in pine stands need to be given graphic design of figures. increased attention, and remediation measures should be implemented quickly in case of their occurrence (Zahrad- ník & Zahraníková 2014). 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Journal

Forestry Journalde Gruyter

Published: Mar 1, 2023

Keywords: silviculture; ecology; threats; wood production; European forests

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