Pine (Pinus spp.) Species in Europe: Ecology, Silviculture, and Ecosystem Services—A Review

Abstract

Pines (genus Pinus) are a diverse group of conifers widely distributed across the Northern Hemisphere, comprising more than 110 species worldwide. In Europe, pines are a defining component of forests, occupying environments that range from dry Mediterranean lowlands to high-elevation alpine zones. They support a wide spectrum of ecosystem services, from timber and non-wood products to protection, biodiversity, and cultural values. In this review, we bring together ecological traits, silvicultural practices, and management considerations for ten European pine species. This review examines ten major European pine species, including Scots pine (Pinus sylvestris), maritime pine (P. pinaster), stone pine (P. pinea), Aleppo pine (P. halepensis), black pine (P. nigra), mountain pine (P. mugo), Swiss stone pine (P. cembra), Turkish pine (P. brutia), Bosnian pine (P. heldreichii), and Macedonian pine (P. peuce). Together, these species span a wide range of European environments, with Pinus sylvestris showing the broadest distribution. Despite the extensive literature on pine species, information on their ecological, silvicultural, and management attributes remains fragmented across disciplines. To bridge this gap, this review draws on an expert-curated core of scientific and technical literature complemented by a structured literature search adapted from the PRISMA framework, compiling within a single framework their main ecological and silvicultural characteristics, altitudinal range, growth patterns, stand diversity, and management practices and the principal goods and ecosystem services they provide. The comparative analysis highlighted distinct functional groups among European pines, illustrating contrasting adaptive strategies across environmental gradients, with altitudinal distributions ranging from sea level to 2400 m. Mediterranean species, such as P. halepensis, P. pinaster, and P. brutia, generally showed higher drought and fire adaptation, whereas mountain taxa such as P. cembra, P. mugo, and P. heldreichii displayed greater cold tolerance but increased vulnerability due to restricted climatic niches and slow regeneration. Broadly distributed species, such as P. sylvestris and P. nigra, exhibited intermediate ecological strategies, with responses strongly influenced by regional climatic conditions. By integrating evidence on distribution patterns, functional traits, growth patterns, and management practices, this review provides a comparative framework to support adaptive forest management and informed species selection, helping to safeguard forest functions and their associated benefits under ongoing climate change.

1. Introduction

Pine species are distributed worldwide [1] and represent the largest share of growing stock in European forests [2]. The role of pine stands goes far beyond timber production. Several woody and non-woody products, protection, conservation, and other goods and services are associated with their stands and forests [3,4]. Ecological and silvicultural traits of the pine species and site (soil, climate, and topography) influence their distribution [5]. Beyond climate, geology, and site conditions, the evolutionary diversification and functional traits of Pinus are also strongly shaped by fire regimes, with species expressing different life history strategies in relation to recurrent, predictable burning [6]. As climate change increases the frequency and severity of natural disturbances across Europe, the vulnerability of forest ecosystems is rising, threatening forest structure, productivity, and the provision of key ecosystem services [7,8,9]. In this context, synthesizing current knowledge on the ecological, functional, and silvicultural characteristics of European pine species is essential to support adaptive forest management and informed species selection under changing environmental conditions. Ten native pine species are found in Europe [3,5]: Pinus sylvestris (Linnaeus, 1753), Pinus pinaster (Aiton, 1789), Pinus pinea (Linnaeus, 1753), Pinus halepensis (Miller, 1768), Pinus nigra (J.F.Arnold, 1785), Pinus mugo (Turra, 1764), Pinus cembra (Linnaeus, 1753), Pinus brutia (Tenore, 1815), Pinus heldreichii (Christ, 1863), and Pinus peuce (Grisebach, 1846). While some species are linked to specific regions, others have a wider distribution. For example, P. pinaster, P. pinea, and P. halepensis are naturally distributed around the Mediterranean region, north and southwards, and P. nigra, P. brutia, P. mugo, P. cembra, P. peuce, and P. heldreichii are restricted by altitude to central Europe and the Balkans [5]. Inversely, P. sylvestris’s natural distribution ranges from Spain to the Balkans southwards and from Ireland to Denmark northwards and is naturalized in central and northern Europe and in vast areas of Asia [5,10]. Also, the trees’ and stands’ main productions vary: whereas P. sylvestris, P. pinaster, P. halepensis, and P. nigra are mainly used for timber production [5,10], P. pinea is used for fruit due to its edible kernels for human consumption [11] and P. cembra and P. mugo are valuable for protection [5,12]. The ten European pine species examined in this review display contrasting ecological strategies but also share traits that are highly relevant for adaptive forest management.

Although European pines have been widely studied, most existing reviews focus on single species or specific regions. What is still missing is a broader perspective that brings together functional traits, ecological strategies, and management implications across the full Mediterranean–Boreal gradient. This review seeks to address these gaps by assembling ecological, functional, and silvicultural information for ten native European pine species within a single comparative framework. Our aim is to clarify how these species behave in their environmental niches, adaptive traits and management requirements, and how these traits can inform climate adaptive forestry. The study is organized as follows: Section 2 informs the review methodology; Section 3 outlines species distributions; and Section 4 presents species-specific ecological and silvicultural traits, silviculture, main goods and services, and diversity of P. sylvestris (Section 4.1), P. pinaster (Section 4.2), P. pinea (Section 4.3), P. halepensis (Section 4.4), P. nigra (Section 4.5), P. mugo and P. cembra (Section 4.6), and P. brutia, P. heldreichii, and P. peuce (Section 4.7). Section 5 synthesizes cross-species patterns and discusses adaptive management, and Section 6 concludes and identifies priorities for future research. The ecological and silvicultural traits of the European pines species and their distribution help clarify their potential uses and management practices best suited to different site conditions, providing a solid basis for adaptive strategies that sustain forest functions and associated benefits under a changing climate.

2. Review Methodology

This review combines a narrative synthesis with a structured literature search procedure adapted from the PRISMA framework. The aim was to reduce selection bias while providing a broad and balanced overview of the available evidence. The review was first built on the authors’ expertise and targeted identification of key reference sources, including peer-reviewed studies and technical or institutional publications, such as those made available by EUFORGEN. This initial step produced a core bibliography of 174 references, which provided the main conceptual, ecological, and silvicultural basis for the review.

To update and complement this initial body of literature, a structured search was conducted on 17 June 2026 in two major bibliographic databases, Web of Science and Scopus. The searches combined the terms Pinus, ecology, silviculture, distribution, functional traits, climate change, and the scientific names of the target species. An additional exploratory search using Google’s search tools was conducted to identify potentially relevant studies that might not have been retrieved through conventional database searches. The full search strings and the corresponding number of records retrieved are presented in the Supplementary Material (Table S1).

Because the initial search retrieved a large and diverse body of literature and the core bibliography already provided comprehensive coverage of earlier studies, database screening focused on studies published from 2022 onwards. This allowed the review to remain manageable while prioritizing recent advances on climate change impacts, species distribution, functional traits, and forest management. All records were brought together into a single reference database, along with the 174 references previously selected through expert knowledge and targeted searches. Duplicates were removed through a combination of exact and fuzzy matching using title, authorship, publication year, and DOI when available. This step removed overlaps between Scopus and Web of Science, between the database results and the original reference set, and among the additional records identified through Google Lab. The literature search revealed differences in evidence availability among European pine species, largely reflecting variations in geographic distribution and research efforts. Species with wider geographic ranges, such as P. sylvestris, were associated with a larger evidence base, whereas species with more restricted distributions were comparatively less represented in the literature. After deduplication and cross-checking with the original reference set, 80 additional unique records remained for eligibility assessment. These studies were then assessed for their relevance to the objectives of the review. This process resulted in the retention of 29 records, yielding a final dataset of 204 studies (175 + 29) included in the qualitative synthesis. A qualitative synthesis was adopted because it is better suited to integrating heterogeneous evidence while preserving the contextual interpretation of the findings and supporting a comparative framework across species. Although the combination of expert-selected references and structured database searches allowed for broader coverage of the relevant literature, the authors acknowledge that this approach may introduce some degree of selection bias.

Overall, the review brings together recent scientific evidence, foundational literature, and relevant technical and institutional sources. The evidence synthesized in this review reflects the inherent variability of ecological systems arising from regional conditions, management contexts, and methodological differences among studies. Broad generalizations based on specific scenarios or modeling projections were deliberately avoided given the uncertainties associated with their assumptions and extrapolation.

3. Distribution of Pine Species in Europe

In Europe, ten native pine species were described, namely P. brutia (Turkish pine), P. cembra (Swiss pine), P. halepensis (Aleppo pine), P. heldreichii (Bosnian pine), P. mugo (mountain pine), P. nigra (European black pine, Austrian pine), P. peuce (Macedonian pine), P. pinaster (maritime pine), P. pinea (umbrella pine, stone pine), and P. sylvestris (Scots pine) [5]. P. sylvestris is native to Spain, Ireland, the United Kingdom, France, Belgium, The Netherlands, Denmark, Germany, Italy, Bosnia and Herzegovina, Croatia, Hungary, Austria, and Slovakia, but it is naturalized from Spain to China (Figure 1).

Figure 1. Distribution of P. sylvestris. Data source: [5].

P. pinaster, P. pinea, and P. halepensis are naturally distributed mainly in the Mediterranean region, ranging from Portugal to Greece and extending beyond Europe from Morocco to Lebanon (Figure 2). P. pinaster has been found outside of its natural range distribution in Algeria, France, Iran, Italy, Morocco, Portugal, and Spain; P. pinea in Albania, Croatia, Cyprus, France, Greece, Italy, Lebanon, Spain, and Turkey; and P. halepensis in Albania, Algeria, France, Greece, Italy, Jordan, Lebanon, Malta, Morocco, Spain, Tunisia, and Turkey [5]. The occurrence of P. halepensis in Umbria has been attributed to a historical introduction from the eastern Mediterranean. Schiller and Brunori [13] have discussed several possible pathways, including introduction by Syrian monks and later Crusaders, considering the former hypothesis to be best supported by historical evidence. P. brutia, P. cembra, P. heldreichii, P. mugo, P. nigra, and P. peuce range from Spain to Greece, Bulgaria, and Romania, extending beyond Europe to Azerbaijan and Iraq (Figure 3) [5]. P. brutia has been found outside of its natural range in Armenia, Bulgaria, Cyprus, Georgia, Greece, Iran, Israel, the Russian Federation, Syria, Turkey, and Ukraine; P. cembra in Austria, France, Slovakia, Slovenia, Switzerland, and Ukraine; P. heldreichii in Albania, Bosnia and Herzegovina, Greece, Italy, Montenegro, and North Macedonia; P. mugo in Austria, Bosnia and Herzegovina, Bulgaria, Croatia, Czechia, France, Germany, Greece, Italy, North Macedonia, Poland, Romania, Serbia, Slovakia, Slovenia, Spain, and Ukraine; P. nigra in Albania, Austria, Bulgaria, Croatia, Cyprus, Greece, Italy, Montenegro, Morocco, North Macedonia, Romania, the Russian Federation, Serbia, Slovenia, Spain, and Turkey; and P. peuce in Albania, Bosnia and Herzegovina, Bulgaria, Greece, Montenegro, North Macedonia, and Serbia [5].

Figure 2. Distribution of P. pinaster, P. pinea and P. halepensis. Data source: [5].

Figure 3. Distribution of P. brutia, P. cembra, P. heldreichii, P. mugo, P. nigra, and P. peuce. Data source: [5].

The current distribution of European pine species reflects major biogeographical gradients shaped by climatic constraints, topography, and human intervention through afforestation. Mediterranean species, such as P. halepensis, P. pinaster, P. pinea, and P. brutia, are largely confined to regions with hot, dry summers. In contrast, mountain taxa, including P. cembra, P. mugo, P. heldreichii, and P. peuce, are found in cooler environments at higher elevations. P. sylvestris and, to a lesser extent, P. nigra have broader ecological tolerances, occupying a wide range of climatic conditions across Europe.

Climate change is expected to substantially alter the abundance and distribution patterns of pine species across Europe [14,15,16]. P. sylvestris is projected to retreat from central and southeast Europe, with its distribution shifting northwards and towards higher elevations in response to increasing drought stress. At the same time, Mediterranean pine species, including P. nigra, P. pinaster, Pinus pinea, and P. halepensis, are expected to expand their ranges northwards into parts of central Europe [15,17]. In contrast, montane species such as P. cembra, P. mugo, P. heldreichii, and P. peuce are likely to experience habitat loss and range contraction as suitable high-elevation environments become increasingly limited [14,18,19]. The magnitude and direction of these distributional changes will depend not only on climatic drivers but also on species-specific dispersal capacity, adaptive potential, land use dynamics, and forest management practices.

4. Characterization, Silviculture, and Products and Services of the Pine Species

4.1. Pinus sylvestris (Scots Pine)

Scots pine (Table 1) is an early successional (pioneer) species with a large natural distribution range, spreading from the Pyrenees in north Spain to the Russian Far East and from Scandinavia to the inner alpine dry valleys in Switzerland and northern Italy. It is the second most widespread conifer tree species in the World after Juniper spp. [20]. Scots pine is a lowland species but occurs until altitudes of more than 2000 m asl [21]. There is a strong discrepancy between the fundamental and realized niche of Scots pine. In the latter, the species is restricted to very poor and acidic sites, e.g., sand dunes, shallow soils on rocks, or fluviatile sands, where competition from other species is low [22]. The species is well-adapted to subcontinental and continental climates (average temperature range from −5 to 20 °C) with variable annual rainfall (300 to 1800 mm) and grows best on soils not affected by stagnant moisture or ground water. Scots pine has a variable root system, often developing a tap root [23]. Litter shows a wide carbon to nitrogen relationship, which is related to a slow decomposition rate.

In the last centuries, Scots pine was often planted to restore degraded landscapes, e.g., on nitrogen-limited (litter-raked) sites. That means that the species was cultivated far beyond its natural range, so it is still covering a significant share of forestland in northern Germany [24] and Poland [25]. The shift from nitrogen-limited forest ecosystems to well-supplied sites in the recent past has boosted volume growth. In the same location, many stands today show higher site indices than some decades before. Scots pine is a moderately water-consuming tree species. In view of enhancing the soil water conditions in a mixed pine stand, the admixture component reduces evapotranspiration losses and therefore improves seepage rates. This is important because in drought-prone northeast Germany, summer precipitation is nearly completely evapotranspirated, i.e., precipitation of winter fills up soil water, whose surpluses feed seepage [24,25].

Scots pine is quite a robust tree species in its core distribution range. Nevertheless, a range of insect species can periodically affect P. sylvestris in so-called mass outbreaks, such as gypsy moth (Lymantria dispar L.), pine tree lappet moth (Dendrolimus pini L.), or pine processionary moth (Thaumetopoea pityocampa Denis and Schiffermüller) [26]. Mixing Scots pine with broadleaved tree species has been the most promising method to mitigate these classical pine pests. Recently, fungi diseases such as the Diplodia shoot decline have emerged. At the southern distribution boundary, e.g., in the inner alpine dry valleys, Scots pine suffers from a disease complex and is gradually substituted by Quercus pubescens (Willd). The disease complex is driven by drought and reinforced by insects, the pine nematode, land use, and additional factors such as mistletoe damage [27]. Currently, in several regions in Germany, Scots pine is affected by cumulative heat and drought events combined with biotic damaging agents, such as pine bark beetles and fungi, with high rates of mortality and salvage cutting [28]. Thus, in Austria and Germany, Scots pine, especially in pure and homogeneous secondary stands, has recently been identified as one of the species most adversely affected by climate change and is not considered a suitable large-scale substitute for Picea abies (L.).

As a pioneer species, Scots pine shows rapid growth in the juvenile stage. Mean annual increments range from 8 to 14 m³ha⁻¹, varying by ecoregion. In Germany, the latest National Forest Inventory (BWI IV) specifies a mean annual increment (MAI) of around 8 m³ha⁻¹y⁻¹ for Scots pine. Thus, Scots pine’s performance is average and lies between European beech (Fagus sylvatica L.) and oak (Quercus spp.) but is far below the more productive conifers such as Picea abies, Abies alba (Mill.), or the exotic Pseudotsuga menziesii (Mirb.) Franco) [10]. Scots pine is not a very responsive species. If released from competitors, the remaining trees respond well only at young ages. With increasing age, pine trees cannot compensate for an increasing standing space. On a European gradient from northeast to southwest, central European pine stands occupy a mean growth level [29].

Growth trends for Scots pine have been increasing for a long time due to nitrogen emissions (for south Germany) and warmer winter temperatures. However, increasing sensitivity to summer drought and high air temperatures has negatively exposed Scots pines to climate change, which cannot be compensated by the positive growth stimulus of warmer winter temperatures [30]. Scots pines’ mortality has sharply increased in recent years due to climate change. In particular, bark beetles and wood-boring pests have become more aggressive and replace previous damaging agents. High mortality occurs in the mid latitudes and can be assessed by looking at the steepening self-thinning line. Tree-level acclimation strategies, such as lowering leaf area and enhancing water use efficiency, as well as stand-level adjustments, including self-thinning, are apparently insufficient to prevent drought-induced Scots pine mortality [31].

The preferred type of regeneration in Scots pine stands in central European lowlands is natural regeneration on mostly medium to poor sites. Widespread regeneration methods for Scots pine are still clearcut and strip cut. In Scandinavia, the seed tree system also plays a significant role in naturally regenerating Scots pine stands. Common to all of these systems is the creation of bare land conditions for establishing a sufficient amount of seedlings. As a climatically robust species, Scots pine tolerates intensive radiation and frost events, thus showing the typical characteristics of an early-successional pioneer species [10]. Alternatives to clear or strip cutting are still rare for this pine species. Shelterwood cutting or retention systems have a long tradition; the forests, in general, were managed as even-aged pure stands. The suitability of Continuous-Cover Systems for pine management is still questioned due to the difficulty of establishing a vertically structured forest with a light-demanding species. In central Europe, only very few examples exist where such forests can be seen in practice. Even the pine forest of Bärenthoren (near Berlin, Germany), which was the model forest for Alfred Möller to develop his Continuous-Cover Forestry (CCF) concept in the 1910s (The “Dauerwald” [32]), was a two-layered open pine forest with low standing volumes due to litter raking and a lack of nitrogen supply.

Pine cones germinate when there is contact with the mineral soil. Recent studies show that seedling density and the spatial occurrence of seedlings strongly depend on soil scarification and overstory density. Moreover, many other ecological factors influence the success of germination and seedlings’ establishment, such as diaspore supply, browsing, water conditions of the soil, and herbaceous competition. Especially for seedlings and saplings less than 50 cm in height, negative correlations with browsing intensity and herb competition by Vaccinium spp. have been identified.

A gradient study across Europe [29] showed that volume increment decreases with increasing thinning intensity, particularly on fertile sites and in northern Europe. Height growth is slightly higher in thinned stands compared to controls because thinning is lowering drought stress among trees, especially when precipitation is low [29]. Thinning has a particularly strong positive effect on diameter development, e.g., on the future crop trees of a stand. However, these gains should be balanced against reduced stand volume production. Heavy crown thinning at an early stage, combined with artificial pruning, is recommended for the production of high-value timber.

In general, thinning increases the stability of Scots pine stands in the long term, e.g., by positively influencing the h/d ratio of trees. Nevertheless, immediately after thinning, stands are often vulnerable, at least for a short period. As storm damage increases with stand and tree height [33], thinning at an early stage is an important measure for stabilization purposes. Moreover, thinning has been proven to improve resilience (recovery) after drought, at least at younger ages. However, the more frequently drought years occur, the lower trees’ resilience. There is evidence that the water use efficiency of Scots pine depends on provenance, as revealed by the international provenance trial Chorin 85 from 1908, e.g., French provenances are more stress-tolerant than Baltic provenances, but the former having a lower volume performance. Nevertheless, analyses of climatic stress in Scots pine populations confirm that especially the rear edge (e.g., southern) provenances will be prone to future mortality [34]. Provenances should be selected according to the future climate of the target region, as Wegener et al. [35] highlight.

Thinning has only a marginal effect on stand microclimate (e.g., relative soil humidity, relative humidity); mixture with broadleaves and thus a higher leaf area index (LAI) and changes over time in temperature or understory day length are much more significant. Intensive thinning leads to tapered wood and big knots but good allometry concerning stability (height/diameter values). However, wood quality is reduced, along with density reduction. The effect of overyielding mixtures of Scots pine and Fagus sylvatica has been proven in several studies, especially in a landmark investigation along a European gradient from southwest to northeast Europe [36,37]. Due to their complementarity in resource utilization, Scots pine and Fagus sylvatica produced standing volume increases of +12% and stand volume growth increases of +8% compared to the weighted mean of the neighboring monocultures. Overyielding is observed, e.g., in mixtures with European beech. One reason is the more efficient packing of the trees in mixtures per area.

Table 1. Characteristics of Pinus sylvestris.

Indicator	Details	References
Ecological and silvicultural characteristics	Light-demanding pioneer species, less competitive; medium volume increment level; highly vulnerable to many biotic damaging agents, prone to fire damage and medium drought-resistant; many provenances due to large natural distribution range	[22]
Silviculture	Short to medium rotation species; suited for clear and patch cutting systems; regeneration best on mineral soil; difficult in CCF systems	[37]
Main goods and services	Mass production timber and industrial, rarely high-quality wood; diverse NWFP, such as resin	[10]
Diversity	Only in mixed stands and stands with microhabitats	[37]

Table 1. Characteristics of Pinus sylvestris.

Indicator	Details	References
Ecological and silvicultural characteristics	Light-demanding pioneer species, less competitive; medium volume increment level; highly vulnerable to many biotic damaging agents, prone to fire damage and medium drought-resistant; many provenances due to large natural distribution range	[22]
Silviculture	Short to medium rotation species; suited for clear and patch cutting systems; regeneration best on mineral soil; difficult in CCF systems	[37]
Main goods and services	Mass production timber and industrial, rarely high-quality wood; diverse NWFP, such as resin	[10]
Diversity	Only in mixed stands and stands with microhabitats	[37]

In Germany, 22.000 ha of pure stands were converted annually into mixed (and sometimes uneven-aged) stands between 2000 and 2017. Scots pine conversion into mixed Scots pine–broadleaved stands is a frequent measure on fertile sites in northeast Germany. The applied technique is underplanting or natural regeneration in the case of animal seed dispersal. To establish mixed forests, European beech or even oak species can be planted or sowed under fairly dense pine forests. In the case of enough light, Scots pine grows rapidly in the first development stages and outcompetes other competing tree species. The crop tree concept is a feasible option to select superior trees in the stand and to produce valuable timber. Here, artificial pruning is inevitable to maintain high-quality timber. Target diameter harvesting starts at a diameter at breast height (dbh) of 40 to 60 cm, the mature trees being removed by creating small clear or patch cuts [10]. The culmination of value production in northeast Germany lies between 70 and 90 years. Due to its behavior as a light-demanding pioneer species, Scots pine tends to develop even-aged cohorts. A specific silvicultural system to provide good seed supply and to enhance the value production of a pine stand is the seed tree system or the hold-over of trees with potentially high value for another forest generation. Finally, thinning positively influences a diverse range of ecosystem services, such as Non-Wood Forest Products (NWFP) (e.g., mushrooms and berries), or habitats for specific species, such as the capercaillie (Tetrao urogallus L.). Moreover, Scots pine stands are important recreation forests in many northern (boreal) countries [10].

Mixing pine, especially with shade-tolerant or intermediate tree species, such as Fagus sylvatica or Tilia spp., increases the functional diversity and structural heterogeneity of forests [38], whereas the competitive advantages of the tree species in mixtures change annually due to climate variability [39]. Thus, thinning reduces tree mortality, which is also disadvantageous, because deadwood is important for antagonists and high species diversity [29]. The rationale for mixing other tree species with Scots pine is evident: productivity (overyielding) [36], resilience [40], higher faunal biodiversity, and economic advantages due to diversification are the most important points in this context [41]. In a landmark study, Jactel et al. [40] showed that diversity makes forest ecosystems more resilient against pests and diseases. The underlying effects are more habitats for diverse species and thus more antagonists, preventing mass outbreaks of damaging agents. Moreover, in a recent study, Lasek et al. [42] showed that historical admixture has enhanced the genetic diversity and potential adaptive capacity of Polish Scots pine seed stands.

Species richness in pine stands in general enhances faunal diversity in forest ecosystems [29] and diversifies production risks. The most important admixed tree species to P. sylvestris in central Europe are Fagus sylvatica, Quercus spp., Picea abies, and Betula pendula (Roth). They can be admixed in rows or patches to the target species on a spatial scale and below the canopy of an old stand for regeneration. The underplanted broadleaved species component can serve either as the new forest generation or as a trainer species to enhance the quality of the target tree [37].

4.2. Pinus pinaster (Maritime Pine)

Maritime pine (Table 2) has a broad ecological range supported by high genetic variation and phenotypic plasticity. Population divergence is largely shaped by summer drought and cold hardiness ([43] and references therein). The high evolvability of key adaptive traits, including plasticity in water use efficiency, is expected to facilitate rapid microevolutionary responses under ongoing climate change [44]. The species thrives from sea level to 800 m asl and above. It prefers mean annual temperatures of 13–15 °C and typically occurs in areas receiving 400–800 mm of annual rainfall or more. The species prefers light-textured, well-drained soils and is acknowledged for its fast growth and intolerance to shade. Flowering occurs from March to May, with cones maturing in autumn [5,45,46].

Regeneration can be achieved artificially via planting or direct seeding, with initial stocking densities typically ranging from 1100 to 2800 trees ha⁻¹. The species also regenerates after fire through serotinous cones [47]. These cones may open gradually over several years or rapidly following a fire, thereby promoting post-fire recruitment. In naturally regenerating post-fire stands (common in Mediterranean countries), initial densities can be extremely high, ranging from 7500 to 90,000 seedlings ha⁻¹ during the first 20 years [48]. Such natural regeneration is often used to reduce establishment costs. However, this process faces a significant bottleneck during the first year, as seedling survival is frequently compromised by summer drought and intense water stress [47]. De Frutos et al. [49] demonstrated that group selection cuttings can promote natural regeneration and structural diversification in P. pinaster plantations, although early seedling survival remains a critical bottleneck influenced by shrub competition, litter accumulation, and microclimatic conditions.

The species is managed in high forest systems typically as even-aged pure stands, although mixed stands with other pines (e.g., P. halepensis, P. nigra, P. pinea, P. sylvestris) or with broadleaved species (e.g., Castanea sativa, Quercus robur, Eucalyptus spp.) have also been reported [46]. In pure stands, traditional management regimes are oriented towards roundwood production, starting from relatively low initial densities (1100–1200 trees ha⁻¹) and harvesting at minimum rotation ages of 40–50 years. Thinning from below is the most commonly applied method; in timber-oriented systems, 3–4 thinnings are typically recommended between 10 and 30 years, resulting in a final stocking of approximately 300 trees ha⁻¹. To regulate P. pinaster stand density, spacing factors have been widely used (e.g., [50,51,52] and references therein). Luís and Fonseca [53] proposed the use of the Stand Density Index (SDI) to define optimal density ranges, suggesting 60% and 35% of SDI as the upper and lower bounds of the growth density interval. Fonseca and Duarte [52] further recommended a spacing factor of 0.21 to limit understory development, reduce fire vulnerability, and maximize timber yield. Other authors recommend defining thinning intensity based on basal area, with the appropriate level depending on initial stand density and management objectives. According to Erdozain [54], basal area removals of 23%–69%, particularly 50%–70% in dense stands, enhance natural regeneration by increasing seedling establishment and early survival, especially under warm and dry conditions. Short rotation management of maritime pine has also been proposed for energy production, with clearcut ages reduced to approximately 15–20 years. Cotton [55] describes a regime with an initial stand density of ~2500 trees ha⁻¹ and harvest at 15 years. Fonseca and Lousada [56] analyzed a comparable option for post-fire naturally regenerated stands characterized by very high initial densities (~40,000 trees ha⁻¹) and harvest at 20 years. These regimes represent short rotation systems that resemble “energy crops” in which harvested biomass is primarily used for bioenergy, pulp, panels, or pilings. Fonseca and Lousada [56] report that reducing the rotation from 45 to 20 years markedly decreases total volume and stem dimensions, limiting value for large-sized timber but offering advantages when the main objective is the production of small-sized material.

The species is susceptible to diseases and pests, such as processionary caterpillar (Thaumetopoea pityocampa Denis and Schiffermüller) and to root rot pathogen (Heterobasidion annosum (Fries) Brefeld). In addition, these resources face substantial biological risk from the pine wood nematode (Bursaphelenchus xylophilus (Steiner and Bührer) Nickle), as climate change is expected to increase the pathogenicity and expand the distribution of the nematode species [57]. To combat threats such as pine wilt disease and increasing aridity, silvicultural strategies are shifting toward large-scale mixed forest conversion [58]. In addition, adaptive silviculture based on moderate thinning, structural heterogeneity, and the retention of large trees has been identified as a key strategy to enhance drought resilience in dry-edge P. pinaster forests [59].

The wood of P. pinaster is characterized by long tracheids, which provide high axial cohesion and favorable mechanical properties, making it highly versatile. Accordingly, it is used in a wide range of products, from low-value assortments such as packaging and pallets to higher-value applications including structural timber, flooring, carpentry, furniture, veneer, particleboard, plywood, poles, and sleepers [46]. Resin tapping also represents an important by-product. Beyond its economic value, P. pinaster plays a key role in carbon sequestration, dune stabilization, and biodiversity conservation by supporting diverse understory communities (for further details, see [46]).

Table 2. Characteristics of Pinus pinaster.

Indicator	Details	References
Ecological and silvicultural characteristics	Altitude up to 800 m, although altitudes up to 2000 m are observed. Climate tolerance: Mean annual temperature 13–15 °C, minimum 1–2 °C, absolute minimum –15 °C, absolute maximum 40 °C. Annual precipitation: 400–800 mm and larger than 800 mm. Tolerances: Frost sensitivity: Winter, low; spring, high. Wind sensitivity: Low; with salt: low. Mean sensitivity to drought. Resistant to fire and wind. Soil: Avoids soils with bad drainage, compacted, humid, and calcareous soils. Ecology: Intolerant to shade. Longevity 150–300 years. Growth is fast.	[5,45,46]
Silviculture	Propagation: Flowers occur March–May. Fruit maturation occurs in autumn. Seed: Produces 8–30 kg of seed per tree, with an average of 16,000 to 17,000 seeds per kg. Dissemination occurs via wind and animals. The species is typically managed as even-aged, pure high forest stands, originating from either natural or artificial regeneration. Uneven-aged and mixed stands are less common. Potential issues: Sensitive to pests and diseases, particularly the pine wood nematode. Exhibits post-fire regeneration through serotinous cones.	[5,45,46,47,60,61,62,63]
Main goods and services	Products: Timber, biomass, and resin. Main uses: Timber, wood panels, paper, pellets, resin, protection, and nursery substrate (bark). Suitable for forest restoration in dry, sandy, nutrient-poor soils. Use in fire-prone areas due to serotinous cones. Plantations in coastal or windy environments, though salt wind sensitivity varies.	[5,45,64]
Diversity	International Union for Conservation of Nature (IUCN) status: Least Concern (LC), wide distribution along the western Mediterranean. Genetic diversity structured in regional provenances. Key conservation strategies: Maintain local provenances, avoid genetic erosion in reforestations, promote mixed stands to increase ecosystem diversity.	[65]

Table 2. Characteristics of Pinus pinaster.

Indicator	Details	References
Ecological and silvicultural characteristics	Altitude up to 800 m, although altitudes up to 2000 m are observed. Climate tolerance: Mean annual temperature 13–15 °C, minimum 1–2 °C, absolute minimum –15 °C, absolute maximum 40 °C. Annual precipitation: 400–800 mm and larger than 800 mm. Tolerances: Frost sensitivity: Winter, low; spring, high. Wind sensitivity: Low; with salt: low. Mean sensitivity to drought. Resistant to fire and wind. Soil: Avoids soils with bad drainage, compacted, humid, and calcareous soils. Ecology: Intolerant to shade. Longevity 150–300 years. Growth is fast.	[5,45,46]
Silviculture	Propagation: Flowers occur March–May. Fruit maturation occurs in autumn. Seed: Produces 8–30 kg of seed per tree, with an average of 16,000 to 17,000 seeds per kg. Dissemination occurs via wind and animals. The species is typically managed as even-aged, pure high forest stands, originating from either natural or artificial regeneration. Uneven-aged and mixed stands are less common. Potential issues: Sensitive to pests and diseases, particularly the pine wood nematode. Exhibits post-fire regeneration through serotinous cones.	[5,45,46,47,60,61,62,63]
Main goods and services	Products: Timber, biomass, and resin. Main uses: Timber, wood panels, paper, pellets, resin, protection, and nursery substrate (bark). Suitable for forest restoration in dry, sandy, nutrient-poor soils. Use in fire-prone areas due to serotinous cones. Plantations in coastal or windy environments, though salt wind sensitivity varies.	[5,45,64]
Diversity	International Union for Conservation of Nature (IUCN) status: Least Concern (LC), wide distribution along the western Mediterranean. Genetic diversity structured in regional provenances. Key conservation strategies: Maintain local provenances, avoid genetic erosion in reforestations, promote mixed stands to increase ecosystem diversity.	[65]

4.3. Pinus pinea (Umbrella Pine, Stone Pine)

Umbrella pine (Table 3) is considered a pioneer species well-adapted to a wide range of mean annual precipitations (300–1500 mm) and mean annual temperatures (10–18 °C) and altitudes (0–1000 m asl) [5,45]. It develops well in most soils, except for those that are excessively compact with poor drainage and/or very calcareous [45]. Its sensitivity to winter frosts is low and to spring ones moderate [5]. It is resistant to wind and to wind with salt [45] and has moderate sensitivity to drought [5]. It is sensitive to fire [45] and several pests and diseases [66,67]. In former years, the western conifer seed bug, Leptoglossus occidentalis Heidemann, was the biotic agent causing the greatest damage to cones [68].

It is a semi-shade-tolerant species in its juvenile phase, becoming light-demanding at maturity [5,45]. Its longevity is about 150–300 years [45], and it has an intermediate growth rate, though it depends on the climate. Tree growth in P. pinea is primarily limited by water availability rather than temperature. While mild winters and adequate autumn–spring precipitation promote radial growth [69,70,71], prolonged spring and summer drought reduce cambial activity and disproportionately affect younger trees because of their shallower root systems [69,70,71,72,73,74]. These patterns indicate that water availability during the growing season is the principal ecological constraint shaping productivity across Mediterranean environments. The negative effects of low precipitation (and drought) can be mitigated by mist precipitation (in areas with high relative air humidity) that can incorporate considerable amounts of water in the soil [75]. Soil nutrient and water holding capacity influences growth, with lower growth rates at poor sites [73,76].

Flowering occurs from March to May and fruiting in the spring [45], with cones needing 4 years to mature [77]. Fruiting starts at 15–20 years [45] or sooner in grafted individuals [78]. Tree production varies between 1 and about 300 cones per tree and is related to stand density and climate [79]. The number of fully developed pine nuts and kernels varies from 2 to 183 per cone, corresponding to a weight of 10 and 27 g [80,81]. There is a trend towards heavier seeds in drier climates [18]. However, climate change and Leptoglossus occidentalis have been causing a reduction in the number of fully developed kernels per pine nut [68]. Dispersal of seeds ranges from 0 to 10 m by gravity due to its weight [82], but distance may be increased by animals [83].

P. pinea is frequently managed in agroforestry systems, with fruit production as its main product [66,78]. The stands have lower density for fruit and higher density for timber-oriented stands [66]. Many stands are pure even-aged [73,84,85], whereas others are mixed even-aged [71,85] or pure or mixed uneven-aged [84,86,87]. Management of the stands with low density is related to their main product being fruit. Flowering and fruiting occur mainly in the outer crown, and thus the larger the crown surface the higher the fruit production [79,80,88]. Yet, there is wide variability in cone and seed production between trees and years [66,79,80,88,89,90,91,92], which is related to the climate of the three years between flowering and seed maturation [89]. In general, P. pinea presents one mast year in 3–6-year cycles [66]. The number of pine nuts per cone also has high variability within and between years and trees. This variability is related to the site, with a decrease in production linked to a decrease in precipitation and an increase in temperature [45,78,81]. The most frequent stand structure is high forest even-aged, with clearcut systems and artificial regeneration [84,93] through plantation of plants produced in containers [94] or with uniform shelterwood system [84], with rotations of about 100–120 years [84,93].

Natural regeneration of P. pinea is constrained by both dispersal limitation and water availability [95,96,97]. Relatively heavy seeds (not well-adapted to long-distance wind dispersal) result in regeneration concentrated close to parent trees unless animal dispersers extend dispersal distances or artificial regeneration complements the natural one, while successful establishment depends largely on favorable moisture conditions during the first growing season. Furthermore, cone harvesting may further constrain natural regeneration by reducing the availability of viable seeds. Consequently, natural regeneration is often patchy and irregular, requiring artificial regeneration to complement seedling recruitment in larger canopy openings [84]. The success of natural regeneration depends on years of good production and suitable precipitation. Uneven-aged stands are described in the literature as three to five cohorts [84,98] under irregular shelterwood or selection systems. These systems enable the perpetuity of the stand with more or less constant crown cover and production [98]. Mixed stands of P. pinea with other species (Quercus suber L. [99], Quercus rotundifolia Lam. [99,100], Quercus faginea Lam. [100], P. pinaster [71,101], and P. halepensis [93,102]) are also referenced.

Overall, the growth and reproductive performance of P. pinea are primarily driven by water availability, while stand structure affects the intensity of competition [71,73,74,87]. Silvicultural interventions such as thinning improve performance by reducing water competition rather than directly stimulating growth in both the stem and the crown [103,104]. In mixed stands, competition tends to be lower (due to niche separation), which enhances growth [71,87]. The number of thinnings varies according to the model of silviculture, and the main goal is to promote crown development to enhance cone production. Pruning is also prescribed to form a stem of about 2–4 m, as well as the control of spontaneous vegetation to reduce competition and fire hazards [98].

Table 3. Characteristics of Pinus pinea.

Indicator	Details	References
Ecological and silvicultural characteristics	Altitude up to 1000 m. Climate tolerance: Mean annual temperature 10–18 °C, minimum −2–7 °C, absolute minimum −19 °C, absolute maximum 41 °C. Annual precipitation 300–1500 mm or higher. Tolerances: Frost sensitivity: Winter low, spring high. Wind sensitivity: Low; with salt, low. Moderate sensitivity to drought. Sensitive to pests, diseases, and fire. Soil: Avoid excessively compact, with poor drainage and/or very calcareous. Ecology: Intolerant to shade. Longevity 150–300 years. Growth: Intermediate. Reproduction: Flowers occur March–May. Fruit maturation occurs in spring of the 4th year. Seed: Seed production per tree is variable, with an average of 1500 to 200 seeds per kg. Dispersal: Occurs via gravity and animals.	[5,45,105]
Silviculture	Managed as even-aged, pure high forest stands from natural or artificial regeneration. Uneven-aged and mixed stands are less common. Potential issues: Sensitive to pests, diseases, and fire.	[45].
Main goods and services	Products: Fruit, timber, biomass, and resin. Main uses: Edible kernels, biomass, protection. Suitable for forest restoration in sandy, nutrient-poor soils.	[5,45]
Diversity	International Union for Conservation of Nature (IUCN) status: Least Concern (LC), with wide distribution along the western Mediterranean. Genetic diversity is low. Key conservation strategies: Implementation of in situ conservation network.	[63,106]

Table 3. Characteristics of Pinus pinea.

Indicator	Details	References
Ecological and silvicultural characteristics	Altitude up to 1000 m. Climate tolerance: Mean annual temperature 10–18 °C, minimum −2–7 °C, absolute minimum −19 °C, absolute maximum 41 °C. Annual precipitation 300–1500 mm or higher. Tolerances: Frost sensitivity: Winter low, spring high. Wind sensitivity: Low; with salt, low. Moderate sensitivity to drought. Sensitive to pests, diseases, and fire. Soil: Avoid excessively compact, with poor drainage and/or very calcareous. Ecology: Intolerant to shade. Longevity 150–300 years. Growth: Intermediate. Reproduction: Flowers occur March–May. Fruit maturation occurs in spring of the 4th year. Seed: Seed production per tree is variable, with an average of 1500 to 200 seeds per kg. Dispersal: Occurs via gravity and animals.	[5,45,105]
Silviculture	Managed as even-aged, pure high forest stands from natural or artificial regeneration. Uneven-aged and mixed stands are less common. Potential issues: Sensitive to pests, diseases, and fire.	[45].
Main goods and services	Products: Fruit, timber, biomass, and resin. Main uses: Edible kernels, biomass, protection. Suitable for forest restoration in sandy, nutrient-poor soils.	[5,45]
Diversity	International Union for Conservation of Nature (IUCN) status: Least Concern (LC), with wide distribution along the western Mediterranean. Genetic diversity is low. Key conservation strategies: Implementation of in situ conservation network.	[63,106]

Potential issues are related to drought, forest fires, and pest and diseases affecting both trees and fruits [63,107]. A specific constraint to growth is drought. Post-drought growth is able to reach pre-drought levels if precipitation is suitable, which is related to the increase in water in the soil and a reduction in evaporation and evapotranspiration [72,108,109]. Repeated drought events might affect the species’ capacity for recovery due to physiological changes [71,72]. However, P. pinea show a trend towards being more affected by drought than Quercus ilex or Juniperus oxycedrus [110], but they are less affected than P. pinaster [111]. Changes in the climate with the increase in aridity likely drive P. pinea northwards and in altitude [112]. Yet, close to nature silviculture promoting uneven-aged structures through selection systems with light-intensity cuts showed a trend towards the maintenance of growth [86].

Three main products are identified in P. pinea stands [66], namely fruit, timber, and resin. The fruit is actually the main product of these systems due to its high nutritional value and market price [113,114,115]. Quantity and spatial and temporal variability of cone production [79,88,90,91,116] and pine nut and kernel efficiency per cone [80,90,91,113] influence the forest system’s sustainability. Manual cone harvest is labor-intensive and dangerous, and mechanical harvest can reduce harvest time and costs and improve labor security [11,117]. The profitability of umbrella pine stands managed for timber and fruit is related to the number of cones per tree and the cone market price [118].

P. pinea stands, both pure and mixed, are heterogeneous in dimension and spatial distribution of the trees and enable the germination and growth of a wide range of species in the understory and also provide services such as protection [119], conservation [120], recreation [121], and carbon storage and sequestration [122]. The factors that influence diversity are the number of species, which are related to the soil, the microclimate, and the stand structure. Understory vegetation increases (in number of species and density) with the decrease in crown cover, especially understory species intolerant to shade (including natural regeneration) [119,120,123]. Moreover, the soil and the microclimate create habitats suited to several species, creating different patterns of distribution [120].

4.4. Pinus halepensis (Aleppo Pine)

Aleppo pine (Table 4), native to the Mediterranean basin, is considered a pioneer plastic species [124]. It develops from 0 to 2000 m asl [45], reaching 1700 m asl in Morocco [1]. It is distributed in climates with mean annual temperatures of 11–23 °C [45,125] and mean annual precipitations of 350–700 mm, up to 1300 mm [125,126]. It has high frost sensitivity both in winter and spring, low sensitivity to wind and wind with salt [45], and high tolerance to drought [124,127], but it is susceptible to fires [126,128] due to its high flammability [129]. It is threatened by several pests, particularly the pine processionary moth (Thaumetopoea pityocampa Denis and Schiffermüller), bark beetles (Orthotomicus erosus Wollaston), and longhorn beetles (Monochamus galloprovincialis Olivier), among other, and diseases including pitch canker (Fusarium circinatum Nirenberg and O’Donnell) and canker (Crumenulopsis sororia (Karsten) Groves) [126]. It can develop in most soils, except for sandy soils with superficial water tables and very high humidity [45,105], withstanding calcareous conditions [105,130] and soils polluted by heavy metals [131,132]. It is a shade-intolerant species [45,105] with 100–150 years of longevity and high growth rates [124].

Table 4. Characteristics of Pinus halepensis.

Indicator	Details	References
Ecological and silvicultural characteristics	Altitude up to 2000 m. Climate tolerance: Mean annual temperature 11–19 °C, minimum −2–10 °C, absolute minimum −15 °C, absolute maximum 43 °C. Annual precipitation: 350–700 mm. Tolerances: Frost sensitivity: Winter, high; spring, high. Wind sensitivity: Low; with salt: low. Low sensitivity to drought. Sensitive to pests, diseases, and fire. Soil: Avoid sandy soils with superficial water table and very humid. Ecology: Intolerant to shade. Longevity 100–150 years. Growth: Intermediate. Propagation: Flowers occur March–May. Fruit maturation occurs in autumn. Seed: Produces 1.6–4 kg of seed per tree, with an average of 50,000 to 60,000 seeds per kg. Dispersal: Occurs via wind and animals.	[45,105,126,133]
Silviculture	Managed as even-aged, pure high forest stands, from natural or artificial regeneration. Mixed stands are less frequent and uneven-aged are scarce. Potential issues: Pests, diseases, fire, and allelopathy.	[93,126,133,134,135,136]
Main goods and services	Products: Timber, resin, edible nuts, and biomass. Main uses: Timber, resin, biomass, protection. Suitable for forest restoration in dry, sandy, nutrient-poor soils, especially calcareous polluted soils.	[131,132,133,137]
Diversity	International Union for Conservation of Nature (IUCN) status: Least Concern (LC), wide distribution along the western Mediterranean. Genetic diversity is geographically structured. Key conservation strategies: Implementation of in situ conservation networks, maintain local provenances, avoid genetic erosion in reforestations.	[137,138]

Flowering occurs in March–May and cones mature 3 years after flowering in autumn [45,137], and fruiting starts at 10–15 years [91]. It produces 1.6–4 kg of seed per tree and an average of 50,000–60,000 seed per kg [133]. The seed is light with a wing [137] with a dispersal up to 100 m [139]. This species is frugal, adapted to sites with poor soils [124,126] and seasonal and annual variability in precipitation, likely due to its well-developed root system [140] and resilience, particularly in adult trees [141]. Growth of P. halepensis is primarily controlled by water availability, with stand density determining the intensity of competition for this limiting resource. It decreases from sub-humid (0.95 m³ha⁻¹yr⁻¹) to arid (0.67 m³ha⁻¹yr⁻¹) sites, whereas it is higher in better fertility sites [133]. In general, growth increases with the decrease in hydric stress [142]. Inversely, in drought periods, growth decreases mainly due to competition for water (the higher the denser of the stand) [143], as well as fruit production [144]. Arid sites frequently have lower stand densities to reduce competition between trees [145].

It is frequently managed for timber and/or protection. High forest pure even-aged stands are frequent [93,133,146,147], along with mixed stands with P. pinea [145,148], P. pinaster [145,148], P. nigra [148], Quercus ilex Linnaeus [135], Quercus suber Linnaeus [93], and Juniperus spp. [149]. Uneven-aged stands are still being developed [136]. Many high forest even-aged stands often originate from natural regeneration and with high densities [150]. Thinning is an important silvicultural practice to enable growth and to reduce fire risk by decreasing crown continuity, both horizontal and vertical [147]. It is frequently associated with pruning to create vertical discontinuity [129]. Thinning methods vary and can result in a more or less regular tree spatial distribution or a clustered one [147]. Regardless of spatial distribution, there is an increase in diameter growth in the first two years post-thinning [147,151], declining 4–5 years after thinning [152], which indicates its periodical nature [129]. Spatial heterogeneity created by thinning in clusters [147] and other land uses further reduces fire risk [129]. Furthermore, in sites with periodical droughts, thinning increases diameter growth [153], and the growth rate is higher in thinning with heavy intensity than moderate intensity [154]. Moreover, the differences between provenances in terms of growth are small [155]. Admixtures of P. halepensis with P. pinaster, P. pinea, or P. nigra have lower yields than pure stands, which is due to the similarity of the pine species’ traits and the predominance of competitive interactions [148]. In arid conditions, mixed stands of P. halepensis with P. pinaster have higher yields than their pure counterparts, likely due to facilitation interactions. It has been observed that P. halepensis can affect the volume growth of P. pinea, but the former is not affected by the latter [148].

Natural regeneration is frequent, especially in stands after fires [150]. P. halepensis produces both non-serotinous and serotinous cones [156], with a higher proportion of the latter in stands prone to fire [157]. The serotinous cones form a seed bank in the trees [156,157], which enhances natural regeneration.

The rapid colonization ability of P. halepensis, for example, in Greece, alters successional trajectories in Mediterranean pine forests. High seed production, wider dispersal range, and rapid juvenile growth enable the species to outcompete slower-regenerating pines, such as P. pinea, following disturbance. As a result, repeated disturbances may gradually shift forest composition towards P. halepensis dominance, with consequences for associated plant and animal communities [82,158]. Besides competition for resources, P. halepensis may also influence community assembly through allelopathic interactions. The release of secondary metabolites can suppress germination and the early development of understory vegetation, potentially reinforcing pine dominance after establishment and reducing opportunities for the recruitment of competing species [159].

In stands dominated by P. halepensis, fire, in the short term, changes the composition and abundance of plant species [160], with an increase in non-native ones [161], but negatively affects ecosystem services [160]. After fire events, P. halepensis’s resilience is higher when the site is close to their climatic niche. Furthermore, the resilience of this species is higher than that of P. nigra [162]. Climate change with aridity increase is likely to reduce areas suitable for this species in Greece [163]. Moreover, aridity might lead to a reduction of the ecosystem functions and multifunctionality of these forest systems [164].

The high productivity and ecological plasticity of P. halepensis allow it to provide multiple ecosystem services (e.g., timber, resin, edible nuts, and biomass) simultaneously. Relatively fast growth (e.g., compared to P. sylvestris and P. nigra) [165] supports timber and biomass production, while extensive root systems stabilize degraded soils and reduce erosion. These characteristics, combined with high drought tolerance, explain the widespread use of the species in Mediterranean restoration programs. Nevertheless, increasing aridity and fire frequency may reduce the long-term capacity of these forests to maintain multiple ecosystem functions [124,127,165]. Its total volume production is quite variable, ranging from 66 to 251 m³ha⁻¹ in Tunisia, 108 to 350 m³ha⁻¹ in Algeria, and 114 to 285 m³ha⁻¹ in Spain [133]. It produces an average of 1.7–3.3 kg tree⁻¹y⁻¹ of resin that can be used to improve, e.g., biofuels, food preservation, and medicinal products [166]. Its kernels have high nutritional value and are used for human consumption, for example, in Tunisia. The production of cones is higher (in number and weight) in sub-humid than in semi-arid regions, with optimum seed production for a crown diameter of about 8 m [133]. Due to its high growth rate, it is able to store more biomass and thus sequester more carbon when compared with other pine species [165,167]. Many stands have a protection function. Stands from natural and artificial regeneration reduce erosion risk [168,169]. It can tolerate soils polluted with heavy metals (e.g., Zn, Pb, and Cd), supporting its use for phytostabilization [131,132].

P. halepensis stands host a range of herbaceous and shrub species, enhancing diversity [4,170], with more than 300 species in the understory [171]. The understory of uneven-aged stands bears higher diversity, likely due to gaps in the overstory, enhancing the development of vegetation compared to even-aged stands [170]. Diversity of plants in the understory increases with the decrease in the overstory crown cover [170,171], whereas the composition of the understory seems to be related to the overstory’s spatial distribution [171]. Yet, due to its dense regeneration [158] and allelopathic effects [159], it may change and/or restrict the development of the understory plants, thus altering the stand’s diversity. Thinning can promote management through a multi-objective approach, enhancing tree and understory resilience [156] and increasing understory vegetation density [171,172]. The effects of thinning and the control of spontaneous vegetation increase understory diversity, but these effects are temporary, decreasing from the second year after thinning onwards [172].

4.5. Pinus nigra (European Black Pine, Austrian Pine)

European black pine (Table 5) is a montane conifer that has a broad ecological range and exhibits significant morphological and genetic variability, being typically divided into six main subspecies: P. nigra subsp. salzmannii (Pirenean black pine), P. nigra subsp. nigra (Austrian pine), P. nigra subsp. laricio (Corsican/Calabrian black pine), P. nigra subsp. pallasiana (Crimerian/Turkish black pine), P. nigra subsp. dalmatica (Dalmatian black pine), and P. nigra subsp. mauretanica (African black pine) [65,173,174,175]. The species typically occurs from ≈350 m asl in Italy to ≈2200 m asl in the Taurus Mountains, with an optimal altitudinal range of 800–1500 m. However, this range varies considerably among sites and subspecies ([5,173,175] and references herein). It is well-adapted to temperate mountain climates, thriving in mean annual temperatures ranging from 9 to 13 °C [176], and it can tolerate extreme temperatures, withstanding absolute minimums as low as −28.8 °C [177]. Annual precipitation lies between 400 and 1000 mm depending on the subspecies [173], with the lower limit indicating its ability to endure relatively dry conditions compared to other montane conifers. It prefers deep (30–40 cm), well-drained, non-compacted soils, and, although it can grow on calcareous substrates, it tends to perform better in decarbonated soils. Its nutrient requirements are modest, making it suitable for marginal or degraded sites [5,176]. Biologically, the species is known for its exceptional longevity [173], often exceeding 400 years. Juvenile growth is relatively rapid; however, the species is strictly light-demanding [5], which has important implications for stand dynamics and silvicultural management. Flowering typically occurs in May, and cone development spans two growing seasons, with seed maturation taking place in October of the second year [173,177]. Seed production is around 30–40 seeds per cone [173]. Seed dispersal is primarily anemochorous (wind-dispersed) [61,178].

Table 5. Characteristics of Pinus nigra.

Indicator	Details	References
Ecological and silvicultural characteristics	Altitude range: 350–2200 m, with an optimal altitudinal range of 800–1500 m, though considerable variation is observed depending on the site/subspecies. Climate tolerance: Mean annual temperature 9–13 °C, minimum −5 °C, maximum 20 °C. Annual precipitation 400–1000 mm, although higher totals, between 650 and 2500 mm, are considered optimal, depending on the subspecies. Tolerances: No sensitivity to spring frost. Low sensitivity to wind and wind with salt. Low sensitivity to drought. Ecology: Intolerant to shading. Longevity 400 years. Early growth is fast.	[5,173,175,176,177]
Silviculture	Propagation: Flowers in May. Fruit maturation occurs in October. Age of fruiting is 10–20 years. Produces 30–40 seeds per cone. Dissemination occurs via wind and animals. Seed viability is good. Black pine is typically managed as even-aged, pure high forest stands. Potential Issues: Sensitive to pests and diseases.
Main goods and services	Products: Timber, biomass, and resin. Main uses: Timber, fuel, and protection. Suitable for high-altitude reforestation and soil protection in steep terrain. High frost and wind resistance make it appropriate for mountain restoration areas.	[5]
Diversity	IUCN status: Least Concern. High genetic structuring, with numerous distinct subspecies and provenances across Europe, often isolated.	[65]

P. nigra is usually managed as pure, even-aged high forest stands [176], but it can also be found in mixed stands with other pine species, particularly P. sylvestris or P. halepensis [173,179]. In Mediterranean countries, depending on management objectives and site conditions, it can be managed using various systems, including small-scale clearcutting (generally 1–3 hectares), uniform or irregular shelterwood systems, and selection cutting [180]. Stand management typically begins with pre-commercial thinning between 15 and 30 years of age, reducing stand density to approximately 1300–1500 trees per hectare. Subsequent selective thinning is implemented every 10–15 years until the end of the rotation period, typically 80–100 years, to promote the growth of future crop trees ([175] and references herein). A review by Vacek et al. [175] found that mature black pine stands averaged 334 ± 187 m³ ha⁻¹ in total standing volume, with a mean annual increment of about 4.9 ± 2.0 m² ha⁻¹ yr⁻¹. In addition to timber and biomass production (alongside resin extraction), it provides important ecosystem services, especially in erosion control on steep mountain slopes and in the rehabilitation and afforestation of degraded areas [5]. In several regions in Europe, forests dominated by P. nigra are included in the Natura 2000 network (habitat type 9530), highlighting their ecological significance and their role in supporting diverse plant and animal communities [179]. Due to climate warming, the distribution of black pine is expected to change significantly in the future, although this response will likely vary by geographic region. In Mediterranean areas, rising temperatures are linked to increased water stress, which can negatively impact the growth of this species. In contrast, central Europe is expected to experience improved climatic conditions, promoting the expansion of black pine in that region ([174] and references herein). The species is considered a candidate for assisted migration in dry forest sites of central Europe and as a substitute for indigenous conifers in future climate scenarios due to its adaptability [181].

The species is threatened by several factors, including pests, diseases, and wildfires. Major biotic threats include fungi like Dothistroma pini Hulbary and Sphaeropsis sapinea (Fries) Fuckel, the pine processionary caterpillar (Thaumetopoea pityocampa Denis and Schiffermüller), pine wilt disease caused by the pine wood nematode (Bursaphelenchus xylophilus (Steiner & Bührer) Nickle), and the bark beetle Ips pini (Say) ([174] and references herein). Large wildfires pose a significant risk, as well; while the species can tolerate low-intensity fires, it cannot survive severe wildfires. Its ability to regenerate relies on the survival of adult trees, which limits its capacity for rapid recovery [179]. For further details, see the review by Vacek et al. [175]. Recent evidence [162] further indicates that P. nigra shows relatively low post-fire resilience, particularly in stands located near the dry edge of its climatic niche.

4.6. Pinus mugo (Mountain Pine) and Pinus cembra (Swiss Stone Pine)

The mountain pine complex (P. mugo s.l. or P. mugo agg.) (Table 6) exhibits high variability and comprises numerous botanical subspecies. It includes the nominal subspecies P. mugo subsp. mugo, as well as other subspecies and their hybrids, representing the full spectrum of genetic and morphological variation within the species [182]. P. mugo is a European conifer complex with fragmented, “island-like” distribution across the main mountain systems of central and southern Europe. Its western occurrences include the Pyrenees and Iberian mountain ranges, extending through the French Massif Central, Vosges, Black Forest, Jura, and the Alps, continuing southeast into the Dinaric Alps. Eastern populations reach the Eastern and Southern Carpathians and the Rhodope Mountains of Bulgaria and Greece. Southern outposts occur in Spanish and central Italian mountains, while the northern natural boundary includes the Western Carpathians and parts of the Sudetes along the Czech–Slovak–Polish and Czech–German border regions [183,184]. The complex spans a wide altitudinal range from approximately 200 m in low mountain areas up to 2700 m asl, reaching its greatest density in exposed subalpine and alpine zones [184,185].

Morphologically, the complex displays considerable variation in growth form, branching patterns, and habit, reflecting adaptations to harsh climates, poor soils, and high-altitude exposure [184]. Some subspecies rarely exceed 0.5 m, while others may reach 20 m, adopting a tree-like form. The growth of P. mugo subsp. mugo is extremely slow, with seedlings increasing in height by only about 2–3 cm per year, ranking it among Europe’s slowest-growing trees. Its growth form is highly variable and typically irregular. The species usually develops as a low shrub, 2–3 m tall, with numerous stems and branches that spread close to the ground. It is a long-lived species, with a lifespan of 250–300 years. Depending on local conditions, it forms dense shrublands or small forest patches [185].

P. mugo subsp. mugo is fully frost-hardy, light-demanding, and undemanding with regard to soil. It occupies some of the most challenging habitats, including peat bog edges, rocky ridges, steep scree slopes, and limestone outcrops [185]. P. mugo subsp. Mugo’s extensive branched root system stabilizes loose soils and helps prevent erosion, avalanches, and landslides in mountainous areas. The wood, while hard and elastic, is suitable for fuel and crafting small items. The species is widely used in horticulture, with many cultivars planted for ornamental purposes. Its needles are rich in vitamin C and carotenoids, making them valuable for beverages that support the immune system and prevent scurvy, while cones and buds are processed into syrups, liquors, or essential oils with antioxidant properties [184,185]. It maintains overall genetic diversity, though isolated populations in southern and eastern Europe may be vulnerable. Its main threats are human activities, such as pasture expansion, ski area development, and fires, along with climate change, reducing suitable habitats. Fungal diseases, including white snow mold caused by Phacidium infestans P. Karst. and black snow mold caused by Herpotrichia juniperi (Duby) Petrak), affect needles but rarely cause tree mortality, while insect pests are not a major concern [184,185].

Table 6. Characteristics of Pinus mugo.

Indicator	Details	References
Ecological and silvicultural characteristics	Highly variable, including multiple subspecies and hybrids. It has a fragmented “island-like” distribution across central and southern European mountains. Altitude: From 200 m to 2700 m. Morphology: Diverse with tree or shrub habits. Tolerances: Frost-hardy. Soils: Tolerant of poor soils, occupying harsh habitats. Ecology: Intolerant to shade. Longevity: 250–300 years. Growth: Very slow and irregular growth form.	[183,184,185]
Silviculture	Forms dense shrublands or small forest patches. Highly resilient to harsh climates, frost, and poor soils. Root system stabilizes soil, reducing erosion, avalanches, and landslides. Minimal pest and disease issues; isolated populations may need monitoring.	[184,185]
Main goods and services	Products: Wood, biomass, protection. Main uses: Small crafts, firewood, horticultural value as ornamental cultivars, syrups, liquors, and essential oils. Ecological services include soil stabilization and erosion control.	[184,185]
Diversity	Maintains overall genetic diversity, though some isolated southern and eastern populations are vulnerable. Threats include human land use (pasture, ski development, fires) and climate change, reducing suitable habitats. Fungal diseases affect needles and roots but rarely cause mortality; insects are minor threats.	[184,185]

Table 6. Characteristics of Pinus mugo.

Indicator	Details	References
Ecological and silvicultural characteristics	Highly variable, including multiple subspecies and hybrids. It has a fragmented “island-like” distribution across central and southern European mountains. Altitude: From 200 m to 2700 m. Morphology: Diverse with tree or shrub habits. Tolerances: Frost-hardy. Soils: Tolerant of poor soils, occupying harsh habitats. Ecology: Intolerant to shade. Longevity: 250–300 years. Growth: Very slow and irregular growth form.	[183,184,185]
Silviculture	Forms dense shrublands or small forest patches. Highly resilient to harsh climates, frost, and poor soils. Root system stabilizes soil, reducing erosion, avalanches, and landslides. Minimal pest and disease issues; isolated populations may need monitoring.	[184,185]
Main goods and services	Products: Wood, biomass, protection. Main uses: Small crafts, firewood, horticultural value as ornamental cultivars, syrups, liquors, and essential oils. Ecological services include soil stabilization and erosion control.	[184,185]
Diversity	Maintains overall genetic diversity, though some isolated southern and eastern populations are vulnerable. Threats include human land use (pasture, ski development, fires) and climate change, reducing suitable habitats. Fungal diseases affect needles and roots but rarely cause mortality; insects are minor threats.	[184,185]

The Swiss stone pine (P. cembra) (Table 7) is an alpine timberline species with a focus on the inner alpine granite and gneiss landscapes. It occurs at altitudes between 1000 and 2300 m asl. Average annual temperature varies between 0 and 5 °C and annual rainfall between 500 and 2000 mm. As a timberline species, P. cembra is very robust and frost-tolerant, bearing minimum temperatures of −30 °C. It is a very long-lived tree species, with ages up to 1000 years. Moreover, stone pine grows on diverse soils, including poor and dry soils. Like all species found in the Alpine region today, P. cembra has had an eventful history. Repeated glacial expansion and retreat in central Europe meant that the area of Swiss stone pine was constantly changing. Fossil plant remains (pollen, macro-remains) prove that it probably survived the last ice age along the central and eastern southern edge of the Alps, in the Hungarian Plain, and around the Carpathians. The return to the Alps took place mainly westward into the Eastern Alps and north/westward into the Central and Western Alps. The Carpathians were mostly colonized by nearby populations [186]. This is also reflected in the spatial genetic structure, which reveals five genetic lines of kinship along the east–west axis of the Carpathians and the Alps. The eastern group is particularly dominant in the Carpathians and is thought to have originated mainly from refugia close to the Carpathians; it shows a genetic connection to the Eastern Alps. The three genetic groups in the Western Alps differentiated during the return migration after the last maximum glaciation. In Switzerland, two genetic groups dominate, corresponding to the two main areas in the east and west (Engadin, Valais). Knowledge of this spatial pattern makes it possible, e.g., to detect long-distance seed transfer or to select suitable seed (origin) [19].

The Swiss stone pine relies heavily on the dispersal of its seeds by animals [178], primarily the spotted nutcracker (Nucifraga caryocatactes). From late summer onwards, nutcrackers harvest the seeds as a key food resource and cache them in the soil for winter. A significant proportion of these seeds is not retrieved, thus contributing to pine regeneration. However, the caching sites are often in locations that are unsuitable for germination, meaning successful recruitment depends on the proportion of unrecovered seeds that end up in suitable locations [187]. Caches in forest openings or above the treeline, which are occasionally created, may become particularly important for establishment in the context of climate change, although natural regeneration can be slow and is strongly influenced by micro-site conditions.

Only selected micro-sites with small groups (three to five Swiss stone pines) should be planted. Favorable micro-sites are elevated areas and areas around old tree stumps or tree wood that has been left lying on the slope. During the first 20 years, Swiss stone pines are susceptible to fungal diseases (especially Gremmeniella shoot dieback), which spread under prolonged snow cover. This became very clear in the high-altitude reforestation project on the Stillberg (Davos, canton of Graubünden), where practically all Swiss stone pines in hollows and between avalanche barriers with long snow cover died due to Gremmeniella shoot dieback. Further afforestation failures are commented on by Fragnière et al. [188] from the Swiss canton of Fribourg. Planted Swiss stone pines are susceptible to damage by wildlife and are often swept away, knocked down, and gnawed at by hoofed game. Especially in areas where there are no old trees, the pressure from wildlife on this tree species is enormous. Reforestation without wildlife protection measures is therefore unlikely to be successful [189].

The Swiss stone pine prefers open forest structures. Although it is well-adapted to mountainous locations, it struggles when there are large numbers of trees, especially in the shade of faster-growing species such as Picea abies [12]. In order to preserve the Swiss stone pine even in the face of increasing competition, it should be promoted as early as possible through targeted young forest management or later through the removal of competing species. This will allow the Swiss stone pine to take on valuable mixed tree species functions even outside of its predicted future range should spruce come under increasing pressure from beetle damage in those areas. The following aspects should be taken into account when planting Swiss stone pines [12]: select suitable small sites, avoiding gullies and snow-covered hollows; plant in small groups with spaces between them; mix with other high-altitude tree species (e.g., P. mugo, Larix spp.); use regional planting material in pots, taking into account a possible shift in altitude under future climate conditions; where possible, grow plants at high altitudes (pre-adaptation); and consider planting above the current tree line.

The earliest evidence of the use of Swiss stone pine wood dates back to the Bronze Age (1400 BC). From the 16th century onwards, in addition to its use as firewood, for example, in salt works, a wide range of uses are known. The soft wood, which is easy to work with, was used to make everyday objects for mountain farming, such as bowls, milk utensils, and spoons. Due to its antibacterial and insect-repellent properties, Swiss stone pine wood was popular for furniture used to store goods (cupboards, flour chests). In addition to wood, forest litter was also used in certain regions for animal feed and bedding until the mid-20th century. This had serious consequences for the regeneration of Swiss stone pine: because the raw humus layer was removed, not only did the seedbed disappear but often also the seedlings developing in it. Out of concern for forest regeneration, the use of litter and cones was restricted or prohibited in many areas from 1900 onwards. In addition to the use of litter, regionally compensated hunting of nutcrackers, which continued into the 20th century, hampered natural regeneration. Today, more and more non-forestry products are being manufactured. In addition to these Swiss stone pine products, for which no figures on added value are available, the Swiss stone pine also has an idealistic and, above all, aesthetic value. The centuries-old, weather-beaten symbolic trees are admired, and richly structured Swiss stone pine or mixed Swiss stone pine forests are valued. This landscape quality is considered an important argument for the attractiveness of a tourist destination and thus for local added value [190].

In order to preserve Swiss stone pine forests as diverse habitats at the transition to the alpine zone under future climate conditions, an integrative approach is needed that promotes biodiversity and various forest functions and services, such as protection, use, and recreation. Local conditions and silvicultural history are important factors in assessing where and how Swiss stone pine should be promoted. Most of the genetic diversity of P. cembra is found within populations, with low differentiation between main regions, making the conservation of large, interfertile, and autochthonous populations essential for the species’ long-term adaptability and survival [191]. The two fungal pathogens Swiss stone pine snow fungus (Gremmenia infestans) and Gremmeniella shoot dieback (Gremmeniella abietina) play a decisive role in the regeneration dynamics of Swiss stone pine and influence its spatial distribution. Moreover, the diversity of mixed P. cembra stands can be at risk due to altitudinal shifts and, finally, range loss [12].

Table 7. Characteristics of Pinus cembra.

Indicator	Details	References
Ecological and silvicultural characteristics	High-altitude species (1000–2300 m), adapted to cold climates (0–5 °C; down to −30 °C), long-lived (up to 1000 years), growing on diverse, including poor and dry, soil. Light-demanding timberline species in the Central European Alps (typical mixture with Larix decidua and Picea abies); propagation by birds.	[186]
Silviculture	Strip and patch cut for mixed-stand regeneration; good regeneration in open forest structures.	[189]
Main goods and services	Protection forest; valuable solid timber for furniture and indoor utilization.	[190]
Diversity	Most diversity is found within populations; low differentiation between main regions; risk of altitudinal shift and range loss.	[12,191]

Table 7. Characteristics of Pinus cembra.

Indicator	Details	References
Ecological and silvicultural characteristics	High-altitude species (1000–2300 m), adapted to cold climates (0–5 °C; down to −30 °C), long-lived (up to 1000 years), growing on diverse, including poor and dry, soil. Light-demanding timberline species in the Central European Alps (typical mixture with Larix decidua and Picea abies); propagation by birds.	[186]
Silviculture	Strip and patch cut for mixed-stand regeneration; good regeneration in open forest structures.	[189]
Main goods and services	Protection forest; valuable solid timber for furniture and indoor utilization.	[190]
Diversity	Most diversity is found within populations; low differentiation between main regions; risk of altitudinal shift and range loss.	[12,191]

4.7. Pinus brutia (Turkish Pine), Pinus heldreichii (Bosnian Pine), and Pinus peuce (Macedonian Pine)

Turkish pine (Table 8) is a Mediterranean pine species closely related to P. halepensis, with which it is capable of natural hybridization [192,193]. The two species are frequently discussed together in the literature [126,194]. It is predominantly distributed along the eastern Mediterranean, with its main populations in Turkey, Crete, Cyprus, Syria, and Lebanon, and smaller, scattered occurrences in Iraq and Iran. Turkey alone hosts the vast majority of the species’ forests, accounting for approximately 3.8 million hectares of the more than 4 million hectares of total forest area [193,195]. The broad climatic tolerance of P. brutia underpins its ecological success across the eastern Mediterranean [193]. Although the species is well-adapted to seasonal drought, its optimum performance in relatively humid Mediterranean environments indicates that it combines drought tolerance with a strong capacity to exploit favorable moisture conditions [193,196]. This ecological strategy distinguishes it from the more xeric P. halepensis and partly explains their contrasting distributions across Mediterranean climatic gradients [193]. It typically occurs from 1400 m asl in the southern Mediterranean, although locally it can reach higher elevations, such as 1650 m asl in the Taurus Mountains of Turkey [126,194]. However, it occurs most optimally at elevations between 800 and 850 m asl [193]. P. brutia is a light-demanding and relatively fast-growing tree species, potentially reaching heights of 35 m and diameters at breast height of up to 150 cm. It achieves its best growth in humid climates where annual rainfall exceeds 900–1000 mm. P. brutia grows mainly on well-drained rocky and calcareous soils, especially rendzina over limestone or marl, and tolerates a wide range of soil types, except for poorly drained ones [193].

P. brutia is the most important conifer species in Turkey [195]. It has both high ecological and economic significance [185,197]. It has a mean annual increment of 2–3 m³ha⁻¹, with maximum yields reaching 12–15 m³ha⁻¹ [193]. Fire represents a major evolutionary driver of P. brutia’s ecology. Traits such as serotinous cones, abundant seed production, and rapid juvenile root development form an integrated regeneration strategy that enables rapid recolonization after disturbance. These adaptations explain both the persistence of the species under recurrent Mediterranean fire regimes and its widespread use in post-fire restoration [193].

Table 8. Characteristics of Pinus brutia.

Indicator	Details	References
Ecological and silvicultural characteristics	Mediterranean pine, related to P. halepensis (can hybridize); adapted to hot, dry summers and mild, wet winters; grows in semi-arid to humid zones; tolerates a wide temperature range (12–20 °C) and precipitation (400–2000 mm); prefers well-drained rocky/calcareous soils; elevation range sea level–1400 m (optimally 800–850 m); light-demanding, fast-growing; fire-adapted (serotinous cones, deep taproot, high seed production); culturally significant (ornamental, historical resin use in winemaking).	[126,185,192,193,196]
Silviculture	Naturally regenerates well (especially after fire); also regenerated by planting; even-aged systems (clearcutting, strip clearcutting) preferred; shelterwood applicable under certain conditions; used in afforestation, reforestation, soil protection, and windbreaks; mean annual increment 2–3 m3 ha−1, max 12–15 m3 ha−1.	[126,185,193]
Main goods and services	Timber: Construction, carpentry, industry, pulp, firewood; seeds: culinary uses; resin: traditional uses (e.g., winemaking); ornamental/landscaping; ecological services: soil protection, windbreak, reforestation.	[126,185,193]
Diversity	Generally stable genetic diversity; locally affected by ecological pressures (frequent wildfires, insect herbivores like Matsucoccus josephii and Thaumatopoea pityocampa, fungal pathogens like Crumenulopsis sororia); resilient to fire, but some populations vulnerable to disturbances.	[126,185,193]

Table 8. Characteristics of Pinus brutia.

Indicator	Details	References
Ecological and silvicultural characteristics	Mediterranean pine, related to P. halepensis (can hybridize); adapted to hot, dry summers and mild, wet winters; grows in semi-arid to humid zones; tolerates a wide temperature range (12–20 °C) and precipitation (400–2000 mm); prefers well-drained rocky/calcareous soils; elevation range sea level–1400 m (optimally 800–850 m); light-demanding, fast-growing; fire-adapted (serotinous cones, deep taproot, high seed production); culturally significant (ornamental, historical resin use in winemaking).	[126,185,192,193,196]
Silviculture	Naturally regenerates well (especially after fire); also regenerated by planting; even-aged systems (clearcutting, strip clearcutting) preferred; shelterwood applicable under certain conditions; used in afforestation, reforestation, soil protection, and windbreaks; mean annual increment 2–3 m3 ha−1, max 12–15 m3 ha−1.	[126,185,193]
Main goods and services	Timber: Construction, carpentry, industry, pulp, firewood; seeds: culinary uses; resin: traditional uses (e.g., winemaking); ornamental/landscaping; ecological services: soil protection, windbreak, reforestation.	[126,185,193]
Diversity	Generally stable genetic diversity; locally affected by ecological pressures (frequent wildfires, insect herbivores like Matsucoccus josephii and Thaumatopoea pityocampa, fungal pathogens like Crumenulopsis sororia); resilient to fire, but some populations vulnerable to disturbances.	[126,185,193]

The preference for even-aged silvicultural systems reflects the regeneration ecology of P. brutia. As a light-demanding pioneer species, successful seedling establishment depends on large canopy openings that provide sufficient light and reduce competition. Consequently, clearcutting and strip cutting generally mimic the natural post-disturbance conditions under which the species regenerates most successfully, whereas shelterwood systems are mainly appropriate where ecological constraints limit complete canopy removal [193]. The ecological plasticity of P. brutia allows the species to deliver multiple ecosystem services across environmentally heterogeneous landscapes. Its ability to establish on dry, rocky sites while maintaining relatively high productivity makes it particularly valuable for afforestation, watershed protection, and restoration of degraded Mediterranean ecosystems, where both timber production and protective functions are required. Its timber is highly versatile and is used in construction, carpentry, industry, pulp production, and as firewood, while its seeds are utilized in pastry-making. Their resin has a long history of use, including in Greek winemaking, and the tree is also valued as an ornamental species in landscaping [126,193,194].

Despite maintaining relatively high genetic diversity across most of its range, the adaptive capacity of local populations may increasingly depend on disturbance regimes. While the species is well-adapted to periodic wildfire, repeated fires combined with insect outbreaks and emerging pathogens may reduce regeneration success and alter long-term population dynamics under climate change [126,193,194].

Overall, P. brutia is a fire-adapted, light-demanding Mediterranean pioneer with broad ecological plasticity that enables successful establishment across a wide range of dry environments. Its persistence is supported by rapid post-fire regeneration, abundant seed production, and tolerance of nutrient-poor soils. However, increasing drought, recurrent high-severity wildfires, and emerging biotic threats may challenge its long-term resilience in parts of its range. Future management should therefore prioritize the use of natural regeneration where appropriate, maintaining stand structures that reduce fire hazard, and conserving genetic diversity to enhance adaptive capacity under climate change.

Pinus heldreichii, also known as Bosnian pine (Table 9), is an endemic and relict pine species from the Tertiary flora [198] native to the Balkans and a few isolated populations in the Apennine Peninsula.

Table 9. Characteristics of Pinus heldreichii.

Indicator	Details	References
Ecological and silvicultural characteristics	Endemic, relict Tertiary pine; native to the Balkans and parts of the Apennines; long-lived, slow-growing; occurs at 900–2500 m, reaching alpine timberline; prefers steep limestone/dolomite slopes but tolerates siliceous and ultrabasic soils; highly adapted to harsh mountain climates (cold winters, heavy snow, strong winds); frost-tolerant, pest- and pollution-resistant; main threats: forest fires, illegal logging, intensive grazing.	[138,198,199]
Silviculture	Occurs in mixed stands with silver fir and European beech but can form pure stands; enhances soil stabilization; used in high-altitude reforestation and afforestation and erosion control; reforestation usually by planting 3-year-old seedlings; strong colonizing potential; well-suited to dry, sunny, high-altitude sites.	[138,199]
Main goods and services	Primarily ornamental and landscape value; wood rarely exploited commercially but usable for general construction; contributes to ecosystem services such as soil stabilization, biodiversity support, and ecological resilience in mountain ecosystems.	[138]
Diversity	Populations fragmented across western and central–western Balkans and southern Italian Apennines; genetic diversity threatened by fragmentation and limited gene flow; some taxonomic uncertainty (possible distinction between P. heldreichii and P. leucodermis); listed as low-risk by IUCN but natural distribution highly restricted.	[138,199]

Exceptional longevity is a key component of the ecological strategy of P. heldreichii. Rather than relying on rapid growth or frequent regeneration, the species persists through long individual lifespans and tolerance of harsh environmental conditions, allowing populations to survive infrequent disturbance and prolonged climatic variability. The restriction of P. heldreichii to high-elevation (900–2500 m asl), rocky habitats (steep limestone, dolomite slopes) reflects its specialization for environments characterized by low temperatures, shallow soils (siliceous and ultrabasic bedrocks), and limited water availability [194]. These ecological conditions reduce competition from faster-growing tree species but also restrict opportunities for range expansion, making the species particularly sensitive to climatic changes that alter mountain ecosystems [199]. It performs well in dry, sunny environments, where it can reach heights up to 30 m and diameters up to 150 cm. It has a high frost tolerance and resistance to pests (including Diplodia tip blight caused by the fungal disease Diplodia pinea (Desm.) J. Kickx f. (syn. Sphaeropsis sapinea (Fr.) Dyko and Sutton) and atmospheric pollutants (such as sulphur dioxide, hydrogen fluoride, nitrogen dioxide, and ozone). The combination of cold tolerance, drought and pest resistance, and the ability to establish on nutrient-poor calcareous substrates enables P. heldreichii to occupy ecological niches unavailable to many other European conifers. The main threats are forest fires, illegal logging, and intensive grazing [138].

P. heldreichii typically occurs in mixed stands with silver fir (Abies alba) and European beech (Fagus sylvatica), but it can also form pure stands at higher elevations. Its tolerance of harsh mountain conditions, together with its capacity to establish on steep, rocky slopes, contributes to slope stabilization and erosion control, making the species an important component of mountain ecosystem functioning. Natural regeneration is often irregular in the harsh mountain environments occupied by P. heldreichii, making artificial regeneration an important management tool. Population persistence depends largely on successful seedling recruitment, which represents one of the weakest stages in the life cycle of the species. Thus, reforestations with P. heldreichii are typically done by planting (three-year-old seedlings). The primary value of Bosnian pine lies in ornamental and landscape applications rather than in timber production. Although its wood can be used for general construction, it is rarely exploited commercially. The species is widely employed in reforestation and high-altitude afforestation projects, as well as for erosion control in environmentally sensitive areas. Its combination of ecological resilience, pest resistance, and aesthetic qualities makes it a valuable component of both natural and managed mountain ecosystems [138].

The biology and ecology of P. heldreichii remain poorly studied [199]. Native populations occur in the western Balkan Peninsula (Albania and Greece), the central–western Balkans (Bosnia, Serbia, Montenegro, and Bulgaria), and the southern Italian Apennines (Monte Pollino). However, the taxonomic and phylogenetic relationships between the Italian and Balkan populations remain unresolved. As summarized in [138], differences in morphological traits, including tree height, bark characteristics, and twig and foliage color, have led to contrasting taxonomic interpretations. Some authors recognize P. leucodermis and P. heldreichii as distinct species, whereas other consider all populations to belong to a single species or treat P. leucodermis as a variety of P. heldreichii.

Bosnian pine is listed as low-risk on the IUCN Red List, but its natural distribution is highly limited and fragmented, mainly due to forest fires. In Italy, it occurs in only five small, isolated stands. This fragmentation reduces gene flow and increases genetic drift, putting at least three populations at risk of losing significant genetic diversity in future generations [138].

In summary, P. heldreichii is a long-lived, stress-tolerant mountain conifer adapted to the harsh environmental conditions of high-elevation Mediterranean ecosystems. Its tolerance of cold, drought, poor calcareous soils, and strong winds, together with its resistance to pests and air pollution, enable it to persist in habitats where few other tree species can thrive. Despite these adaptations, its naturally fragmented distribution, limited natural regeneration, forest fires, illegal logging, and intensive grazing threaten the long-term viability of many populations. Future management should therefore prioritize the conservation of genetically diverse populations, protection of natural regeneration through grazing control, continued use in restoration of degraded mountain habitats, and monitoring of population responses to climate change.

Macedonian pine (P. peuce) (Table 10) is a relict Balkan conifer specialized for cold, humid mountain environments. Its current distribution reflects the interaction between climatic constraints, topography, soil conditions, and historical biogeographic processes rather than altitude alone. The species occupies a broad altitudinal range, from about 600 to 2300 m asl [200,201], extending from the lower sub-mountain belt to the upper subalpine forest belt [202]. However, its ecological optimum lies between 1700 and 1900 m asl [203], where a combination of low temperatures, high atmospheric humidity, prolonged snow cover, and reduced competitive pressure from other tree species creates favorable conditions for persistence. At optimal sites, P. peuce can reach impressive dimensions, attaining heights of 35–40 m and diameters of 150 cm diameter at breast height [200]. In contrast, near the tree line and in harsher environments, growth is naturally constrained, and individuals may adopt shrub-like forms, reflecting the species’ ability to adjust morphology to severe climatic stress [202]. It is well-adapted to cold mountain climates characterized by long winters, short summers, and high air humidity. Its growth is positively influenced by high temperatures at the end of the previous growing season and negatively affected by cold winters. It tolerates heavy snowfall and exhibits relatively high resistance to severe winter cold and is hardy to temperatures between −23.3 °C and −28.8 °C. Although the species is generally considered tolerant of moisture variability and is not regarded as strongly drought sensitive, increasing summer water deficits, particularly during August, can negatively influence growth by reducing carbon assimilation during the active growing period [200]. This suggests that while P. peuce is well-adapted to cold climates, its future performance may increasingly depend on changes in seasonal water availability. Its ability to tolerate wind exposure and atmospheric pollution further contributes to its ecological resilience in exposed mountain environments [202].

Table 10. Characteristics of Pinus peuce.

Indicator	Details	References
Ecological and silvicultural characteristics	Altitudinal range: 600–2300 m; tolerates cold mountain climates, heavy snow, and frost (−23.3 °C to −28.8 °C); prefers humid conditions and light, humus-rich soils; slow early growth (“grass stage” 5–6 years); bark thin for ~30 years; Tertiary relict species with scattered native distribution in the Balkans; listed as Near Threatened.	[200,201,202,203,204]
Silviculture	Naturally forms pure or mixed stands (with Picea abies, P. sylvestris, P. mugo, Abies alba, P. nigra, P. heldreichii, Fagus sylvatica); intermediate shade tolerance; slow seedling establishment requires early competition control; used for afforestation on harsh terrain, erosion-prone sites, low-yielding stands, and forest restoration; can be regenerated naturally; no longer significantly exploited for timber in its native range; introduced in N. Europe and N. America, occasionally naturalized.	[200,201,202]
Main goods and services	Goods: Soft, light, durable wood (construction, furniture, carving), resin (chemical industry, optics, pharmacy, traditional medicine), ornamental uses. Services: Soil and water protection, biodiversity conservation, ecosystem stability, high-mountain forest restoration.	[200,201,202]
Diversity	High morphological and genetic variation: crown shape, branching, bark texture, strobili color; two main ecotypes (var. typica—west, var. vermiculata—east of Vardar River); hybridization potential with several pines; significant heritability and needle terpene variation among populations; adaptability to wide environmental conditions.	[202]

Soil relationships indicate substantial ecological plasticity but also reveal important regeneration constraints. In its native range, P. peuce grows mainly on soils derived from acid parent materials but also on serpentine, siliceous substrates and occasionally carbonate soils. Outside of the Balkans, the species demonstrates a broad edaphic tolerance, growing successfully on substrates ranging from flushed peats and podzols to sandy soils under poor- to medium-nutrient conditions [201]. This broad soil tolerance reflects physiological adaptability, although optimal development is generally associated with light, well-drained, humus-rich soils that provide favorable conditions for root development and nutrient acquisition. Poor performance on heavy, compacted, or strongly competitive grass-dominated soils indicates that regeneration is strongly influenced by microsite conditions and interspecific competition. The species’ deep and extensive root system represents an important functional adaptation, providing mechanical stability on steep mountain slopes, improving resistance to erosion, and allowing persistence under nutrient-poor and physically demanding conditions. P. peuce is relatively resistant to insect pests and fungal diseases, primarily due to the harsh high-mountain climate of its natural range and the high resin content of its tissues, though it is not entirely immune. As indicated above, the species shows high ecological adaptability [202]. It is a Tertiary relict species that survived major range contractions during the Pleistocene glaciations. Today, it is restricted to two disjunctive populations in the central Balkans, between 41° and 43° N, separated by the Vardar River valley; the western population is spread across Albania and Montenegro and the eastern population in western Bulgaria, with additional scattered occurrences in North Macedonia and northern Greece. It typically occupies north- and northwest-facing slopes near the timberline, where cold temperatures and high humidity prevail [201,202].

P. peuce is wind-pollinated, and its cones follow a two-year developmental cycle. Seed maturation occurs in September–October of the second year, followed by seed dispersal [200,202]. The species exhibits very slow early growth during its first five to six years (“grass stage”) before establishing strong vertical growth, which is believed to be under strong genetic control. The bark remains thin for up to 30 years, making young trees susceptible to deer browsing and fire damage [201].

Although it is one of the most valuable conifer species in the Balkans, P. peuce occupies a limited native area of about 30,000 ha. Due to overexploitation and habitat loss, is currently classified as Near Threatened and included in the IUCN Red List of Species [204]. P. peuce is no longer significantly exploited for timber production in its native range. However, it is a valuable species for high-mountain silviculture and used for afforestation on harsh terrain and erosion-prone sites, improving low-yielding stands and restoring or expanding degraded forest limits. It can also be regenerated naturally. Seedling establishment is slow and early competition control is essential to enable the transition from a prolonged grass stage to vertical growth. Introduced into northern Europe and North America, it has naturalized locally but is not considered invasive, although monitoring is recommended due to its broad site tolerance [200].

The species naturally forms pure stands and mixed stands, most commonly with Norway spruce (Picea abies), Scots pine (P. sylvestris), mountain pine (P. mugo), silver fir (Abies alba), black pine (P. nigra), Bosnian pine (P. heldreichii), and European beech (Fagus sylvatica). In mixed stands, P. peuce combines well with Picea abies or Abies alba, often forming two-storied structures that exploit complementary ecological and growth characteristics. Its intermediate shade tolerance allows it to compete successfully in such mixtures [201,202,203].

P. peuce wood is soft, light, of homogenous structure, thick, and durable. Historically, it was utilized for light construction, furniture, and woodcarving, while its resin was valued in the chemical industry, optics, pharmacy, and traditional medicine. In northern Europe and North America, it is cultivated as a timber tree in plantations and as ornamental. Ecologically, P. peuce is one of the most important conifers in the Balkans, as it provides a large number of ecosystem services, including soil and water protection and biodiversity conservation [202]. P. peuce exhibits considerable morphological and genetic variation across its range, including differences in crown shape, branching habits, bark texture, and strobili color. Two main ecotypes, var. typica and var. vermiculata, correspond to the western and eastern parts of the Vardar River. The species has successfully hybridized with several other pines (P. strobus, P. parviflora, P. monticola, P. ayacahuite, P. lambertiana, P. excelsa, and P. flexilis), producing vigorous seedlings with competitive growth and disease resistance. Genetic studies across Balkan populations indicate significant heritability and potential for selection, while needle terpene profiles reveal affinities among populations from Serbia, Montenegro, Greece, and Kosovo [202].

P. peuce is adapted to harsh mountain environments through high frost tolerance, resistance to snow and exposure, a deep root system, and a slow growth strategy that supports long-term persistence. However, its restricted distribution, fragmented populations, climate change, increasing drought stress, habitat degradation, and limited regeneration capacity represent major threats. Management priorities should focus on conserving genetic diversity, protecting high-elevation habitats, promoting natural regeneration, and maintaining resilient populations through adaptive forest management.

5. Cross-Species Synthesis and Management Implications

The ten pine species described in this article have some similar traits, like shade intolerance, can be cultivated in diverse soil conditions, and, in general, have timber of good quality. Yet, specific traits differentiate them, such as optimal climatic and altitudinal ranges. The information compiled from the literature reflects differences in regional conditions, management history, and site-specific factors, with some findings relying mainly on observational studies. Therefore, the patterns identified should be interpreted within the context of the variability inherent to the available evidence. Nevertheless, they provide a comparative overview of the characteristics reported among species. Based on this synthesis, Table 11 summarizes the main functional, ecological, and management attributes of the ten species analyzed, including shade tolerance, drought resistance, fire adaptation, growth rate, key goods and services, and major vulnerabilities. Their functional diversity offers opportunities to meet a wide range of management objectives—from timber production to protection and non-wood products—provided that species are matched to site conditions and emerging climatic risks.

Table 11. Overview of functional and silvicultural characteristics of ten native European pine species. Classes: Low (L), Medium (M), High (H).

Species	Biogeograph. Type	Altitude Range (m)	Drought Tolerance	Cold Tolerance	Fire Adaptation	Growth Rate	Goods and Services	Vulnerabilities
P. sylvestris	Boreal–continental	0–2000	M/H	H	L	M	Timber, protection	Drought, pests, fungi
P. pinaster	Mediterranean–Atlantic	0–800	M	M	H	H	Timber, resin	Pine wood nematode, fire
P. pinea	Mediterranean	0–600	M	M	M	M	Pine nuts, timber	Prolonged drought
P. halepensis	Mediterranean dry	0–600	H	L	H	H	Protection, timber	Invasiveness, fire
P. nigra	Sub-montane	600–1800	M	H	L	M	Timber, protection	Drought in southern range, pests, diseases, and wildfires
P. mugo	Alpine	1200–2400	M	H	M	L	Protection	Climate warming
P. cembra	Alpine	1500–2400	L	H	L	L	Protection	Slow regeneration
P. brutia	Eastern Mediterranean	0–800	M	L	M	H	Timber	Fire, drought
P. heldreichii	Balkan–alpine	800–2200	M	H	L	L	Protection	Fragmented populations
P. peuce	Balkan montane	1200–2200	M	H	L	M	Timber	Climate change

Note: The qualitative classes (Low, Medium, High) represent a literature-based synthesis of the predominant evidence reported for each species regarding drought tolerance, cold tolerance, fire adaptation, and growth rate. The categories are intended for comparative purposes and should not be interpreted as absolute or quantitative measures.

The qualitative classes assigned to drought tolerance, cold tolerance, fire adaptation, and growth rate represent a literature-based synthesis of the predominant evidence available for each species. These categories are intended to facilitate comparisons among species and should be interpreted as relative rather than absolute or quantitative measures. For simplification, the goods and services column highlights the principal products or functions most closely associated with each species (timber, resin, pine nuts, and protection). Broader ecosystem services, such as biodiversity conservation and recreational value, are recognized for many species but were not individually reported because they are common across most European pines.

The comparison across species reveals clear functional patterns along Europe’s environmental gradients. Mediterranean pines such as P. halepensis, P. pinaster, and P. brutia share a suite of drought-tolerant and fire-adaptive traits that allow them to cope with frequent disturbances. In contrast, alpine species like P. cembra, P. mugo, and P. heldreichii are characterized by slow growth, high cold tolerance, and a strong protective function in mountain landscapes. P. sylvestris and P. nigra occupy intermediate positions, combining broad ecological amplitude with increasing vulnerability to drought in their southern ranges. P. halepensis is one of the most drought-tolerant and fire-adapted Mediterranean pines, although it has also been reported as a potential invader in some non-native environments.

Regarding present-day distributions and expected scenarios, Mediterranean species may expand into areas that become warmer and drier, although increasing drought intensity and more frequent disturbances may reduce regeneration success at the southern margins of their distributions. Conversely, high-elevation species with restricted climatic niches are expected to experience habitat contraction as warming reduces the availability of cold environments, making them particularly vulnerable to future climate change. Species with broad ecological ranges, such as P. sylvestris and P. nigra, are likely to show contrasting regional responses, with declining suitability in southern Europe and persistence or expansion towards northern latitudes. Additionally, it is important to recognize that the current distributions of European pine species (Section 3) are dynamic, as climate change is anticipated to alter species’ ranges.

This synthesis reveals three broad functional groups of Mediterranean and European pine species. The first comprises fire-adapted pines, characterized by traits such as serotiny and rapid post-fire regeneration, which confer high resilience in fire-prone landscapes. The second includes high-mountain pines, long-lived and highly tolerant of cold conditions, which play a key role in slope stabilization, watershed protection, and the maintenance of other ecosystem functions in alpine environments. The third group encompasses ecologically versatile species with broad environmental tolerances and moderate phenotypic plasticity, making them suitable for a wide range of ecological conditions and management objectives. Although these groups simplify a continuum of ecological strategies, they provide a useful framework for understanding species’ responses to environmental change and for matching species selection and silvicultural practices to local site conditions under future climate scenarios. A comparative classification of climate change vulnerability and adaptive capacity was developed based on the individual species profiles and the cross-species synthesis (Supplementary Material, Table S2). Mediterranean fire-adapted species (P. halepensis, P. pinaster and P. brutia) generally demonstrate a higher level of adaptability thanks to their drought tolerance, fire-related characteristics, and regeneration strategies. However, they may still be susceptible to the effects of increasing aridity and more frequent disturbances. In contrast, high-elevation species (P. cembra, P. mugo, and P. heldreichii) are particularly vulnerable due to their restricted climatic niches, slow growth, and limited opportunities for upward migration. Broadly distributed species, such as P. sylvestris and P. nigra, exhibit more variable responses, with vulnerability depending heavily on regional climatic conditions and population context. Overall, adaptive capacity appears to be associated with a combination of ecological plasticity, genetic diversity, and functional traits that enable persistence under changing conditions. These patterns reinforce the importance of site-specific management approaches, including diversification, adaptive thinning, and the maintenance of structural heterogeneity to enhance forest resilience. In general, Mediterranean species are better suited to drought-prone and fire-prone environments, whereas alpine species remain appropriate for high-elevation protective forests. Broadly distributed species may provide greater management flexibility but require careful matching of provenances and site conditions, particularly in regions projected to experience increasing drought stress. Building on these findings, the evidence synthesized in this review highlights several priorities for adaptive forest management. Increasing stand diversity through mixtures with drought-tolerant broadleaved species can enhance resilience while reducing pest and disease risks. Silvicultural interventions, particularly thinning intensity and stand density, should be adjusted to alleviate water stress, especially in Mediterranean and continental regions where drought is expected to intensify. Increasing drought stress may also amplify the impacts of pests and diseases while interacting with more frequent and severe wildfires, reinforcing the need for integrated management strategies that simultaneously address these multiple disturbance agents. In landscapes facing increasing fire frequency, management should favor species with strong post-fire regeneration capacity, such as P. halepensis and P. pinaster, whereas extensive monocultures of more vulnerable species, particularly P. sylvestris in drought-prone areas, should be avoided. Maintaining genetic diversity and selecting seed sources and provenances better suited to projected climatic conditions will become increasingly important for enhancing adaptive capacity, maintaining forest productivity, and improving long-term resilience. Finally, conserving the protective functions of high-elevation forests, especially those dominated by P. cembra and P. heldreichii, should remain a priority given their importance for ecosystem stability and the delivery of critical ecosystem services.

Overall, the comparative framework presented here facilitates cross-species comparisons and supports decision making in adaptive forest management. Future meta-analyses using comparable datasets could further refine these comparisons through quantitative assessments of species’ responses.

6. Conclusions

The distribution of the ten European native pine species largely reflects differences in their ecological traits and environmental requirements. Variations in climatic and soil tolerances, regeneration strategies, and responses to disturbance help to explain both their present-day ranges and the silvicultural models followed across the continent. In turn, this ecological and management diversity is mirrored in the wide range of goods and ecosystem services supplied by pine forests across Europe well beyond timber production. While some pines are primarily managed for wood in pure or mixed stands under even- or uneven-aged systems (e.g., P. sylvestris and P. pinaster), others are chiefly valued for non-wood products, such as edible seeds (P. pinea), for combined production and ecological objectives (e.g., P. halepensis), or for their protective role in harsh mountain environments (e.g., P. heldreichii, P. cembra). In the context of ongoing climate change, such diversity represents a key resource for adaptive forest management. Aligning species (and, where appropriate, genetic material) with site conditions, disturbance regimes (including fires), and emerging threats, while preserving diversity in both composition and structure and applying suitable silvicultural systems, will be central to maintaining forest functions and the benefits they provide in the coming decades. From a management perspective, species selection should be aligned with local site conditions, climatic projections, and management objectives. Adaptive silvicultural practices, including diversification, appropriate thinning regimes, and the maintenance of structural heterogeneity, together with long-term monitoring of forest responses, will be essential to support resilient European pine forests. Knowledge gaps remain regarding the performance of different provenances and their responses to extreme climatic events. Particular attention should also be given to the conservation of genetic resources, especially for populations located at the warm and dry margins of species distributions, which may represent valuable reservoirs of adaptive diversity under future climatic conditions. For some species and ecological contexts, available evidence is still largely based on observational studies, highlighting the value of further experimental assessments to better support adaptive management strategies. Future quantitative analyses based on comparable datasets could also refine the understanding of species’ responses and management options. Overall, the comparative framework developed in this review can support evidence-based species selection, adaptive forest management, and forest policy decisions under changing climatic conditions.