Limited contribution of low-productive forests to green infrastructure along a temperate to boreal biogeographic gradient

Limited contribution of low-productive forests to green infrastructure along a temperate to boreal biogeographic gradient | Research Square window.SnipcartSettings = { analytics: { enabled: false } }; (function() { var accessVector = localStorage.getItem('access_vector') || ''; window.dataLayer = window.dataLayer || []; if (accessVector) { window.dataLayer.push({ user: { profile: { profileInfo: { snid: accessVector } } } }); } })(); (function(w,d,s,l,i){w[l]=w[l]||[];w[l].push({'gtm.start':new Date().getTime(),event:'gtm.js'});var f=d.getElementsByTagName(s)[0],j=d.createElement(s),dl=l!='dataLayer'?'&l='+l:'';j.async=true;j.src='https://www.googletagmanager.com/gtm.js?id='+i+dl;f.parentNode.insertBefore(j,f);})(window,document,'script','dataLayer','GTM-K279D39R'); Browse Preprints In Review Journals COVID-19 Preprints AJE Video Bytes Research Tools Research Promotion AJE Professional Editing AJE Rubriq About Preprint Platform In Review Editorial Policies Our Team Advisory Board Help Center Sign In Submit a Preprint Cite Share Download PDF Research Article Limited contribution of low-productive forests to green infrastructure along a temperate to boreal biogeographic gradient Bengt Gunnar Jonsson, Jakub Witold Bubnicki, Grzegorz Mikusiński, and 4 more This is a preprint; it has not been peer reviewed by a journal. https://doi.org/ 10.21203/rs.3.rs-9379486/v1 This work is licensed under a CC BY 4.0 License Status: Under Review Version 1 posted 6 You are reading this latest preprint version Abstract Context: Green infrastructure (GI) is an EU policy concept addressing functional connectivity to support biodiversity conservation and ecosystem services. In regions with significant forest fragmentation, establishing functional GI presents a challenge. Objectives We evaluate the contribution of low-productive forests (LPF) in networks of existing High Conservation Value Forests (HCVF) in Sweden, and discuss conservation implications for GI across different forest types and ecoregions. Methods Three characteristics for LPF were assessed: area extent of different forest types, patch size distribution, and role in functional landscape connectivity across five ecoregions. The primary data sources were a database of known HCVF and the national land cover data that includes data on LPF. Results LPF are dominant in the mountain region, both for all forests and among HCVF classified as low-productive. For other ecoregions, the extent of LPF was lower. Pine forests dominate LPF in all regions below the mountain region. For pine forests outside the mountain region, the inclusion of LPF increased the area of functional HCVF habitat for forest generalist species by 5–8% and for forest specialists by 6–16%. For other forest types, there was minimal improvement in connectivity. Overall, most LPF occur as small fragments, particularly when considering individual forest types. Conclusion LPF can strengthen GI in some regions and some forest types. However, they cannot replace the need for additional protection and restoration of productive forests to ensure functional connectivity. The role of LPFs as a contribution to conservation objectives is limited and related to species confined to LPFs. Biodiversity Connectivity Landscape planning Habitat loss Fragmentation Policy implementation Figures Figure 1 Figure 2 Figure 3 Figure 4 Introduction Green infrastructure (GI) is a European Union policy concept emphasizing the necessity of maintaining landscape-scale functional connectivity of representative habitat types to support biodiversity conservation and ecosystem services provisioning (EC 2013). In regions where natural forest habitats have been lost and fragmented due to historical and ongoing land use practices, establishing a functional GI presents a substantial challenge. Here, recent research (e.g., Orlikowska et al. 2020; Angelstam et al. 2020, 2026; Joks et al. 2023) indicates that areas designated for biodiversity conservation are mostly inadequate in quality, size and too fragmented to be ecologically functional. The prevailing commercially managed forest matrix often acts as resistance or barrier to functional connectivity (Villard and Haché 2012, Svensson et al. 2019, Angelstam et al. 2020, 2026, Mikkonen et al. 2023). However, within commercially managed forest landscapes, low-productive forest (LPF) stands excluded from management due to economic and regulatory limitations, may sustain structural and functional attributes typical for natural forests, such as old and dead trees, multilayered canopy and open and sun-exposed conditions (Sandström et al. 2020). From a GI perspective, LPFs may contribute naturalness attributes such as 1) characteristics of primary and undisturbed forests, 2) increasing heterogeneity of largely homogenous landscapes, 3) temporal continuity and 4) spatial connectivity among fragmented forest patches of significant conservation importance. While site productivity and species richness are commonly positively correlated as indicators of biodiversity value (Gillman and Wright 2006; Hämäläinen et al. 2024), LPFs provide valuable habitats for biodiversity and refugia for some forest species that are rare in managed forests (Uliczka and Angelstam 1999, Hämäläinen et al. 2020). This characteristic of LPFs is partially explained by longer temporal forest- and tree-continuity (e.g. Gustafsson et al. 2025). Forest productivity is primarily governed by water availability, soil properties, and climate. Interacting disturbance regimes further shape tree species composition, canopy structure, and spatial heterogeneity (e.g., Forrester et al. 2016). Productivity generally declines with increasing latitude and altitude, for example in tree-line transitions to alpine and tundra systems (Körner 2021). Although productive stands dominate temperate and boreal forest regions, local edaphic and topographic constraints, such as peat substrates, drought-prone soils, steep slopes, and shallow rocky soil, can limit growth, producing LPFs as discrete patches within otherwise productive forests. Sweden provides a compelling case study of challenges to conserve, manage and restore functional habitat networks as GI (Angelstam et al. 2020, 2026). With a predominant portion of the forests not providing favourable habitat conditions (Angelstam et al. 2020), securing the last remnants of natural habitats for biodiversity conservation remain a core challenge (Jonsson et al. 2019, Angelstam et al. 2026). However, in spite of current re-iterated policy targets, such as the Convention on Biological Diversity (CBD), the Aichi targets (CBD 2010) and the CBD Kunming-Montreal framework's third target on protected areas (CBD 2022), the functionality of GI is deemed inadequate in Sweden due to limited area protection and insufficient conservation management (Angelstam et al. 2020, Swedish Forest Agency 2022). To establish sufficient amounts of representative habitats with functional connectivity at a landscape and regional scale remains a major political and economic challenge (Angelstam et al. 2023, Wang et al. 2025, Svensson et al. 2026). Therefore, evaluating the potential of LPFs to support and strengthen habitat networks for various forest types is crucial. Out of Sweden’s total forest cover of 28 million hectares, approximately 23.5 million hectares are classified as productive, capable of supporting commercial forestry due to a volume growth potential at an annual rate of >1m 3 ha -1 over a 100-year rotation period. Conversely, the remaining 4.4 million hectares are categorized as low-productive forests (SLU 2025). Legislation currently restricts commercial forestry in LPFs (Swedish Forest Agency 2025). Hence, since the establishment of the present commercial rotation forestry system in the mid-20 th century, Swedish LPFs have essentially been set aside. Certain LPFs are identified as key Natura 2000 habitat types (CEC 1992) for conservation, including bog woodlands (91D0) and Fennoscandian deciduous swamp woods (9080). However, in the case of the predominant Natura 2000 habitat type, Western taiga (9010), the significance of LPFs lies in their potential capacity to strengthen the overall ecological functionality of habitat networks in managed forest landscapes. The primary objective of national environmental goals, EU directives, and CBD agreements is to halt biodiversity loss. This implies that conservation efforts must transcend mere quantitative measures, by also considering how functional GI can be established to ensure viable populations of species demanding higher levels of naturalness in the long term (Pressey et al. 2007). It is within this context that the role of LPFs should be evaluated, and especially their potential contribution to biodiversity conservation as elements of functional GI in different forest types and biogeographic regions. Therefore, to preserve functional GI through conservation, management, and restoration, the role of LPFs must be carefully examined. The objective of this study is to evaluate the potential role of LPFs in the network of both protected and non-protected High Conservation Value Forests (HCVF) as a contribution to GI functionality in Sweden. Our investigation is based on comprehensive national databases created using remote sensing and field surveys. Consequently, our research contributes to evaluating the on-the-ground effects of conservation policies concerning the role of LPFs in supporting functional green infrastructure across different forest types in different forest ecoregions. Study area The study area covers the entire land area of Sweden, which spans a north-south gradient of 1600 km from the temperate lowland ecoregion (nemoral) in the south to boreal and mountain ecoregions in the north (Ahti et al. 1968). Forests is the dominating (69%) land cover, with significant regional variations in main forest types (SLU 2025). For regional analysis, we have applied the five biogeographical ecoregions commonly used for official forest statistics in Sweden (SCB 2025, Figure 1), based on biogeographic conditions, main landscape characteristics. Forestry practices focused on wood production have a long history in Sweden. While substantial forest areas were converted to agricultural land during Millenia in southern Sweden (Berglund 1992), since the 13th century, the mining industry has impacted forest structure in south-central ecoregions. Large-scale industrial forestry, however, began in the early 1800s in southwestern Sweden (Bladh 1995; Angelstam et al. 2013), followed by a "timber frontier" expanding into northern Sweden from the late 1800s (Östlund et al. 1997). Concerns regarding sustained yield forestry in the early 1900s led to a management system involving rotation forestry based on clear-cut harvesting, soil scarification, artificial planting, pre-commercial cleaning and commercial thinning (Angelstam et al. 2022), which has been fully implemented since the mid-1900s (Naumborg 2024). Concurrently, significant areas of bogs (approximately 1.5 million ha) were drained to encourage wood production. At present, around 200,000 ha (1%) of Swedish forests undergo clear-cut annually, with an additional 200,000 ha being subject to commercial and pre-commercial thinning (SLU 2025). Currently, formally protected areas on productive land contribute most in terms of habitat quality (Kyaschenko et al. 2022), and these cover 6.3 % of Sweden’s productive forest land, and 9.4% of all forest land (Statistics Sweden 2025). However, there is a stark contrast in the proportion of protected areas between the remote mountain forests in northwestern Sweden (59.1%, all forest land), and the rest of the country (4.2%; ibid.) where wood production dominates. In addition, 5.9% is voluntarily set aside on productive forest land largely due to the widespread adoption of forest certification schemes such as FSC and PEFC (ibid.). While LPFs cover 16% of Sweden’s total forest area, the majority (11%) are located outside formally protected areas (SCB 2025). Methods To evaluate the significance of LPFs, three characteristics were assessed for each of the five ecoregions: area extent of different forest types, distribution of patch size, and role in functional landscape connectivity. Area of different forest types The primary source utilized for delineating various forest types was the national land cover data of Sweden (NMD; Swedish EPA 2019) generated during the period 2017-2019. This comprehensive, raster based, land cover map covers the entire country, mapping various forest types with a minimum mapping unit of 0.01 hectares. To detect LPFs, supplementary land cover data on productivity (Keskitalo et al. 2019) was employed. This LPF dataset, a primary source of data for our analysis, integrates satellite data (Sentinel-2), laser scanning, soil characteristics, moisture content, and climate. Validation assessment (at a 20-meter radius scale) was conducted using data from the National Forest Inventory (40,100 plots) and the National Inventory of the Landscape in Sweden (2,200 plots; Keskitalo et al. 2019), revealing high precision in identifying productive forests (99%) and somewhat lower accuracy for LPFs (75%) when considering the predominant land cover class within the 20-meter plot (Nilsson et al. 2020). It is important to highlight that certain large forest owners define LPFs from a management perspective and incorporate technical management constraints such as steep terrain, based on their internal stand level inventories. Consequently, the actual areal extent of LPFs excluded from logging activities may differ from the estimates obtained from the NMD. The initial NMD land cover classes encompasses seven forest types occurring on mineral and wet soils (totalling 14 combinations). To simplify while retaining biological significance, the two soil types were combined across all forest types. Norway spruce ( Picea abies ) forest and mixed conifer forests were grouped into one category, as were hardwood and mixed hardwood-deciduous forests (only present in the southern ecoregions), thus generating five distinct forest types; 1) deciduous (mainly birch dominated), 2) hardwood (dominated by broadleaf tree species with a southern distribution), 3) mixed deciduous/coniferous, 4) Scots pine ( Pinus sylvestris ), 5) spruce and mixed coniferous. Statistical analyses and data visualizations were performed using GRASS GIS 8.4 (GRASS Development Team, 2025) and custom Python 3.10 scripts. Two publicly available national spatial datasets served as input: 1) a compilation of High Conservation Value Forests (HCV forests) in Sweden, produced by the Swedish Environmental Protection Agency (Swedish EPA 2016 and subsequent updates); and 2) the National Land Cover Data (NMD), also produced by the Swedish Environmental Protection Agency (Swedish EPA 2019). The NMD dataset includes the LPF layer as described above. The density of LPF was visualized by implementing a 100 km 2 moving window across Sweden. Patch size In terms of patch size, the distribution of LPFs was evaluated based on four size-categories: 50-500 ha, >500 ha. A threshold was employed to delineate LPFs at a 1 ha pixel scale, selecting pixels with total LPF cover above 50%, i.e., hectare pixels dominated by LPF. Patches were defined as connected LPF pixels (8-neighbours connectivity), with patch size computed for each patch. Functional connectivity To assess the potential impact of low-productive forests on functional connectivity, we calculated how their addition to existing HCVF enhances habitat network functionality. The functional connectivity of HCVF, both with and without the addition LPF, was assessed using the algorithm outlined in Angelstam et al. (2020). Spatial modelling parameters were set to represent two virtual species with differing habitat requirements i.e. with higher and lower demands in respect to HCVF availability (Table 1). Forest patches at 1 ha scale were delineated using the same rules as for the patch size analysis and further defined by buffering HCVF- pixels by 100 m (200 m distance between patches) for specialist species and by 200 m (400 m distance between patches) for generalist species. Patches meeting minimum size criteria (50 and 5 ha respectively for specialist and generalist species) were selected, using moving window sizes of 2x2 km (specialist species) and 5x5 km (generalist species) to represent the landscape scale. A minimum habitat availability threshold of 20% was applied to identify functional HCVF networks at the landscape scale (cf. Hanski 2011; Svancara et al. 2005). The outcome was a mask of interconnected functional areas derived through the aforementioned steps. The actual area was calculated as the sum of all pixels from the original layer intersecting with this mask. This process was repeated for all forest types and ecoregions under two scenarios: 1) HCVF only and 2) combined HCVF and LPF. The results are visualized as the distribution of hotspots and coldspots based on the distance to the nearest functional patch, and hence show habitat network functionality across Sweden. Table 1 Characteristics of the two virtual species used in the analysis of functional connectivity. The threshold for habitat availability at landscape level is set to 20% for both generalist and specialist species Buffer size Patch size limit Landscape size Generalist species 100 m 5 ha 5x5 km Specialist species 200 m 50 ha 2x2 km Results Area of different forest types LPFs in Sweden are primarily concentrated to the mountain region with 58% of all forests and 77% of HCVF classified as LPF (Table 2). For the other ecoregions, the fraction of LPFs is substantially lower, varying from 19% and 11% in the north boreal, 12% and 10% in south boreal SE, 8% and 9% in hemiboreal, and 6% and 7% in the nemoral ecoregions. Pine is the dominant forest type among LPFs in all forest ecoregions below the mountain region, while deciduous forest dominates in the mountain region. A series of maps in Figure 2 illustrate the density of different LPF types. Table 2 Total area of LPF and its fraction (%) of the individual forest types and out of delineated HCV forests in the five Swedish forest ecoregions. NA denotes a forest type missing in the region Region Forest type LPF area (kha) % of forest type % of HCVF Mountain Deciduous 837.8 94 69 Mountain Hardwood NA NA NA Mountain Mixed 183.0 48 87 Mountain Pine 185.8 49 87 Mountain Spruce & mixed 220.0 27 92 Mountain All 1426.6 58 77 North boreal Deciduous 140.6 25 9 North boreal Hardwood NA NA NA North boreal Mixed 200.8 19 10 North boreal Pine 513.3 21 10 North boreal Spruce & mixed 152.1 12 17 North boreal All 1006.9 19 11 South boreal Deciduous 82.3 15 10 South boreal Hardwood 0.4 2 12 South boreal Mixed 147.1 13 10 South boreal Pine 549.8 17 9 South boreal Spruce & mixed 112.7 4 14 South boreal All 892.2 12 10 Hemiboreal Deciduous 72.6 11 8 Hemiboreal Hardwood 16.5 4 4 Hemiboreal Mixed 41.5 6 6 Hemiboreal Pine 309.4 15 10 Hemiboreal Spruce & mixed 33.1 2 7 Hemiboreal All 473.1 8 9 Nemoral Deciduous 10.4 9 10 Nemoral Hardwood 4.4 2 14 Nemoral Mixed 6.7 6 6 Nemoral Pine 20.5 15 5 Nemoral Spruce & mixed 3.1 1 4 Nemoral All 45.1 6 7 Patch size Habitat patch size plays a crucial role in assessing habitat network functionality. Small to intermediate sized patches dominate LPFs across all forest ecoregions, except for the mountain region where almost 59% of all LPFs are located in patches exceeding 500 hectares. For the other forest ecoregions, the two smallest patch size intervals strongly dominate (Figure 3). For the individual forest types, pine forest and deciduous forest are relatively well represented in the >5-50 ha class while other forest types are mainly restricted to patches less than 5 ha (Table 3). Table 3 Patch size distribution of LPF per region, forest type and patch size class for hectare pixels with more than 50% LPF Region Forest type % 500 ha Mountain Deciduous 8.6 15.4 19.7 56.4 Mountain Hardwood 0 0 0 0 Mountain Mixed 47.6 40.7 10.6 1 Mountain Pine 33.3 41 19.8 5.8 Mountain Spruce & mixed 30.8 37.9 30.3 1 Mountain All 7.9 13.6 19.8 58.7 North boreal Deciduous 64.8 28.8 6.4 0 North boreal Hardwood 0 0 0 0 North boreal Mixed 77.4 22 0.6 0 North boreal Pine 53.9 40.8 5.3 0 North boreal Spruce & mixed 77.4 20.3 2.3 0 North boreal All 42.7 45.7 11.3 0.2 South boreal Deciduous 80 16.1 3.9 0 South boreal Hardwood 100 0 0 0 South boreal Mixed 88.9 11.1 0 0 South boreal Pine 61.9 34.7 3.4 0 South boreal Spruce & mixed 88.5 11.5 0 0 South boreal All 54.9 39.8 5.2 0.2 Hemiboreal Deciduous 75.2 24.8 0 0 Hemiboreal Hardwood 100 0 0 0 Hemiboreal Mixed 96.6 3.4 0 0 Hemiboreal Pine 56.6 39.2 3.8 0.5 Hemiboreal Spruce & mixed 90 10 0 0 Hemiboreal All 56.3 39.1 4.2 0.4 Nemoral Deciduous 71 29 0 0 Nemoral Hardwood 100 0 0 0 Nemoral Mixed 88.5 11.5 0 0 Nemoral Pine 65.5 34.5 0 0 Nemoral Spruce & mixed 68 32 0 0 Nemoral All 64.2 35.8 0 0 Functional connectivity The analysis of functional connectivity shows substantial differences in the contribution of LPFs in improving habitat network functionality across different forest types, ecoregions in Sweden, and for less demanding generalist versus more demanding specialist species (Table 4). Table 4 The addition in percent units of LPFs area to existing HCVF in enhancing functionality of habitat networks for different forest types and forest ecoregions. G = generalist species; S = specialist species. NA denotes that the forest type is missing in the region All forest Pine Spruce Mixed Deciduous Hardwood Region G S G S G S G S G S G S Mountain 0 1 0 1 0 0 0 0 1 1 NA NA North boreal 6 15 6 11 1 1 2 3 1 1 NA NA South boreal 8 16 8 13 1 2 0 1 1 0 0 0 Hemiboreal 6 14 8 16 0 1 1 0 1 1 0 0 Nemoral 2 3 5 6 1 0 0 0 0 0 0 0 Given the high density of HCVF and its large overlap with LPF in the mountain region, the inclusion of LPF provides very limited additional GI functionality across forest types. For pine forests below the mountain region, the inclusion of LPFs led to an increase of 5-8% in area of functional HCVF habitat for forest generalist species across ecoregions. Adding LPFs for pine forest specialists showed a higher increase, with figures ranging from 6-16%, notably in the south boreal and hemiboreal ecoregions with 13% and 16% increased connectivity, respectively. The contribution of pine forest to functional connectivity is clearly reflected in the patterns for all forest land although the fraction of connected habitats is obviously larger when including all forest types. For the other four forest types, there is minimal to no improvement in functional connectivity by incorporating LPFs, with figures ranging from 0-3%. The improved connectivity for all forest types and for pine forests caused by adding LPFs is illustrated by the hotspot-coldspot maps in Figure 4 (for the other forest types see Appendix 1). Discussion In forest-dominated landscapes, effective green infrastructure requires functional HCVF-habitat networks. In this study, the contribution of LPFs to habitat functionality was assessed based on their occurrence across Sweden, their patch size distribution and in relation to functional habitat networks of two virtual species representing different network demands. For all these three aspects the contribution of LPFs varied strongly between different forest types and different ecoregions. In the mountain region, LPFs dominates the landscape but given the high share of already identified HCVF forests their additional contribution to habitat functionality is small. Outside the mountain region it is mainly in pine forests that LPFs provide improved connectivity. This is logical since pine forests represent the dominant LPF type outside the mountain region due to the ability of pine to thrive on very poor site types on both dry mineral (rocky outcrops) and mesic to wet mineral and peat organic soils. The areal extent of LPFs in Sweden follows a clear south – northwest gradient mainly reflecting macroclimatic conditions. The vast majority of the LPFs occur in the mountain and boreal ecoregions. Although they do occur in both the hemiboreal region and nemoral region they represent less than 10% of the forest area. Hence, the role of LPFs for improving forest GI varies between biogeographical ecoregions and dominant tree species (forest types). For forests dominated by pine in the mountain ecoregion, parts of the boreal ecoregions and in the eastern part of the hemiboreal ecoregions, they do however contribute to GI functionality by increasing the functional connectivity of forest patches (Fig. 4 ). For other ecoregions, LPFs derive much of their ecological importance as isolated patches providing habitat for species specialised on low-productive forest conditions with natural structural characteristics (Hämäläinen et al. 2020 ). It should be noted, however, that in the mountain forest region most of the LFP are already identified as HCVF and included in functional networks (Svensson et al. 2020 ). The size distribution of LPFs varies across the south-north gradient. Overall, most of the LPFs occur as small fragments (< 5 ha), particularly when considering individual forest types. Only for deciduous forest in the mountain region, larger landscapes were dominated by LPFs, a pattern identified in earlier studies (Angelstam et al. 2020 , 2026 ; Mikusiński et al. 2021 ). In contrast, and especially for the more southern ecoregions, most LPFs occurred in fragments smaller than 5 ha. This fragmentation is natural and primarily caused by variations in local forest site productivity, rather than habitat loss caused by human activity (e.g. Svensson et al. 2019 ). Hence, it is likely that some species with very small area requirements may have favourable conservation status as long as edge effects and climate change remain limited. The analysis of functional connectivity further emphasizes a relatively limited contribution of LPFs. Obviously, the chosen examples of virtual species do not reflect the full gradient of habitat demands of all forest species. However, they do represent relevant examples of habitat demands and landscape connectivity and a commonly used threshold (20%) for amount of habitat at landscape scale (Hanski 2011 ; Svancara et al. 2005 ). It may look contradictory that the increase in connectivity by adding LPFs to the network was higher for specialist species, but it should be noted that that total area of functional networks is consistently higher for generalist species. It is also clear from the analysis that the contribution of LPFs is only evident for forests in general and basically mirrors the contribution of for pine dominated LPFs. For all other forest types, the contribution is negligible. While this study addresses how LPFs contribute to GI, there are several other potential contributions to GI functionality, including formally protected areas, voluntary set asides and nature considerations applied during forest operations through retention practices. Although compilations of the area extents of these categories exists, as their levels of naturalness and other conservation qualities differ one cannot simply add the total areas. Thus, the extent to which conservation targets are met depends on not only on quantities (Angelstam et al. 2020 , 2026 ) as an area’s effectiveness depends on i) quality in terms of degree of naturalness (Puettman et al. 2012; Winter 2012), ii) patch size related to species requirement (Angelstam et al. 2004; Wiens and Milne 1989), iii) functional connectivity allowing species to move between suitable habitats in the landscape (Pither et al. 2023), and iv) longevity in relation to natural disturbances and succession dynamics (Rosenvald and Lõhmus 2023). In the light of the agreed policy targets for protected areas and with their specification to be particularly important for biodiversity through securing functional networks, we question whether LPFs by default should be seen as a contribution to fulfilling conservation targets albeit being legally excluded from commercial forest management, particularly if only their area contribution is considered. In local landscapes and for species with small area requirements, however, LPFs may contribute to functional GI, but here we argue that including locally-relevant quality parameters on top of area is of key importance. In this context, even small pine-dominated LPFs may play an important role as restoration nodes, facilitating more effective recolonisation of restored habitats by specialised species (Mikusiński et al. 2021 ; Ramberg et al. 2026 ). This illustrates the importance of not just considering the area proportion protected to fulfil the intention of the international agreements, but also the particular roles of habitat quality, habitat patch size and functional connectivity for biodiversity conservation (cf. CBD 2022 ). Green infrastructure, like conservation networks more broadly, should be ecologically representative of the ecoregions in which it is implemented (Pressey et al. 2007 ). LPFs in Sweden are not fully representative for all forest types, both for natural reasons—because they encompass only a narrow segment of the forest productivity gradient—and for socio-economic reasons, as they have often been protected due to their comparatively low economic value. This skewed representation, constrains their broader contribution to forest biodiversity conservation in Sweden. Still, naturally multilayered pine forests, the main LPF type, have become rare due to intensive forest management and fire suppression (Berglund and Kuuluvainen 2021 ). Recent research suggests that the historical extent of such forests in northern European dynamic forest landscapes was greater than previously assumed (ibid.). This implies substantial restoration needs. To conclude, our analysis show that although LPFs may strengthen GI in some ecoregions and for some forest types they cannot replace the need for additional protection and restoration of productive forests with high conservation value to ensure functional connectivity. Hence, the role of LPFs as a contribution to national and international conservation objectives is limited and mainly related to species that are confined to these island habitats. Yet they contribute to landscape heterogeneity and may hold aesthetic and recreational values. Declarations Competing Interests The authors have no relevant financial or non-financial interests to disclose Funding This work was supported by Swedish Environmental Protection Agency, grant NV-03728–17, to Bengt Gunnar Jonsson, Horizon 2020 Research and Innovation Programme Grant Agreement No. 101036849 to Johan Svensson and Formas grant 2022–02146 to Per Angelstam Author Contribution All authors contributed to the study conception and design. Material preparation and analysis were performed by Jakub Bubnicki. The first draft of the manuscript was written by Bengt Gunnar Jonsson and all authors commented on previous versions of the manuscript. All authors read and approved the final manuscript Acknowledgments This work was supported by Swedish Environmental Protection Agency, grant NV-03728–17, to Bengt Gunnar Jonsson, Formas Research Council, grant 2022–02146_Formas to Per Angelstam and Horizon 2020 Research Innovation Programme (SUPERB: Upscaling Forest Restoration—SUPERB (forest-restoration.eu)); Grant Agreement No. 101036849 to Johan Svensson. References Ahti T, Hämet-Ahti L, Jalas, J (1968) Vegetation zones and their sections in northwestern Europe. Annales Botanici Fennici 5:69–211 Angelstam P, Andersson M, Isacson DV et al (2013) Learning about the history of landscape use for the future: consequences for ecological and social systems in Swedish Bergslagen. Ambio 42:150–163. https://doi.org/10.1007/s13280-012-0369-z Angelstam P, Asplund B, Bastian O et al (2022) Tradition as asset or burden for transitions from forests as cropping systems to multifunctional forest landscapes: Sweden as a case study. Forest Ecology and Management 505:119895 https://doi.org/10.1016/j.foreco.2021.119895 Angelstam P, Bush T, Manton M (2023) Challenges and solutions for forest biodiversity conservation in Sweden: Assessment of policy, implementation outputs, and consequences. Land 12:1098. https://doi.org/10.3390/land12051098 Angelstam P, Manton M, Green M, Jonsson BG, Mikusiński G, Svensson J, Sabatini FM (2020) Sweden does not meet agreed national and international forest biodiversity targets: A call for adaptive landscape planning. Landscape and Urban Planning 202:103838. https://doi.org/10.1016/j.landurbplan.2020.103838 Angelstam P, Yamelynets T, Manton M, Bubnicki JW, Jonsson BG, Mikusinski G, Svensson J, Dawson L (2026) Environmental history determines forest habitat network functionality: The need for landscape planning in Sweden. Ambio. https://doi.org/10.1007/s13280-026-02353-7 Berglund B (1992) Landscape reconstructions in South Sweden for the past 6000 years. Proceeding of the British Academy 19:25-37 Berglund H, Kuuluvainen T (2021) Representative boreal forest habitats in northern Europe, and a revised model for ecosystem management and biodiversity conservation. Ambio 50:1003–1017. https://doi.org/10.1007/s13280-020-01444-3 Bladh G (1995) Finnskogens landskap och människor under fyra sekler. Forskningsrapport 95:11 Högskolan i Karlstad CBD (2010) Strategic plan for biodiversity 2011–2020 and the Aichi targets. Convention on Biological Diversity, Montreal. Available from https://www.cbd.int/sp/targets/(accessed February 2026) CBD (2022) Kunming-Montreal Global Biodiversity Framework. Convention of Biological https://www.cbd.int/gbf (Accessed February 2026) CEC (1992) Council Directive 92/43/EEC of 21 730 May 1992 on the conservation of natural habitats and of wild fauna and flora. Council of the European Communities, Official 731 Journal of the European Communities 22.7.92, no L 206/7-50 EC (2013) Green Infrastructure (GI) — Enhancing Europe’s natural capital. European Commission, Brussels EC (2020) EU Biodiversity strategy for 2030—Bringing nature back into our lives. European Commission, Brussels Forrester DI, Bauhus J (2016) A Review of Processes Behind Diversity—Productivity Relationships in Forests. Curr Forestry Rep 2:45–61. https://doi.org/10.1007/s40725-016-0031-2 Gillman LN, Wright SD (2006) The influence of productivity on the species richness of plants: a critical assessment. Ecology 87:1234-1243. https://doi.org/10.1890/0012-9658(2006)87[1234:TIOPOT]2.0.CO;2 GRASS Development Team (2025) Geographic Resources Analysis Support System (GRASS) Software, Version 8.4. Open Source Geospatial Foundation. https://grass.osgeo.org, DOI: 10.5281/zenodo.5176030 Gustafsson L, Andersson J, Jonsson F, Jönsson M, Jonsson M, Nordin U, Strengbom J, Johansson V (2025) Remnant continuity forests are essential for sustaining epiphytic biodiversity in boreal production forest landscapes. J Appl Ecol. https://doi.org/10.1111/1365-2664.70090 Hämäläinen A, Strengbom J, Ranius T (2020) Low-productivity boreal forests have high conservation value for lichens. Journal of Applied Ecology 57:43-54. https://doi.org/10.1111/1365-2664.13509 Hämäläinen A, Runnel K, Ranius T, Strengbom J (2024) Diversity of forest structures important for biodiversity is determined by the combined effects of productivity, stand age, and management. Ambio 53:718–729. https://doi.org/10.1007/s13280-023-01971-9 Hanski I (2011) Habitat Loss, the Dynamics of Biodiversity, and a Perspective on Conservation. Ambio 40:248-255. DOI: 10.1007/s13280-011-0147-3 Joks M, Helm A, Kasari-Toussaint L et al (2023) A simulation model of functional habitat connectivity demonstrates the importance of species establishment in older forests. Ecological Modelling 481:110361. https://doi.org/10.1016/j.ecolmodel.2023.110361 Jonsson BG, Svensson J, Mikusinski G, Manton M, Angelstam P (2019) European Union’s last intact forest landscapes are at a value chain crossroad between multiple use and intensified wood production. Forests 10:564. https://doi.org/10.3390/f10070564 Keskitalo C, Naumov V, Ahlkrona E, Jönsson C, Olsson B (2019) Nationella marktäckedata tilläggsskikt produktivitet. Naturvårdsverket, Stockholm Körner C (2021) Plant ecology at high elevations. In Alpine plant life: functional plant ecology of high mountain ecosystems (pp. 1-22). Cham: Springer International Publishing. https://doi.org/10.1007/978-3-030-59538-8_1 Kyaschenko J, Strengbom J, Felton A, Aakala T, Staland H, Ranius T (2022) Increase in dead wood, large living trees and tree diversity, yet decrease in understory vegetation cover: The effect of three decades of biodiversity-oriented forest policy in Swedish forests. Journal of Environmental Management, 313:114993. https://doi.org/10.1016/j.jenvman.2022.114993 Mikkonen N, Leikola N, Lehtomäki J, Halme P, Moilanen A (2023) National high-resolution conservation prioritisation of boreal forests. Forest Ecology and Management 541:121079. https://doi.org/10.1016/j.foreco.2023.121079 Mikusiński G, Orlikowska EH, Bubnicki JW, Jonsson BG, Svensson J (2021). Strengthening the network of high conservation value forests in boreal landscapes. Frontiers in Ecology and Evolution 8:595730. https://doi.org/10.3389/fevo.2020.595730 Naumburg B (ed.) (2024) Forests and forestry in Sweden. Stockholm, The Royal Swedish Academy of Agriculture and Forestry Nilsson M, Ahlkrona E, Jönsson C, Allard A (2020) Regionala jämförelser mellan Nationella Marktäckedata och fältdata från Riksskogstaxeringen och NILS. SLU, Umeå and Metria, Stockholm Orlikowska EH, Roberge JM, Svensson J, Mikusiński G (2020) Hit or miss? Evaluating Natura 2000 sites for conservation of forest bird habitat in Sweden. Global Ecology and Conservation 22, e00939. https://doi.org/10.1016/j.gecco.2020.e00939 Östlund L, Zackrisson L, Axelsson, AL (1997) The history and transformation of a Scandinavian boreal forest landscape since the 19th century. Can. J. For. Res. 27:1198–1206. https://doi.org/10.1139/x97-070 Potapov P et al (2008) Mapping the World's Intact Forest Landscapes by Remote Sensing. Ecology and Society 13:51. https://www.jstor.org/stable/26267984 Pressey RL, Cabeza M, Watts ME, Cowling RM, Wilson KA (2007) Conservation planning in a changing world. Trends in Ecology & Evolution 22:583–592. https://doi.org/10.1016/j.tree.2007.10.001 Ramberg E, Edman M, Granath G, Sjögren J, Strengbom J (2026) Prescribed burning for boreal forest restoration: Evaluating challenges and conservation outcomes. Ambio 55:608–619. https://doi.org/10.1007/s13280-025-02248-z Sandström J, Edman M, Jonsson BG (2020) Rocky pine forests in the High Coast Region in Sweden: structure, dynamics and history. Nature Conservation 38:101–130. https://doi.org/10.3897/natureconservation.38.34870 SCB (2025) Formellt skyddad skogsmark, frivilliga avsättningar, hänsynsytor samt improduktiv skogsmark 2024. MI 41 Rapport 2025:02 SLU (2025) Forest Statistics 2025. Swedish University of Agricultural Sciences, Umeå Svancara LK, Brannon JR, Scott M, Groves CR, Noss RF, Pressey RL (2005) Policy-driven versus evidence-based conservation: a review of political targets and biological needs. BioScience 55:989-995. https://doi.org/10.1641/0006-3568(2005)055[0989:PVECAR]2.0.CO;2 Swedish EPA (2016) A database of High Conservation Value Forests in Sweden. Available online at https://geodata.naturvardsverket.se/nedladdning/land/skogliga_vardekarnor_2016.zip. Updated with new data from the mountain region 2019-2020 Swedish EPA (2019) National landcover database. https:// www.naturvardsverket.se/verktyg-och-tjanster/kartor-och-karttjanster/ nationella-marktackedata Swedish Forest Agency (2022) Levande skogar - Fördjupad utvärdering 2023. Skogsstyrelsen Rapport 2022/12 Swedish Forest Agency (2025) Skogsvårdslagstiftningen – Gällande regler 1 april 2025. Skogsstyrelsen (in Swedish) Svensson J, Andersson J, Sandström P, Mikusiński G, Jonsson BG (2019) Landscape trajectory of natural boreal forest loss as an impediment to green infrastructure Cons. Biol. 33:152-163. https://doi.org/10.1111/cobi.13148 Svensson J, Bubnicki JW, Jonsson BG, Andersson J, Mikusiński G (2020) Conservation significance of intact forest landscapes in the Scandinavian Mountains Green Belt. Landscape Ecology 35:2113–2131. https://doi.org/10.1007/s10980-020-01088-4 Svensson J, Wallin I, Droste N (2026) Makten över skogen. Rapport 3, Skogsstyrelsen, Uliczka H, Angelstam P (1999) Occurrence of epiphytic macrolichens in relation to tree species and age in managed boreal forest. Ecography, 22: 396-405. https://www.jstor.org/stable/3683231 Villard MA, Haché S (2012) Conifer plantations consistently act as barriers to movement in a deciduous forest songbird: A translocation experiment. Biological Conservation 155:33-37. https://doi.org/10.1016/j.biocon.2012.06.007 Wang X, Svensson J, Jonsson BG, Singh NJ, Bubnicki JW, Lopéz-Peinado A, Angelstam P, Mikusiński G, Ardö J (2025). Where to restore: connectivity forest for spatial prioritization in forest landscape restoration. iScience 28:113263. https://doi.org/10.1016/j.isci.2025.113263 Additional Declarations No competing interests reported. Supplementary Files Appendix1.docx Cite Share Download PDF Status: Under Review Version 1 posted Reviewers agreed at journal 18 May, 2026 Reviewers agreed at journal 18 May, 2026 Reviewers invited by journal 22 Apr, 2026 Editor assigned by journal 13 Apr, 2026 Submission checks completed at journal 13 Apr, 2026 First submitted to journal 10 Apr, 2026 You are reading this latest preprint version Research Square lets you share your work early, gain feedback from the community, and start making changes to your manuscript prior to peer review in a journal. As a division of Research Square Company, we’re committed to making research communication faster, fairer, and more useful. We do this by developing innovative software and high quality services for the global research community. Our growing team is made up of researchers and industry professionals working together to solve the most critical problems facing scientific publishing. Also discoverable on Platform About Our Team In Review Editorial Policies Advisory Board Help Center Resources Author Services Accessibility API Access RSS feed Manage Cookie Preferences © Research Square 2026 | ISSN 2693-5015 (online) Privacy Policy Terms of Service Do Not Sell My Personal Information {"props":{"pageProps":{"initialData":{"identity":"rs-9379486","acceptedTermsAndConditions":true,"allowDirectSubmit":false,"archivedVersions":[],"articleType":"Research Article","associatedPublications":[],"authors":[{"id":628957354,"identity":"08f9c154-7aa8-40d5-9ea1-212eccb10c63","order_by":0,"name":"Bengt Gunnar Jonsson","email":"data:image/png;base64,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","orcid":"","institution":"Mid Sweden University","correspondingAuthor":true,"prefix":"","firstName":"Bengt","middleName":"Gunnar","lastName":"Jonsson","suffix":""},{"id":628957355,"identity":"5213b731-3763-4cfe-b086-9d94b3212361","order_by":1,"name":"Jakub Witold Bubnicki","email":"","orcid":"","institution":"Polish Academy of Sciences","correspondingAuthor":false,"prefix":"","firstName":"Jakub","middleName":"Witold","lastName":"Bubnicki","suffix":""},{"id":628957356,"identity":"3788eaf9-762c-47cc-ba1f-bc282b0ac979","order_by":2,"name":"Grzegorz Mikusiński","email":"","orcid":"","institution":"Swedish University of Agricultural Sciences","correspondingAuthor":false,"prefix":"","firstName":"Grzegorz","middleName":"","lastName":"Mikusiński","suffix":""},{"id":628957358,"identity":"1eac7d8e-b4a8-4c64-9fab-60288aa7ec4c","order_by":3,"name":"Johan Svensson","email":"","orcid":"","institution":"Swedish University of Agricultural Sciences","correspondingAuthor":false,"prefix":"","firstName":"Johan","middleName":"","lastName":"Svensson","suffix":""},{"id":628957359,"identity":"5bed9028-8312-4454-b5dd-66eef3622315","order_by":4,"name":"Michael Manton","email":"","orcid":"","institution":"Vytautas Magnus University","correspondingAuthor":false,"prefix":"","firstName":"Michael","middleName":"","lastName":"Manton","suffix":""},{"id":628957361,"identity":"bd49f8e5-83d8-4e5e-b798-68068fe7cf90","order_by":5,"name":"Taras Yamelynets","email":"","orcid":"","institution":"Ivan Franko National University","correspondingAuthor":false,"prefix":"","firstName":"Taras","middleName":"","lastName":"Yamelynets","suffix":""},{"id":628957364,"identity":"442027af-f0cd-4541-aec8-418106472d52","order_by":6,"name":"Per Angelstam","email":"","orcid":"","institution":"University of Inland Norway","correspondingAuthor":false,"prefix":"","firstName":"Per","middleName":"","lastName":"Angelstam","suffix":""}],"badges":[],"createdAt":"2026-04-10 12:24:48","currentVersionCode":1,"declarations":"","doi":"10.21203/rs.3.rs-9379486/v1","doiUrl":"https://doi.org/10.21203/rs.3.rs-9379486/v1","draftVersion":[],"editorialEvents":[],"editorialNote":"","failedWorkflow":false,"files":[{"id":108235360,"identity":"567b5fde-8f51-4f61-8d49-5395bc62a85a","added_by":"auto","created_at":"2026-04-30 18:38:01","extension":"png","order_by":1,"title":"Figure 1","display":"","copyAsset":false,"role":"figure","size":187888,"visible":true,"origin":"","legend":"\u003cp\u003e\u003cem\u003eSwedish forest ecoregions used on the study\u003c/em\u003e\u003c/p\u003e","description":"","filename":"1.png","url":"https://assets-eu.researchsquare.com/files/rs-9379486/v1/dfae8846268242e3ce5204b7.png"},{"id":108235362,"identity":"9d8df391-09c5-4ce0-9a1d-cb5e8f855714","added_by":"auto","created_at":"2026-04-30 18:38:02","extension":"png","order_by":2,"title":"Figure 2","display":"","copyAsset":false,"role":"figure","size":387221,"visible":true,"origin":"","legend":"\u003cp\u003e\u003cem\u003eMaps showing the proportion of LPF for all forests and for five different forest types (based on dominant species) using a forest filtering algorithm with a moving window of 100 km\u003c/em\u003e\u003csup\u003e\u003cem\u003e2\u003c/em\u003e\u003c/sup\u003e\u003c/p\u003e","description":"","filename":"2.png","url":"https://assets-eu.researchsquare.com/files/rs-9379486/v1/47169f28d53527a957e053cc.png"},{"id":108491760,"identity":"4413d424-d0a8-490a-8a3b-068800b80f89","added_by":"auto","created_at":"2026-05-05 09:55:30","extension":"png","order_by":3,"title":"Figure 3","display":"","copyAsset":false,"role":"figure","size":30928,"visible":true,"origin":"","legend":"\u003cp\u003e\u003cem\u003eSize distribution of low productive forests for connected hectare pixels with more than 50% LPF\u003c/em\u003e\u003c/p\u003e","description":"","filename":"3.png","url":"https://assets-eu.researchsquare.com/files/rs-9379486/v1/776cf1dbd1e331d87995ce0f.png"},{"id":108235361,"identity":"4feb8713-9fc5-4792-ade5-6e9daea047c5","added_by":"auto","created_at":"2026-04-30 18:38:01","extension":"jpeg","order_by":4,"title":"Figure 4","display":"","copyAsset":false,"role":"figure","size":1594586,"visible":true,"origin":"","legend":"\u003cp\u003e\u003cem\u003eHotspot – coldspot maps showing the distance to nearest functional patch and the improved connectivity for all forests (upper row) taken together and for pine forests (lower row) by adding LPFs for generalist and specialised species. Percentages in the inserted tables show the fraction of the region that represents functional networks from the mountain region (R1) to the nemoral region (R5)\u003c/em\u003e\u003c/p\u003e","description":"","filename":"4.jpeg","url":"https://assets-eu.researchsquare.com/files/rs-9379486/v1/2b3a112cb040de2f21f8679d.jpeg"},{"id":108804093,"identity":"5def920b-e01d-49fb-97c0-0ae15a70ffa2","added_by":"auto","created_at":"2026-05-08 15:15:40","extension":"pdf","order_by":0,"title":"","display":"","copyAsset":false,"role":"manuscript-pdf","size":2626331,"visible":true,"origin":"","legend":"","description":"","filename":"manuscript.pdf","url":"https://assets-eu.researchsquare.com/files/rs-9379486/v1/744b87ff-f616-43ff-b03a-9d466464cf84.pdf"},{"id":108235359,"identity":"47fb05af-2bb5-4305-8553-de1e1fea8ddc","added_by":"auto","created_at":"2026-04-30 18:38:01","extension":"docx","order_by":0,"title":"","display":"","copyAsset":false,"role":"supplement","size":939953,"visible":true,"origin":"","legend":"","description":"","filename":"Appendix1.docx","url":"https://assets-eu.researchsquare.com/files/rs-9379486/v1/cc63a6c1bebd8e0162215d55.docx"}],"financialInterests":"No competing interests reported.","formattedTitle":"Limited contribution of low-productive forests to green infrastructure along a temperate to boreal biogeographic gradient","fulltext":[{"header":"Introduction","content":"\u003cp\u003eGreen infrastructure (GI) is a European Union policy concept emphasizing the necessity of maintaining landscape-scale functional connectivity of representative habitat types to support biodiversity conservation and ecosystem services provisioning (EC 2013). In regions where natural forest habitats have been lost and fragmented due to historical and ongoing land use practices, establishing a functional GI presents a substantial challenge. Here, recent research (e.g., Orlikowska et al. 2020; Angelstam et al. 2020, 2026; Joks et al. 2023) indicates that areas designated for biodiversity conservation are mostly inadequate in quality, size and too fragmented to be ecologically functional. The prevailing commercially managed forest matrix often acts as resistance or barrier to functional connectivity (Villard and Hach\u0026eacute; 2012, Svensson et al. 2019, Angelstam et al. 2020, 2026, Mikkonen et al. 2023).\u003c/p\u003e\n\u003cp\u003eHowever, within commercially managed forest landscapes, low-productive forest (LPF) stands excluded from management due to economic and regulatory limitations, may sustain structural and functional attributes typical for natural forests, such as old and dead trees, multilayered canopy and open and sun-exposed conditions (Sandstr\u0026ouml;m et al. 2020). From a GI perspective, LPFs may contribute naturalness attributes such as 1) characteristics of primary and undisturbed forests, 2) increasing heterogeneity of largely homogenous landscapes, 3) temporal continuity and 4) spatial connectivity among fragmented forest patches of significant conservation importance.\u003c/p\u003e\n\u003cp\u003eWhile site productivity and species richness are commonly positively correlated as indicators of biodiversity value (Gillman and Wright 2006; H\u0026auml;m\u0026auml;l\u0026auml;inen et al. 2024), LPFs provide valuable habitats for biodiversity and refugia for some forest species that are rare in managed forests (Uliczka and Angelstam 1999, H\u0026auml;m\u0026auml;l\u0026auml;inen et al. 2020). This characteristic of LPFs is partially explained by longer temporal forest- and tree-continuity (e.g. Gustafsson et al. 2025). Forest productivity is primarily governed by water availability, soil properties, and climate. Interacting disturbance regimes further shape tree species composition, canopy structure, and spatial heterogeneity (e.g., Forrester et al. 2016). Productivity generally declines with increasing latitude and altitude, for example in tree-line transitions to alpine and tundra systems (K\u0026ouml;rner 2021). Although productive stands dominate temperate and boreal forest regions, local edaphic and topographic constraints, such as peat substrates, drought-prone soils, steep slopes, and shallow rocky soil, can limit growth, producing LPFs as discrete patches within otherwise productive forests.\u0026nbsp;\u003c/p\u003e\n\u003cp\u003eSweden provides a compelling case study of challenges to conserve, manage and restore functional habitat networks as GI (Angelstam et al. 2020, 2026). With a predominant portion of the forests not providing favourable habitat conditions (Angelstam et al. 2020), securing the last remnants of natural habitats for biodiversity conservation remain a core challenge (Jonsson et al. 2019, Angelstam et al. 2026). However, in spite of current re-iterated policy targets, such as the Convention on Biological Diversity (CBD), the Aichi targets (CBD 2010) and the CBD Kunming-Montreal framework\u0026apos;s third target on protected areas (CBD 2022), the functionality of GI is deemed inadequate in Sweden due to limited area protection and insufficient conservation management (Angelstam et al. 2020, Swedish Forest Agency 2022). To establish sufficient amounts of representative habitats with functional connectivity at a landscape and regional scale remains a major political and economic challenge (Angelstam et al. 2023, Wang et al. 2025, Svensson et al. 2026). Therefore, evaluating the potential of LPFs to support and strengthen habitat networks for various forest types is crucial.\u003c/p\u003e\n\u003cp\u003eOut of Sweden\u0026rsquo;s total forest cover of 28 million hectares, approximately 23.5 million hectares are classified as productive, capable of supporting commercial forestry due to a volume growth potential at an annual rate of \u0026gt;1m\u003csup\u003e3\u003c/sup\u003e ha\u003csup\u003e-1\u003c/sup\u003e over a 100-year rotation period. Conversely, the remaining 4.4 million hectares are categorized as low-productive forests (SLU 2025). Legislation currently restricts commercial forestry in LPFs (Swedish Forest Agency 2025). Hence, since the establishment of the present commercial rotation forestry system in the mid-20\u003csup\u003eth\u003c/sup\u003e century, Swedish LPFs have essentially been set aside. Certain LPFs are identified as key Natura 2000 habitat types (CEC 1992) for conservation, including bog woodlands (91D0) and Fennoscandian deciduous swamp woods (9080). However, in the case of the predominant Natura 2000 habitat type, Western taiga (9010), the significance of LPFs lies in their potential capacity to strengthen the overall ecological functionality of habitat networks in managed forest landscapes.\u003c/p\u003e\n\u003cp\u003eThe primary objective of national environmental goals, EU directives, and CBD agreements is to halt biodiversity loss. This implies that conservation efforts must transcend mere quantitative measures, by also considering how functional GI can be established to ensure viable populations of species demanding higher levels of naturalness in the long term (Pressey et al. 2007). It is within this context that the role of LPFs should be evaluated, and especially their potential contribution to biodiversity conservation as elements of functional GI in different forest types and biogeographic regions. Therefore, to preserve functional GI through conservation, management, and restoration, the role of LPFs must be carefully examined.\u003c/p\u003e\n\u003cp\u003eThe objective of this study is to evaluate the potential role of LPFs in the network of both protected and non-protected High Conservation Value Forests (HCVF) as a contribution to GI functionality in Sweden. Our investigation is based on comprehensive national databases created using remote sensing and field surveys. Consequently, our research contributes to evaluating the on-the-ground effects of conservation policies concerning the role of LPFs in supporting functional \u0026nbsp;green infrastructure across different forest types in different forest ecoregions.\u003c/p\u003e\n\u003ch2 id=\"_Toc189659340\"\u003eStudy area\u003c/h2\u003e\n\u003cp\u003eThe study area covers the entire land area of Sweden, which spans a north-south gradient of 1600 km from the temperate lowland ecoregion (nemoral) in the south to boreal and mountain ecoregions in the north (Ahti et al. 1968). Forests is the dominating (69%) land cover, with significant regional variations in main forest types (SLU 2025). For regional analysis, we have applied the five biogeographical ecoregions commonly used for official forest statistics in Sweden (SCB 2025, Figure 1), based on biogeographic conditions, main landscape characteristics.\u003c/p\u003e\n\u003cp\u003eForestry practices focused on wood production have a long history in Sweden. While substantial forest areas were converted to agricultural land during Millenia in southern Sweden (Berglund 1992), since the 13th century, the mining industry has impacted forest structure in south-central ecoregions. Large-scale industrial forestry, however, began in the early 1800s in southwestern Sweden (Bladh 1995; Angelstam et al. 2013), followed by a \u0026quot;timber frontier\u0026quot; expanding into northern Sweden from the late 1800s (\u0026Ouml;stlund et al. 1997). Concerns regarding sustained yield forestry in the early 1900s led to a management system involving rotation forestry based on clear-cut harvesting, soil scarification, artificial planting, pre-commercial cleaning and commercial thinning (Angelstam et al. 2022), which has been fully implemented since the mid-1900s (Naumborg 2024). Concurrently, significant areas of bogs (approximately 1.5 million ha) were drained to encourage wood production. At present, around 200,000 ha (1%) of Swedish forests undergo clear-cut annually, with an additional 200,000 ha being subject to commercial and pre-commercial thinning (SLU 2025).\u003c/p\u003e\n\u003cp\u003eCurrently, formally protected areas on productive land contribute most in terms of habitat quality (Kyaschenko et al. 2022), and these cover 6.3 % of Sweden\u0026rsquo;s productive forest land, and 9.4% of all forest land (Statistics Sweden 2025). However, there is a stark contrast in the proportion of protected areas between the remote mountain forests in northwestern Sweden (59.1%, all forest land), and the rest of the country (4.2%; ibid.) where wood production dominates. In addition, 5.9% is voluntarily set aside on productive forest land largely due to the widespread adoption of forest certification schemes such as FSC and PEFC (ibid.). While LPFs cover 16% of Sweden\u0026rsquo;s total forest area, the majority (11%) are located outside formally protected areas (SCB 2025).\u003c/p\u003e"},{"header":"Methods","content":"\u003cp\u003eTo evaluate the significance of LPFs, three characteristics were assessed for each of the five ecoregions: area extent of different forest types, distribution of patch size, and role in functional landscape connectivity.\u003c/p\u003e\n\u003ch3 id=\"_Toc189659342\"\u003eArea of different forest types\u003c/h3\u003e\n\u003cp\u003eThe primary source utilized for delineating various forest types was the national land cover data of Sweden (NMD; Swedish EPA 2019) generated during the period 2017-2019. This comprehensive, raster based, land cover map covers the entire country, mapping various forest types with a minimum mapping unit of 0.01 hectares. To detect LPFs, supplementary land cover data on productivity (Keskitalo et al. 2019) was employed. This LPF dataset, a primary source of data for our analysis, integrates satellite data (Sentinel-2), laser scanning, soil characteristics, moisture content, and climate. Validation assessment (at a 20-meter radius scale) was conducted using data from the National Forest Inventory (40,100 plots) and the National Inventory of the Landscape in Sweden (2,200 plots; Keskitalo et al. 2019), revealing high precision in identifying productive forests (99%) and somewhat lower accuracy for LPFs (75%) when considering the predominant land cover class within the 20-meter plot (Nilsson et al. 2020). It is important to highlight that certain large forest owners define LPFs from a management perspective and incorporate technical management constraints such as steep terrain, based on their internal stand level inventories. Consequently, the actual areal extent of LPFs excluded from logging activities may differ from the estimates obtained from the NMD.\u003c/p\u003e\n\u003cp\u003eThe initial NMD land cover classes encompasses seven forest types occurring on mineral and wet soils (totalling 14 combinations). To simplify while retaining biological significance, the two soil types were combined across all forest types. Norway spruce (\u003cem\u003ePicea abies\u003c/em\u003e) forest and mixed conifer forests were grouped into one category, as were hardwood and mixed hardwood-deciduous forests (only present in the southern ecoregions), thus generating five distinct forest types; 1) deciduous (mainly birch dominated), 2) hardwood (dominated by broadleaf tree species with a southern distribution), 3) mixed deciduous/coniferous, 4) Scots pine (\u003cem\u003ePinus sylvestris\u003c/em\u003e), 5) spruce and mixed coniferous.\u003c/p\u003e\n\u003cp\u003eStatistical analyses and data visualizations were performed using GRASS GIS 8.4 (GRASS Development Team, 2025) and custom Python 3.10 scripts. Two publicly available national spatial datasets served as input: 1) a compilation of High Conservation Value Forests (HCV forests) in Sweden, produced by the Swedish Environmental Protection Agency (Swedish EPA 2016 and subsequent updates); and 2) the National Land Cover Data (NMD), also produced by the Swedish Environmental Protection Agency (Swedish EPA 2019). The NMD dataset includes the LPF layer as described above. The density of LPF was visualized by implementing a 100 km\u003csup\u003e2\u003c/sup\u003e moving window across Sweden.\u003c/p\u003e\n\u003ch3 id=\"_Toc189659344\"\u003ePatch size\u003c/h3\u003e\n\u003cp\u003eIn terms of patch size, the distribution of LPFs was evaluated based on four size-categories: \u0026lt;5 ha, 5-50 ha, \u0026gt;50-500 ha, \u0026gt;500 ha. A threshold was employed to delineate LPFs at a 1 ha pixel scale, selecting pixels with total LPF cover above 50%, i.e., hectare pixels dominated by LPF. Patches were defined as connected LPF pixels (8-neighbours connectivity), with patch size computed for each patch.\u003c/p\u003e\n\u003ch3 id=\"_Toc189659345\"\u003eFunctional connectivity\u003c/h3\u003e\n\u003cp\u003eTo assess the potential impact of low-productive forests on functional connectivity, we calculated how their addition to existing HCVF enhances habitat network functionality. The functional connectivity of HCVF, both with and without the addition LPF, was assessed using the algorithm outlined in Angelstam et al. (2020). Spatial modelling parameters were set to represent two virtual species with differing habitat requirements i.e. with higher and lower demands in respect to HCVF availability (Table 1). Forest patches at 1 ha scale were delineated using the same rules as for the patch size analysis and further defined by buffering HCVF- pixels by 100 m (200 m distance between patches) for specialist species and by 200 m (400 m distance between patches) for generalist species. Patches meeting minimum size criteria (50 and 5 ha respectively for specialist and generalist species) were selected, using moving window sizes of 2x2 km (specialist species) and 5x5 km (generalist species) to represent the landscape scale. A minimum habitat availability threshold of 20% was applied to identify functional HCVF networks at the landscape scale (cf. Hanski 2011; Svancara et al. 2005).\u003c/p\u003e\n\u003cp\u003eThe outcome was a mask of interconnected functional areas derived through the aforementioned steps. The actual area was calculated as the sum of all pixels from the original layer intersecting with this mask. This process was repeated for all forest types and ecoregions under two scenarios: 1) HCVF only and 2) combined HCVF and LPF. The results are visualized as the distribution of hotspots and coldspots based on the distance to the nearest functional patch, and hence show habitat network functionality across Sweden.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003e\u003cem\u003eTable 1\u003c/em\u003e\u003c/strong\u003e\u003cem\u003e\u0026nbsp;Characteristics of the two virtual species used in the analysis of functional connectivity. The threshold for habitat availability at landscape level is set to 20% for both generalist and specialist species\u003c/em\u003e\u003c/p\u003e\n\u003ctable border=\"1\" cellspacing=\"0\" cellpadding=\"0\" align=\"left\" width=\"100%\"\u003e\n \u003ctbody\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 27px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 21px;\"\u003e\n \u003cp\u003e\u003cstrong\u003e\u003cem\u003eBuffer size\u003c/em\u003e\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 23px;\"\u003e\n \u003cp\u003e\u003cstrong\u003e\u003cem\u003ePatch size limit\u003c/em\u003e\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 27px;\"\u003e\n \u003cp\u003e\u003cstrong\u003e\u003cem\u003eLandscape size\u003c/em\u003e\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 27px;\"\u003e\n \u003cp\u003eGeneralist species\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 21px;\"\u003e\n \u003cp\u003e100 m\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 23px;\"\u003e\n \u003cp\u003e5 ha\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 27px;\"\u003e\n \u003cp\u003e5x5 km\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 27px;\"\u003e\n \u003cp\u003eSpecialist species\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 21px;\"\u003e\n \u003cp\u003e200 m\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 23px;\"\u003e\n \u003cp\u003e50 ha\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 27px;\"\u003e\n \u003cp\u003e2x2 km\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003c/tbody\u003e\n\u003c/table\u003e"},{"header":"Results","content":"\u003ch3\u003eArea of different forest types\u003c/h3\u003e\n\u003cp\u003eLPFs in Sweden are primarily concentrated to the mountain region with 58% of all forests and 77% of HCVF classified as LPF (Table 2). For the other ecoregions, the fraction of LPFs is substantially lower, varying from 19% and 11% in the north boreal, 12% and 10% in south boreal SE, 8% and 9% in hemiboreal, and 6% and 7% in the nemoral ecoregions. Pine is the dominant forest type among LPFs in all forest ecoregions below the mountain region, while deciduous forest dominates in the mountain region. A series of maps in Figure 2 illustrate the density of different LPF types.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003e\u003cem\u003eTable 2\u003c/em\u003e\u003c/strong\u003e\u003cem\u003e\u0026nbsp;Total area of LPF and its fraction (%) of the individual forest types and out of delineated HCV forests in the five Swedish forest ecoregions. NA denotes a forest type missing in the region\u003c/em\u003e\u003c/p\u003e\n\u003ctable border=\"1\" cellspacing=\"0\" cellpadding=\"0\" width=\"548\"\u003e\n \u003ctbody\u003e\n \u003ctr\u003e\n \u003ctd nowrap=\"\" valign=\"top\" style=\"width: 106px;\"\u003e\n \u003cp\u003e\u003cstrong\u003e\u003cem\u003eRegion\u003c/em\u003e\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"top\" style=\"width: 106px;\"\u003e\n \u003cp\u003e\u003cstrong\u003e\u003cem\u003eForest type\u003c/em\u003e\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"top\" style=\"width: 106px;\"\u003e\n \u003cp\u003e\u003cstrong\u003e\u003cem\u003eLPF area (kha)\u003c/em\u003e\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"top\" style=\"width: 117px;\"\u003e\n \u003cp\u003e\u003cstrong\u003e\u003cem\u003e% of forest type\u003c/em\u003e\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"top\" style=\"width: 113px;\"\u003e\n \u003cp\u003e\u003cstrong\u003e\u003cem\u003e% of HCVF\u003c/em\u003e\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd nowrap=\"\" valign=\"top\" style=\"width: 106px;\"\u003e\n \u003cp\u003eMountain\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"top\" style=\"width: 106px;\"\u003e\n \u003cp\u003eDeciduous\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"top\" style=\"width: 106px;\"\u003e\n \u003cp\u003e837.8\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"top\" style=\"width: 117px;\"\u003e\n \u003cp\u003e94\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"top\" style=\"width: 113px;\"\u003e\n \u003cp\u003e69\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd nowrap=\"\" valign=\"top\" style=\"width: 106px;\"\u003e\n \u003cp\u003eMountain\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"top\" style=\"width: 106px;\"\u003e\n \u003cp\u003eHardwood\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"top\" style=\"width: 106px;\"\u003e\n \u003cp\u003eNA\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"top\" style=\"width: 117px;\"\u003e\n \u003cp\u003eNA\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"top\" style=\"width: 113px;\"\u003e\n \u003cp\u003eNA\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd nowrap=\"\" valign=\"top\" style=\"width: 106px;\"\u003e\n \u003cp\u003eMountain\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"top\" style=\"width: 106px;\"\u003e\n \u003cp\u003eMixed\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"top\" style=\"width: 106px;\"\u003e\n \u003cp\u003e183.0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"top\" style=\"width: 117px;\"\u003e\n \u003cp\u003e48\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"top\" style=\"width: 113px;\"\u003e\n \u003cp\u003e87\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd nowrap=\"\" valign=\"top\" style=\"width: 106px;\"\u003e\n \u003cp\u003eMountain\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"top\" style=\"width: 106px;\"\u003e\n \u003cp\u003ePine\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"top\" style=\"width: 106px;\"\u003e\n \u003cp\u003e185.8\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"top\" style=\"width: 117px;\"\u003e\n \u003cp\u003e49\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"top\" style=\"width: 113px;\"\u003e\n \u003cp\u003e87\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd nowrap=\"\" valign=\"top\" style=\"width: 106px;\"\u003e\n \u003cp\u003eMountain\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"top\" style=\"width: 106px;\"\u003e\n \u003cp\u003eSpruce \u0026amp; mixed\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"top\" style=\"width: 106px;\"\u003e\n \u003cp\u003e220.0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"top\" style=\"width: 117px;\"\u003e\n \u003cp\u003e27\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"top\" style=\"width: 113px;\"\u003e\n \u003cp\u003e92\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd nowrap=\"\" valign=\"top\" style=\"width: 106px;\"\u003e\n \u003cp\u003e\u003cstrong\u003eMountain\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"top\" style=\"width: 106px;\"\u003e\n \u003cp\u003e\u003cstrong\u003eAll\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"top\" style=\"width: 106px;\"\u003e\n \u003cp\u003e\u003cstrong\u003e1426.6\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"top\" style=\"width: 117px;\"\u003e\n \u003cp\u003e\u003cstrong\u003e58\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"top\" style=\"width: 113px;\"\u003e\n \u003cp\u003e\u003cstrong\u003e77\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd nowrap=\"\" valign=\"top\" style=\"width: 106px;\"\u003e\n \u003cp\u003eNorth boreal\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"top\" style=\"width: 106px;\"\u003e\n \u003cp\u003eDeciduous\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"top\" style=\"width: 106px;\"\u003e\n \u003cp\u003e140.6\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"top\" style=\"width: 117px;\"\u003e\n \u003cp\u003e25\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"top\" style=\"width: 113px;\"\u003e\n \u003cp\u003e9\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd nowrap=\"\" valign=\"top\" style=\"width: 106px;\"\u003e\n \u003cp\u003eNorth boreal\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"top\" style=\"width: 106px;\"\u003e\n \u003cp\u003eHardwood\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"top\" style=\"width: 106px;\"\u003e\n \u003cp\u003eNA\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"top\" style=\"width: 117px;\"\u003e\n \u003cp\u003eNA\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"top\" style=\"width: 113px;\"\u003e\n \u003cp\u003eNA\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd nowrap=\"\" valign=\"top\" style=\"width: 106px;\"\u003e\n \u003cp\u003eNorth boreal\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"top\" style=\"width: 106px;\"\u003e\n \u003cp\u003eMixed\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"top\" style=\"width: 106px;\"\u003e\n \u003cp\u003e200.8\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"top\" style=\"width: 117px;\"\u003e\n \u003cp\u003e19\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"top\" style=\"width: 113px;\"\u003e\n \u003cp\u003e10\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd nowrap=\"\" valign=\"top\" style=\"width: 106px;\"\u003e\n \u003cp\u003eNorth boreal\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"top\" style=\"width: 106px;\"\u003e\n \u003cp\u003ePine\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"top\" style=\"width: 106px;\"\u003e\n \u003cp\u003e513.3\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"top\" style=\"width: 117px;\"\u003e\n \u003cp\u003e21\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"top\" style=\"width: 113px;\"\u003e\n \u003cp\u003e10\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd nowrap=\"\" valign=\"top\" style=\"width: 106px;\"\u003e\n \u003cp\u003eNorth boreal\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"top\" style=\"width: 106px;\"\u003e\n \u003cp\u003eSpruce \u0026amp; mixed\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"top\" style=\"width: 106px;\"\u003e\n \u003cp\u003e152.1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"top\" style=\"width: 117px;\"\u003e\n \u003cp\u003e12\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"top\" style=\"width: 113px;\"\u003e\n \u003cp\u003e17\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd nowrap=\"\" valign=\"top\" style=\"width: 106px;\"\u003e\n \u003cp\u003e\u003cstrong\u003eNorth boreal\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"top\" style=\"width: 106px;\"\u003e\n \u003cp\u003e\u003cstrong\u003eAll\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"top\" style=\"width: 106px;\"\u003e\n \u003cp\u003e\u003cstrong\u003e1006.9\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"top\" style=\"width: 117px;\"\u003e\n \u003cp\u003e\u003cstrong\u003e19\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"top\" style=\"width: 113px;\"\u003e\n \u003cp\u003e\u003cstrong\u003e11\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd nowrap=\"\" valign=\"top\" style=\"width: 106px;\"\u003e\n \u003cp\u003eSouth boreal\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"top\" style=\"width: 106px;\"\u003e\n \u003cp\u003eDeciduous\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"top\" style=\"width: 106px;\"\u003e\n \u003cp\u003e82.3\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"top\" style=\"width: 117px;\"\u003e\n \u003cp\u003e15\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"top\" style=\"width: 113px;\"\u003e\n \u003cp\u003e10\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd nowrap=\"\" valign=\"top\" style=\"width: 106px;\"\u003e\n \u003cp\u003eSouth boreal\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"top\" style=\"width: 106px;\"\u003e\n \u003cp\u003eHardwood\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"top\" style=\"width: 106px;\"\u003e\n \u003cp\u003e0.4\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"top\" style=\"width: 117px;\"\u003e\n \u003cp\u003e2\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"top\" style=\"width: 113px;\"\u003e\n \u003cp\u003e12\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd nowrap=\"\" valign=\"top\" style=\"width: 106px;\"\u003e\n \u003cp\u003eSouth boreal\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"top\" style=\"width: 106px;\"\u003e\n \u003cp\u003eMixed\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"top\" style=\"width: 106px;\"\u003e\n \u003cp\u003e147.1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"top\" style=\"width: 117px;\"\u003e\n \u003cp\u003e13\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"top\" style=\"width: 113px;\"\u003e\n \u003cp\u003e10\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd nowrap=\"\" valign=\"top\" style=\"width: 106px;\"\u003e\n \u003cp\u003eSouth boreal\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"top\" style=\"width: 106px;\"\u003e\n \u003cp\u003ePine\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"top\" style=\"width: 106px;\"\u003e\n \u003cp\u003e549.8\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"top\" style=\"width: 117px;\"\u003e\n \u003cp\u003e17\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"top\" style=\"width: 113px;\"\u003e\n \u003cp\u003e9\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd nowrap=\"\" valign=\"top\" style=\"width: 106px;\"\u003e\n \u003cp\u003eSouth boreal\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"top\" style=\"width: 106px;\"\u003e\n \u003cp\u003eSpruce \u0026amp; mixed\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"top\" style=\"width: 106px;\"\u003e\n \u003cp\u003e112.7\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"top\" style=\"width: 117px;\"\u003e\n \u003cp\u003e4\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"top\" style=\"width: 113px;\"\u003e\n \u003cp\u003e14\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd nowrap=\"\" valign=\"top\" style=\"width: 106px;\"\u003e\n \u003cp\u003e\u003cstrong\u003eSouth boreal\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"top\" style=\"width: 106px;\"\u003e\n \u003cp\u003e\u003cstrong\u003eAll\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"top\" style=\"width: 106px;\"\u003e\n \u003cp\u003e\u003cstrong\u003e892.2\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"top\" style=\"width: 117px;\"\u003e\n \u003cp\u003e\u003cstrong\u003e12\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"top\" style=\"width: 113px;\"\u003e\n \u003cp\u003e\u003cstrong\u003e10\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd nowrap=\"\" valign=\"top\" style=\"width: 106px;\"\u003e\n \u003cp\u003eHemiboreal\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"top\" style=\"width: 106px;\"\u003e\n \u003cp\u003eDeciduous\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"top\" style=\"width: 106px;\"\u003e\n \u003cp\u003e72.6\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"top\" style=\"width: 117px;\"\u003e\n \u003cp\u003e11\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"top\" style=\"width: 113px;\"\u003e\n \u003cp\u003e8\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd nowrap=\"\" valign=\"top\" style=\"width: 106px;\"\u003e\n \u003cp\u003eHemiboreal\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"top\" style=\"width: 106px;\"\u003e\n \u003cp\u003eHardwood\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"top\" style=\"width: 106px;\"\u003e\n \u003cp\u003e16.5\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"top\" style=\"width: 117px;\"\u003e\n \u003cp\u003e4\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"top\" style=\"width: 113px;\"\u003e\n \u003cp\u003e4\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd nowrap=\"\" valign=\"top\" style=\"width: 106px;\"\u003e\n \u003cp\u003eHemiboreal\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"top\" style=\"width: 106px;\"\u003e\n \u003cp\u003eMixed\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"top\" style=\"width: 106px;\"\u003e\n \u003cp\u003e41.5\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"top\" style=\"width: 117px;\"\u003e\n \u003cp\u003e6\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"top\" style=\"width: 113px;\"\u003e\n \u003cp\u003e6\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd nowrap=\"\" valign=\"top\" style=\"width: 106px;\"\u003e\n \u003cp\u003eHemiboreal\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"top\" style=\"width: 106px;\"\u003e\n \u003cp\u003ePine\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"top\" style=\"width: 106px;\"\u003e\n \u003cp\u003e309.4\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"top\" style=\"width: 117px;\"\u003e\n \u003cp\u003e15\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"top\" style=\"width: 113px;\"\u003e\n \u003cp\u003e10\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd nowrap=\"\" valign=\"top\" style=\"width: 106px;\"\u003e\n \u003cp\u003eHemiboreal\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"top\" style=\"width: 106px;\"\u003e\n \u003cp\u003eSpruce \u0026amp; mixed\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"top\" style=\"width: 106px;\"\u003e\n \u003cp\u003e33.1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"top\" style=\"width: 117px;\"\u003e\n \u003cp\u003e2\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"top\" style=\"width: 113px;\"\u003e\n \u003cp\u003e7\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd nowrap=\"\" valign=\"top\" style=\"width: 106px;\"\u003e\n \u003cp\u003e\u003cstrong\u003eHemiboreal\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"top\" style=\"width: 106px;\"\u003e\n \u003cp\u003e\u003cstrong\u003eAll\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"top\" style=\"width: 106px;\"\u003e\n \u003cp\u003e\u003cstrong\u003e473.1\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"top\" style=\"width: 117px;\"\u003e\n \u003cp\u003e\u003cstrong\u003e8\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"top\" style=\"width: 113px;\"\u003e\n \u003cp\u003e\u003cstrong\u003e9\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd nowrap=\"\" valign=\"top\" style=\"width: 106px;\"\u003e\n \u003cp\u003eNemoral\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"top\" style=\"width: 106px;\"\u003e\n \u003cp\u003eDeciduous\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"top\" style=\"width: 106px;\"\u003e\n \u003cp\u003e10.4\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"top\" style=\"width: 117px;\"\u003e\n \u003cp\u003e9\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"top\" style=\"width: 113px;\"\u003e\n \u003cp\u003e10\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd nowrap=\"\" valign=\"top\" style=\"width: 106px;\"\u003e\n \u003cp\u003eNemoral\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"top\" style=\"width: 106px;\"\u003e\n \u003cp\u003eHardwood\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"top\" style=\"width: 106px;\"\u003e\n \u003cp\u003e4.4\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"top\" style=\"width: 117px;\"\u003e\n \u003cp\u003e2\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"top\" style=\"width: 113px;\"\u003e\n \u003cp\u003e14\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd nowrap=\"\" valign=\"top\" style=\"width: 106px;\"\u003e\n \u003cp\u003eNemoral\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"top\" style=\"width: 106px;\"\u003e\n \u003cp\u003eMixed\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"top\" style=\"width: 106px;\"\u003e\n \u003cp\u003e6.7\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"top\" style=\"width: 117px;\"\u003e\n \u003cp\u003e6\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"top\" style=\"width: 113px;\"\u003e\n \u003cp\u003e6\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd nowrap=\"\" valign=\"top\" style=\"width: 106px;\"\u003e\n \u003cp\u003eNemoral\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"top\" style=\"width: 106px;\"\u003e\n \u003cp\u003ePine\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"top\" style=\"width: 106px;\"\u003e\n \u003cp\u003e20.5\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"top\" style=\"width: 117px;\"\u003e\n \u003cp\u003e15\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"top\" style=\"width: 113px;\"\u003e\n \u003cp\u003e5\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd nowrap=\"\" valign=\"top\" style=\"width: 106px;\"\u003e\n \u003cp\u003eNemoral\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"top\" style=\"width: 106px;\"\u003e\n \u003cp\u003eSpruce \u0026amp; mixed\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"top\" style=\"width: 106px;\"\u003e\n \u003cp\u003e3.1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"top\" style=\"width: 117px;\"\u003e\n \u003cp\u003e1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"top\" style=\"width: 113px;\"\u003e\n \u003cp\u003e4\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd nowrap=\"\" valign=\"top\" style=\"width: 106px;\"\u003e\n \u003cp\u003eNemoral\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"top\" style=\"width: 106px;\"\u003e\n \u003cp\u003e\u003cstrong\u003eAll\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"top\" style=\"width: 106px;\"\u003e\n \u003cp\u003e\u003cstrong\u003e45.1\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"top\" style=\"width: 117px;\"\u003e\n \u003cp\u003e\u003cstrong\u003e6\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"top\" style=\"width: 113px;\"\u003e\n \u003cp\u003e\u003cstrong\u003e7\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003c/tbody\u003e\n\u003c/table\u003e\n\u003cp\u003e\u003cstrong\u003ePatch size\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eHabitat patch size plays a crucial role in assessing habitat network functionality. Small to intermediate sized patches dominate LPFs across all forest ecoregions, except for the mountain region where almost 59% of all LPFs are located in patches exceeding 500 hectares. For the other forest ecoregions, the two smallest patch size intervals strongly dominate (Figure 3). For the individual forest types, pine forest and deciduous forest are relatively well represented in the \u0026gt;5-50 ha class while other forest types are mainly restricted to patches less than 5 ha (Table 3).\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003e\u003cem\u003eTable 3\u003c/em\u003e\u003c/strong\u003e\u003cem\u003e\u0026nbsp;Patch size distribution of LPF per region, forest type and patch size class for hectare pixels with more than 50% LPF\u003c/em\u003e\u003c/p\u003e\n\u003ctable border=\"0\" cellspacing=\"0\" cellpadding=\"0\" width=\"551\"\u003e\n \u003ctbody\u003e\n \u003ctr\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 92px;\"\u003e\n \u003cp\u003e\u003cstrong\u003e\u003cem\u003eRegion\u003c/em\u003e\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 116px;\"\u003e\n \u003cp\u003e\u003cstrong\u003e\u003cem\u003eForest type\u003c/em\u003e\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 67px;\"\u003e\n \u003cp\u003e\u003cstrong\u003e\u003cem\u003e% \u0026lt;5 ha\u003c/em\u003e\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 92px;\"\u003e\n \u003cp\u003e\u003cstrong\u003e\u003cem\u003e% 5-50 ha\u003c/em\u003e\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 92px;\"\u003e\n \u003cp\u003e\u003cstrong\u003e\u003cem\u003e% 50-500 ha\u003c/em\u003e\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 92px;\"\u003e\n \u003cp\u003e\u003cstrong\u003e\u003cem\u003e% \u0026gt;500 ha\u003c/em\u003e\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 92px;\"\u003e\n \u003cp\u003eMountain\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 116px;\"\u003e\n \u003cp\u003eDeciduous\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 67px;\"\u003e\n \u003cp\u003e8.6\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 92px;\"\u003e\n \u003cp\u003e15.4\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 92px;\"\u003e\n \u003cp\u003e19.7\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 92px;\"\u003e\n \u003cp\u003e56.4\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 92px;\"\u003e\n \u003cp\u003eMountain\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 116px;\"\u003e\n \u003cp\u003eHardwood\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 67px;\"\u003e\n \u003cp\u003e0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 92px;\"\u003e\n \u003cp\u003e0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 92px;\"\u003e\n \u003cp\u003e0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 92px;\"\u003e\n \u003cp\u003e0\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 92px;\"\u003e\n \u003cp\u003eMountain\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 116px;\"\u003e\n \u003cp\u003eMixed\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 67px;\"\u003e\n \u003cp\u003e47.6\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 92px;\"\u003e\n \u003cp\u003e40.7\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 92px;\"\u003e\n \u003cp\u003e10.6\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 92px;\"\u003e\n \u003cp\u003e1\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 92px;\"\u003e\n \u003cp\u003eMountain\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 116px;\"\u003e\n \u003cp\u003ePine\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 67px;\"\u003e\n \u003cp\u003e33.3\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 92px;\"\u003e\n \u003cp\u003e41\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 92px;\"\u003e\n \u003cp\u003e19.8\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 92px;\"\u003e\n \u003cp\u003e5.8\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 92px;\"\u003e\n \u003cp\u003eMountain\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 116px;\"\u003e\n \u003cp\u003eSpruce \u0026amp; mixed\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 67px;\"\u003e\n \u003cp\u003e30.8\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 92px;\"\u003e\n \u003cp\u003e37.9\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 92px;\"\u003e\n \u003cp\u003e30.3\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 92px;\"\u003e\n \u003cp\u003e1\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 92px;\"\u003e\n \u003cp\u003e\u003cstrong\u003eMountain\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 116px;\"\u003e\n \u003cp\u003e\u003cstrong\u003eAll\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 67px;\"\u003e\n \u003cp\u003e\u003cstrong\u003e7.9\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 92px;\"\u003e\n \u003cp\u003e\u003cstrong\u003e13.6\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 92px;\"\u003e\n \u003cp\u003e\u003cstrong\u003e19.8\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 92px;\"\u003e\n \u003cp\u003e\u003cstrong\u003e58.7\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 92px;\"\u003e\n \u003cp\u003eNorth boreal\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 116px;\"\u003e\n \u003cp\u003eDeciduous\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 67px;\"\u003e\n \u003cp\u003e64.8\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 92px;\"\u003e\n \u003cp\u003e28.8\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 92px;\"\u003e\n \u003cp\u003e6.4\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 92px;\"\u003e\n \u003cp\u003e0\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 92px;\"\u003e\n \u003cp\u003eNorth boreal\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 116px;\"\u003e\n \u003cp\u003eHardwood\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 67px;\"\u003e\n \u003cp\u003e0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 92px;\"\u003e\n \u003cp\u003e0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 92px;\"\u003e\n \u003cp\u003e0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 92px;\"\u003e\n \u003cp\u003e0\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 92px;\"\u003e\n \u003cp\u003eNorth boreal\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 116px;\"\u003e\n \u003cp\u003eMixed\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 67px;\"\u003e\n \u003cp\u003e77.4\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 92px;\"\u003e\n \u003cp\u003e22\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 92px;\"\u003e\n \u003cp\u003e0.6\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 92px;\"\u003e\n \u003cp\u003e0\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 92px;\"\u003e\n \u003cp\u003eNorth boreal\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 116px;\"\u003e\n \u003cp\u003ePine\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 67px;\"\u003e\n \u003cp\u003e53.9\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 92px;\"\u003e\n \u003cp\u003e40.8\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 92px;\"\u003e\n \u003cp\u003e5.3\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 92px;\"\u003e\n \u003cp\u003e0\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 92px;\"\u003e\n \u003cp\u003eNorth boreal\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 116px;\"\u003e\n \u003cp\u003eSpruce \u0026amp; mixed\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 67px;\"\u003e\n \u003cp\u003e77.4\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 92px;\"\u003e\n \u003cp\u003e20.3\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 92px;\"\u003e\n \u003cp\u003e2.3\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 92px;\"\u003e\n \u003cp\u003e0\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 92px;\"\u003e\n \u003cp\u003e\u003cstrong\u003eNorth boreal\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 116px;\"\u003e\n \u003cp\u003e\u003cstrong\u003eAll\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 67px;\"\u003e\n \u003cp\u003e\u003cstrong\u003e42.7\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 92px;\"\u003e\n \u003cp\u003e\u003cstrong\u003e45.7\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 92px;\"\u003e\n \u003cp\u003e\u003cstrong\u003e11.3\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 92px;\"\u003e\n \u003cp\u003e\u003cstrong\u003e0.2\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 92px;\"\u003e\n \u003cp\u003eSouth boreal\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 116px;\"\u003e\n \u003cp\u003eDeciduous\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 67px;\"\u003e\n \u003cp\u003e80\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 92px;\"\u003e\n \u003cp\u003e16.1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 92px;\"\u003e\n \u003cp\u003e3.9\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 92px;\"\u003e\n \u003cp\u003e0\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 92px;\"\u003e\n \u003cp\u003eSouth boreal\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 116px;\"\u003e\n \u003cp\u003eHardwood\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 67px;\"\u003e\n \u003cp\u003e100\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 92px;\"\u003e\n \u003cp\u003e0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 92px;\"\u003e\n \u003cp\u003e0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 92px;\"\u003e\n \u003cp\u003e0\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 92px;\"\u003e\n \u003cp\u003eSouth boreal\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 116px;\"\u003e\n \u003cp\u003eMixed\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 67px;\"\u003e\n \u003cp\u003e88.9\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 92px;\"\u003e\n \u003cp\u003e11.1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 92px;\"\u003e\n \u003cp\u003e0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 92px;\"\u003e\n \u003cp\u003e0\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 92px;\"\u003e\n \u003cp\u003eSouth boreal\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 116px;\"\u003e\n \u003cp\u003ePine\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 67px;\"\u003e\n \u003cp\u003e61.9\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 92px;\"\u003e\n \u003cp\u003e34.7\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 92px;\"\u003e\n \u003cp\u003e3.4\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 92px;\"\u003e\n \u003cp\u003e0\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 92px;\"\u003e\n \u003cp\u003eSouth boreal\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 116px;\"\u003e\n \u003cp\u003eSpruce \u0026amp; mixed\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 67px;\"\u003e\n \u003cp\u003e88.5\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 92px;\"\u003e\n \u003cp\u003e11.5\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 92px;\"\u003e\n \u003cp\u003e0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 92px;\"\u003e\n \u003cp\u003e0\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 92px;\"\u003e\n \u003cp\u003e\u003cstrong\u003eSouth boreal\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 116px;\"\u003e\n \u003cp\u003e\u003cstrong\u003eAll\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 67px;\"\u003e\n \u003cp\u003e\u003cstrong\u003e54.9\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 92px;\"\u003e\n \u003cp\u003e\u003cstrong\u003e39.8\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 92px;\"\u003e\n \u003cp\u003e\u003cstrong\u003e5.2\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 92px;\"\u003e\n \u003cp\u003e\u003cstrong\u003e0.2\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 92px;\"\u003e\n \u003cp\u003eHemiboreal\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 116px;\"\u003e\n \u003cp\u003eDeciduous\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 67px;\"\u003e\n \u003cp\u003e75.2\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 92px;\"\u003e\n \u003cp\u003e24.8\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 92px;\"\u003e\n \u003cp\u003e0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 92px;\"\u003e\n \u003cp\u003e0\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 92px;\"\u003e\n \u003cp\u003eHemiboreal\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 116px;\"\u003e\n \u003cp\u003eHardwood\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 67px;\"\u003e\n \u003cp\u003e100\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 92px;\"\u003e\n \u003cp\u003e0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 92px;\"\u003e\n \u003cp\u003e0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 92px;\"\u003e\n \u003cp\u003e0\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 92px;\"\u003e\n \u003cp\u003eHemiboreal\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 116px;\"\u003e\n \u003cp\u003eMixed\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 67px;\"\u003e\n \u003cp\u003e96.6\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 92px;\"\u003e\n \u003cp\u003e3.4\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 92px;\"\u003e\n \u003cp\u003e0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 92px;\"\u003e\n \u003cp\u003e0\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 92px;\"\u003e\n \u003cp\u003eHemiboreal\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 116px;\"\u003e\n \u003cp\u003ePine\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 67px;\"\u003e\n \u003cp\u003e56.6\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 92px;\"\u003e\n \u003cp\u003e39.2\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 92px;\"\u003e\n \u003cp\u003e3.8\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 92px;\"\u003e\n \u003cp\u003e0.5\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 92px;\"\u003e\n \u003cp\u003eHemiboreal\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 116px;\"\u003e\n \u003cp\u003eSpruce \u0026amp; mixed\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 67px;\"\u003e\n \u003cp\u003e90\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 92px;\"\u003e\n \u003cp\u003e10\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 92px;\"\u003e\n \u003cp\u003e0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 92px;\"\u003e\n \u003cp\u003e0\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 92px;\"\u003e\n \u003cp\u003e\u003cstrong\u003eHemiboreal\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 116px;\"\u003e\n \u003cp\u003e\u003cstrong\u003eAll\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 67px;\"\u003e\n \u003cp\u003e\u003cstrong\u003e56.3\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 92px;\"\u003e\n \u003cp\u003e\u003cstrong\u003e39.1\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 92px;\"\u003e\n \u003cp\u003e\u003cstrong\u003e4.2\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 92px;\"\u003e\n \u003cp\u003e\u003cstrong\u003e0.4\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 92px;\"\u003e\n \u003cp\u003eNemoral\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 116px;\"\u003e\n \u003cp\u003eDeciduous\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 67px;\"\u003e\n \u003cp\u003e71\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 92px;\"\u003e\n \u003cp\u003e29\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 92px;\"\u003e\n \u003cp\u003e0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 92px;\"\u003e\n \u003cp\u003e0\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 92px;\"\u003e\n \u003cp\u003eNemoral\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 116px;\"\u003e\n \u003cp\u003eHardwood\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 67px;\"\u003e\n \u003cp\u003e100\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 92px;\"\u003e\n \u003cp\u003e0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 92px;\"\u003e\n \u003cp\u003e0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 92px;\"\u003e\n \u003cp\u003e0\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 92px;\"\u003e\n \u003cp\u003eNemoral\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 116px;\"\u003e\n \u003cp\u003eMixed\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 67px;\"\u003e\n \u003cp\u003e88.5\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 92px;\"\u003e\n \u003cp\u003e11.5\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 92px;\"\u003e\n \u003cp\u003e0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 92px;\"\u003e\n \u003cp\u003e0\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 92px;\"\u003e\n \u003cp\u003eNemoral\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 116px;\"\u003e\n \u003cp\u003ePine\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 67px;\"\u003e\n \u003cp\u003e65.5\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 92px;\"\u003e\n \u003cp\u003e34.5\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 92px;\"\u003e\n \u003cp\u003e0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 92px;\"\u003e\n \u003cp\u003e0\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 92px;\"\u003e\n \u003cp\u003eNemoral\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 116px;\"\u003e\n \u003cp\u003eSpruce \u0026amp; mixed\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 67px;\"\u003e\n \u003cp\u003e68\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 92px;\"\u003e\n \u003cp\u003e32\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 92px;\"\u003e\n \u003cp\u003e0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 92px;\"\u003e\n \u003cp\u003e0\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 92px;\"\u003e\n \u003cp\u003e\u003cstrong\u003eNemoral\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 116px;\"\u003e\n \u003cp\u003e\u003cstrong\u003eAll\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 67px;\"\u003e\n \u003cp\u003e\u003cstrong\u003e64.2\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 92px;\"\u003e\n \u003cp\u003e\u003cstrong\u003e35.8\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 92px;\"\u003e\n \u003cp\u003e\u003cstrong\u003e0\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 92px;\"\u003e\n \u003cp\u003e\u003cstrong\u003e0\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003c/tbody\u003e\n\u003c/table\u003e\n\u003ch3 id=\"_Toc189659350\"\u003eFunctional connectivity\u003c/h3\u003e\n\u003cp\u003eThe analysis of functional connectivity shows substantial differences in the contribution of LPFs in improving habitat network functionality across different forest types, ecoregions in Sweden, and for less demanding generalist versus more demanding specialist species (Table 4).\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003e\u003cem\u003eTable 4\u003c/em\u003e\u003c/strong\u003e\u003cem\u003e\u0026nbsp;The addition in percent units of LPFs area to existing HCVF in enhancing functionality of habitat networks for different forest types and forest ecoregions. G = generalist species; S = specialist species. NA denotes that the forest type is missing in the region\u003c/em\u003e\u003c/p\u003e\n\u003ctable border=\"0\" cellspacing=\"0\" cellpadding=\"0\" align=\"left\" width=\"100%\"\u003e\n \u003ctbody\u003e\n \u003ctr\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 19px;\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003ctd colspan=\"2\" valign=\"top\" style=\"width: 12px;\"\u003e\n \u003cp\u003e\u003cstrong\u003e\u003cem\u003eAll forest\u003c/em\u003e\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" colspan=\"2\" valign=\"bottom\" style=\"width: 12px;\"\u003e\n \u003cp\u003e\u003cstrong\u003e\u003cem\u003ePine\u003c/em\u003e\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" colspan=\"2\" valign=\"bottom\" style=\"width: 12px;\"\u003e\n \u003cp\u003e\u003cstrong\u003e\u003cem\u003eSpruce\u003c/em\u003e\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" colspan=\"2\" valign=\"bottom\" style=\"width: 11px;\"\u003e\n \u003cp\u003e\u003cstrong\u003e\u003cem\u003eMixed\u003c/em\u003e\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" colspan=\"2\" valign=\"bottom\" style=\"width: 17px;\"\u003e\n \u003cp\u003e\u003cstrong\u003e\u003cem\u003eDeciduous\u003c/em\u003e\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" colspan=\"2\" valign=\"bottom\" style=\"width: 14px;\"\u003e\n \u003cp\u003e\u003cstrong\u003e\u003cem\u003eHardwood\u003c/em\u003e\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 19px;\"\u003e\n \u003cp\u003e\u003cstrong\u003e\u003cem\u003eRegion\u003c/em\u003e\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 5px;\"\u003e\n \u003cp\u003eG\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 6px;\"\u003e\n \u003cp\u003eS\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 5px;\"\u003e\n \u003cp\u003eG\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 6px;\"\u003e\n \u003cp\u003eS\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 6px;\"\u003e\n \u003cp\u003eG\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 6px;\"\u003e\n \u003cp\u003eS\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 5px;\"\u003e\n \u003cp\u003eG\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 5px;\"\u003e\n \u003cp\u003eS\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 8px;\"\u003e\n \u003cp\u003eG\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 8px;\"\u003e\n \u003cp\u003eS\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 8px;\"\u003e\n \u003cp\u003eG\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 6px;\"\u003e\n \u003cp\u003eS\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 19px;\"\u003e\n \u003cp\u003eMountain\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 5px;\"\u003e\n \u003cp\u003e0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 6px;\"\u003e\n \u003cp\u003e1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 5px;\"\u003e\n \u003cp\u003e0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 6px;\"\u003e\n \u003cp\u003e1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 6px;\"\u003e\n \u003cp\u003e0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 6px;\"\u003e\n \u003cp\u003e0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 5px;\"\u003e\n \u003cp\u003e0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 5px;\"\u003e\n \u003cp\u003e0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 8px;\"\u003e\n \u003cp\u003e1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 8px;\"\u003e\n \u003cp\u003e1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 8px;\"\u003e\n \u003cp\u003eNA\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 6px;\"\u003e\n \u003cp\u003eNA\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 19px;\"\u003e\n \u003cp\u003eNorth boreal\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 5px;\"\u003e\n \u003cp\u003e6\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 6px;\"\u003e\n \u003cp\u003e15\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 5px;\"\u003e\n \u003cp\u003e6\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 6px;\"\u003e\n \u003cp\u003e11\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 6px;\"\u003e\n \u003cp\u003e1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 6px;\"\u003e\n \u003cp\u003e1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 5px;\"\u003e\n \u003cp\u003e2\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 5px;\"\u003e\n \u003cp\u003e3\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 8px;\"\u003e\n \u003cp\u003e1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 8px;\"\u003e\n \u003cp\u003e1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 8px;\"\u003e\n \u003cp\u003eNA\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 6px;\"\u003e\n \u003cp\u003eNA\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 19px;\"\u003e\n \u003cp\u003eSouth boreal\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 5px;\"\u003e\n \u003cp\u003e8\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 6px;\"\u003e\n \u003cp\u003e16\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 5px;\"\u003e\n \u003cp\u003e8\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 6px;\"\u003e\n \u003cp\u003e13\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 6px;\"\u003e\n \u003cp\u003e1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 6px;\"\u003e\n \u003cp\u003e2\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 5px;\"\u003e\n \u003cp\u003e0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 5px;\"\u003e\n \u003cp\u003e1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 8px;\"\u003e\n \u003cp\u003e1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 8px;\"\u003e\n \u003cp\u003e0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 8px;\"\u003e\n \u003cp\u003e0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 6px;\"\u003e\n \u003cp\u003e0\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 19px;\"\u003e\n \u003cp\u003eHemiboreal\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 5px;\"\u003e\n \u003cp\u003e6\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 6px;\"\u003e\n \u003cp\u003e14\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 5px;\"\u003e\n \u003cp\u003e8\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 6px;\"\u003e\n \u003cp\u003e16\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 6px;\"\u003e\n \u003cp\u003e0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 6px;\"\u003e\n \u003cp\u003e1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 5px;\"\u003e\n \u003cp\u003e1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 5px;\"\u003e\n \u003cp\u003e0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 8px;\"\u003e\n \u003cp\u003e1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 8px;\"\u003e\n \u003cp\u003e1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 8px;\"\u003e\n \u003cp\u003e0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 6px;\"\u003e\n \u003cp\u003e0\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 19px;\"\u003e\n \u003cp\u003eNemoral\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 5px;\"\u003e\n \u003cp\u003e2\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 6px;\"\u003e\n \u003cp\u003e3\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 5px;\"\u003e\n \u003cp\u003e5\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 6px;\"\u003e\n \u003cp\u003e6\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 6px;\"\u003e\n \u003cp\u003e1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 6px;\"\u003e\n \u003cp\u003e0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 5px;\"\u003e\n \u003cp\u003e0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 5px;\"\u003e\n \u003cp\u003e0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 8px;\"\u003e\n \u003cp\u003e0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 8px;\"\u003e\n \u003cp\u003e0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 8px;\"\u003e\n \u003cp\u003e0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd nowrap=\"\" valign=\"bottom\" style=\"width: 6px;\"\u003e\n \u003cp\u003e0\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003c/tbody\u003e\n\u003c/table\u003e\n\u003cp\u003e\u0026nbsp;\u003c/p\u003e\n\u003cp\u003eGiven the high density of HCVF and its large overlap with LPF in the mountain region, the inclusion of LPF provides very limited additional GI functionality across forest types. For pine forests below the mountain region, the inclusion of LPFs led to an increase of 5-8% in area of functional HCVF habitat for forest generalist species across ecoregions. Adding LPFs for pine forest specialists showed a higher increase, with figures ranging from 6-16%, notably in the south boreal and hemiboreal ecoregions with 13% and 16% increased connectivity, respectively. The contribution of pine forest to functional connectivity is clearly reflected in the patterns for all forest land although the fraction of connected habitats is obviously larger when including all forest types. For the other four forest types, there is minimal to no improvement in functional connectivity by incorporating LPFs, with figures ranging from 0-3%. The improved connectivity for all forest types and for pine forests caused by adding LPFs is illustrated by the hotspot-coldspot maps in Figure 4 (for the other forest types see Appendix 1).\u003c/p\u003e"},{"header":"Discussion","content":"\u003cp\u003eIn forest-dominated landscapes, effective green infrastructure requires functional HCVF-habitat networks. In this study, the contribution of LPFs to habitat functionality was assessed based on their occurrence across Sweden, their patch size distribution and in relation to functional habitat networks of two virtual species representing different network demands. For all these three aspects the contribution of LPFs varied strongly between different forest types and different ecoregions. In the mountain region, LPFs dominates the landscape but given the high share of already identified HCVF forests their additional contribution to habitat functionality is small. Outside the mountain region it is mainly in pine forests that LPFs provide improved connectivity. This is logical since pine forests represent the dominant LPF type outside the mountain region due to the ability of pine to thrive on very poor site types on both dry mineral (rocky outcrops) and mesic to wet mineral and peat organic soils.\u003c/p\u003e \u003cp\u003eThe areal extent of LPFs in Sweden follows a clear south \u0026ndash; northwest gradient mainly reflecting macroclimatic conditions. The vast majority of the LPFs occur in the mountain and boreal ecoregions. Although they do occur in both the hemiboreal region and nemoral region they represent less than 10% of the forest area. Hence, the role of LPFs for improving forest GI varies between biogeographical ecoregions and dominant tree species (forest types). For forests dominated by pine in the mountain ecoregion, parts of the boreal ecoregions and in the eastern part of the hemiboreal ecoregions, they do however contribute to GI functionality by increasing the functional connectivity of forest patches (Fig.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e4\u003c/span\u003e). For other ecoregions, LPFs derive much of their ecological importance as isolated patches providing habitat for species specialised on low-productive forest conditions with natural structural characteristics (H\u0026auml;m\u0026auml;l\u0026auml;inen et al. \u003cspan citationid=\"CR19\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). It should be noted, however, that in the mountain forest region most of the LFP are already identified as HCVF and included in functional networks (Svensson et al. \u003cspan citationid=\"CR45\" class=\"CitationRef\"\u003e2020\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eThe size distribution of LPFs varies across the south-north gradient. Overall, most of the LPFs occur as small fragments (\u0026lt;\u0026thinsp;5 ha), particularly when considering individual forest types. Only for deciduous forest in the mountain region, larger landscapes were dominated by LPFs, a pattern identified in earlier studies (Angelstam et al. \u003cspan citationid=\"CR5\" class=\"CitationRef\"\u003e2020\u003c/span\u003e, \u003cspan citationid=\"CR6\" class=\"CitationRef\"\u003e2026\u003c/span\u003e; Mikusiński et al. \u003cspan citationid=\"CR28\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). In contrast, and especially for the more southern ecoregions, most LPFs occurred in fragments smaller than 5 ha. This fragmentation is natural and primarily caused by variations in local forest site productivity, rather than habitat loss caused by human activity (e.g. Svensson et al. \u003cspan citationid=\"CR44\" class=\"CitationRef\"\u003e2019\u003c/span\u003e). Hence, it is likely that some species with very small area requirements may have favourable conservation status as long as edge effects and climate change remain limited.\u003c/p\u003e \u003cp\u003eThe analysis of functional connectivity further emphasizes a relatively limited contribution of LPFs. Obviously, the chosen examples of virtual species do not reflect the full gradient of habitat demands of all forest species. However, they do represent relevant examples of habitat demands and landscape connectivity and a commonly used threshold (20%) for amount of habitat at landscape scale (Hanski \u003cspan citationid=\"CR21\" class=\"CitationRef\"\u003e2011\u003c/span\u003e; Svancara et al. \u003cspan citationid=\"CR39\" class=\"CitationRef\"\u003e2005\u003c/span\u003e). It may look contradictory that the increase in connectivity by adding LPFs to the network was higher for specialist species, but it should be noted that that total area of functional networks is consistently higher for generalist species. It is also clear from the analysis that the contribution of LPFs is only evident for forests in general and basically mirrors the contribution of for pine dominated LPFs. For all other forest types, the contribution is negligible.\u003c/p\u003e \u003cp\u003eWhile this study addresses how LPFs contribute to GI, there are several other potential contributions to GI functionality, including formally protected areas, voluntary set asides and nature considerations applied during forest operations through retention practices. Although compilations of the area extents of these categories exists, as their levels of naturalness and other conservation qualities differ one cannot simply add the total areas. Thus, the extent to which conservation targets are met depends on not only on quantities (Angelstam et al. \u003cspan citationid=\"CR5\" class=\"CitationRef\"\u003e2020\u003c/span\u003e, \u003cspan citationid=\"CR6\" class=\"CitationRef\"\u003e2026\u003c/span\u003e) as an area\u0026rsquo;s effectiveness depends on i) quality in terms of degree of naturalness (Puettman et al. 2012; Winter 2012), ii) patch size related to species requirement (Angelstam et al. 2004; Wiens and Milne 1989), iii) functional connectivity allowing species to move between suitable habitats in the landscape (Pither et al. 2023), and iv) longevity in relation to natural disturbances and succession dynamics (Rosenvald and L\u0026otilde;hmus 2023).\u003c/p\u003e \u003cp\u003eIn the light of the agreed policy targets for protected areas and with their specification to be particularly important for biodiversity through securing functional networks, we question whether LPFs by default should be seen as a contribution to fulfilling conservation targets albeit being legally excluded from commercial forest management, particularly if only their area contribution is considered. In local landscapes and for species with small area requirements, however, LPFs may contribute to functional GI, but here we argue that including locally-relevant quality parameters on top of area is of key importance. In this context, even small pine-dominated LPFs may play an important role as restoration nodes, facilitating more effective recolonisation of restored habitats by specialised species (Mikusiński et al. \u003cspan citationid=\"CR28\" class=\"CitationRef\"\u003e2021\u003c/span\u003e; Ramberg et al. \u003cspan citationid=\"CR35\" class=\"CitationRef\"\u003e2026\u003c/span\u003e). This illustrates the importance of not just considering the area proportion protected to fulfil the intention of the international agreements, but also the particular roles of habitat quality, habitat patch size and functional connectivity for biodiversity conservation (cf. CBD \u003cspan citationid=\"CR11\" class=\"CitationRef\"\u003e2022\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eGreen infrastructure, like conservation networks more broadly, should be ecologically representative of the ecoregions in which it is implemented (Pressey et al. \u003cspan citationid=\"CR34\" class=\"CitationRef\"\u003e2007\u003c/span\u003e). LPFs in Sweden are not fully representative for all forest types, both for natural reasons\u0026mdash;because they encompass only a narrow segment of the forest productivity gradient\u0026mdash;and for socio-economic reasons, as they have often been protected due to their comparatively low economic value. This skewed representation, constrains their broader contribution to forest biodiversity conservation in Sweden. Still, naturally multilayered pine forests, the main LPF type, have become rare due to intensive forest management and fire suppression (Berglund and Kuuluvainen \u003cspan citationid=\"CR8\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). Recent research suggests that the historical extent of such forests in northern European dynamic forest landscapes was greater than previously assumed (ibid.). This implies substantial restoration needs.\u003c/p\u003e \u003cp\u003eTo conclude, our analysis show that although LPFs may strengthen GI in some ecoregions and for some forest types they cannot replace the need for additional protection and restoration of productive forests with high conservation value to ensure functional connectivity. Hence, the role of LPFs as a contribution to national and international conservation objectives is limited and mainly related to species that are confined to these island habitats. Yet they contribute to landscape heterogeneity and may hold aesthetic and recreational values.\u003c/p\u003e"},{"header":"Declarations","content":"\u003cp\u003e \u003ch2\u003eCompeting Interests\u003c/h2\u003e \u003cp\u003eThe authors have no relevant financial or non-financial interests to disclose\u003c/p\u003e \u003c/p\u003e\u003ch2\u003eFunding\u003c/h2\u003e \u003cp\u003eThis work was supported by Swedish Environmental Protection Agency, grant NV-03728\u0026ndash;17, to Bengt Gunnar Jonsson, Horizon 2020 Research and Innovation Programme Grant Agreement No. 101036849 to Johan Svensson and Formas grant 2022\u0026ndash;02146 to Per Angelstam\u003c/p\u003e\u003ch2\u003eAuthor Contribution\u003c/h2\u003e\u003cp\u003eAll authors contributed to the study conception and design. Material preparation and analysis were performed by Jakub Bubnicki. The first draft of the manuscript was written by Bengt Gunnar Jonsson and all authors commented on previous versions of the manuscript. All authors read and approved the final manuscript\u003c/p\u003e\u003ch2\u003eAcknowledgments\u003c/h2\u003e \u003cp\u003eThis work was supported by Swedish Environmental Protection Agency, grant NV-03728\u0026ndash;17, to Bengt Gunnar Jonsson, Formas Research Council, grant 2022\u0026ndash;02146_Formas to Per Angelstam and Horizon 2020 Research Innovation Programme (SUPERB: Upscaling Forest Restoration\u0026mdash;SUPERB (forest-restoration.eu)); Grant Agreement No. 101036849 to Johan Svensson.\u003c/p\u003e"},{"header":"References","content":"\u003col\u003e\n\u003cli\u003eAhti T, H\u0026auml;met-Ahti L, Jalas, J (1968) Vegetation zones and their sections in northwestern Europe. Annales Botanici Fennici 5:69\u0026ndash;211\u003c/li\u003e\n\u003cli\u003eAngelstam P, Andersson M, Isacson DV et al (2013) Learning about the history of landscape use for the future: consequences for ecological and social systems in Swedish Bergslagen. Ambio 42:150\u0026ndash;163. https://doi.org/10.1007/s13280-012-0369-z\u003c/li\u003e\n\u003cli\u003eAngelstam P, Asplund B, Bastian O et al (2022) Tradition as asset or burden for transitions from forests as cropping systems to multifunctional forest landscapes: Sweden as a case study. Forest Ecology and Management 505:119895 https://doi.org/10.1016/j.foreco.2021.119895\u003c/li\u003e\n\u003cli\u003eAngelstam P, Bush T, Manton M (2023) Challenges and solutions for forest biodiversity conservation in Sweden: Assessment of policy, implementation outputs, and consequences. Land 12:1098. https://doi.org/10.3390/land12051098\u003c/li\u003e\n\u003cli\u003eAngelstam P, Manton M, Green M, Jonsson BG, Mikusiński G, Svensson J, Sabatini FM (2020) Sweden does not meet agreed national and international forest biodiversity targets: A call for adaptive landscape planning. Landscape and Urban Planning 202:103838. https://doi.org/10.1016/j.landurbplan.2020.103838\u003c/li\u003e\n\u003cli\u003eAngelstam P, Yamelynets T, Manton M, Bubnicki JW, Jonsson BG, Mikusinski G, Svensson J, Dawson L (2026) Environmental history determines forest habitat network functionality: The need for landscape planning in Sweden. Ambio. https://doi.org/10.1007/s13280-026-02353-7\u003c/li\u003e\n\u003cli\u003eBerglund B (1992) Landscape reconstructions in South Sweden for the past 6000 years. Proceeding of the British Academy 19:25-37\u003c/li\u003e\n\u003cli\u003eBerglund H, Kuuluvainen T (2021) Representative boreal forest habitats in northern Europe, and a revised model for ecosystem management and biodiversity conservation. Ambio 50:1003\u0026ndash;1017. https://doi.org/10.1007/s13280-020-01444-3\u003c/li\u003e\n\u003cli\u003eBladh G (1995) Finnskogens landskap och m\u0026auml;nniskor under fyra sekler. Forskningsrapport 95:11 H\u0026ouml;gskolan i Karlstad\u003c/li\u003e\n\u003cli\u003eCBD (2010) Strategic plan for biodiversity 2011\u0026ndash;2020 and the Aichi targets. Convention on Biological Diversity, Montreal. Available from https://www.cbd.int/sp/targets/(accessed February 2026)\u003c/li\u003e\n\u003cli\u003eCBD (2022) Kunming-Montreal Global Biodiversity Framework. Convention of Biological https://www.cbd.int/gbf (Accessed February 2026)\u003c/li\u003e\n\u003cli\u003eCEC (1992) Council Directive 92/43/EEC of 21 730 May 1992 on the conservation of natural habitats and of wild fauna and flora. Council of the European Communities, Official 731 Journal of the European Communities 22.7.92, no L 206/7-50\u003c/li\u003e\n\u003cli\u003eEC (2013) Green Infrastructure (GI) \u0026mdash; Enhancing Europe\u0026rsquo;s natural capital. European Commission, Brussels\u003c/li\u003e\n\u003cli\u003eEC (2020) EU Biodiversity strategy for 2030\u0026mdash;Bringing nature back into our lives. European Commission, Brussels\u003c/li\u003e\n\u003cli\u003eForrester DI, Bauhus J (2016) A Review of Processes Behind Diversity\u0026mdash;Productivity Relationships in Forests. Curr Forestry Rep 2:45\u0026ndash;61. https://doi.org/10.1007/s40725-016-0031-2\u003c/li\u003e\n\u003cli\u003eGillman LN, Wright SD (2006) The influence of productivity on the species richness of plants: a critical assessment. Ecology 87:1234-1243. https://doi.org/10.1890/0012-9658(2006)87[1234:TIOPOT]2.0.CO;2\u003c/li\u003e\n\u003cli\u003eGRASS Development Team (2025) Geographic Resources Analysis Support System (GRASS) Software, Version 8.4. Open Source Geospatial Foundation. https://grass.osgeo.org, DOI: 10.5281/zenodo.5176030\u003c/li\u003e\n\u003cli\u003eGustafsson L, Andersson J, Jonsson F, J\u0026ouml;nsson M, Jonsson M, Nordin U, Strengbom J, Johansson V (2025) Remnant continuity forests are essential for sustaining epiphytic biodiversity in boreal production forest landscapes. J Appl Ecol. https://doi.org/10.1111/1365-2664.70090\u003c/li\u003e\n\u003cli\u003eH\u0026auml;m\u0026auml;l\u0026auml;inen A, Strengbom J, Ranius T (2020) Low-productivity boreal forests have high conservation value for lichens. Journal of Applied Ecology 57:43-54. https://doi.org/10.1111/1365-2664.13509\u003c/li\u003e\n\u003cli\u003eH\u0026auml;m\u0026auml;l\u0026auml;inen A, Runnel K, Ranius T, Strengbom J (2024) Diversity of forest structures important for biodiversity is determined by the combined effects of productivity, stand age, and management. Ambio 53:718\u0026ndash;729. https://doi.org/10.1007/s13280-023-01971-9\u003c/li\u003e\n\u003cli\u003eHanski I (2011) Habitat Loss, the Dynamics of Biodiversity, and a Perspective on Conservation. Ambio 40:248-255. DOI: 10.1007/s13280-011-0147-3\u003c/li\u003e\n\u003cli\u003eJoks M, Helm A, Kasari-Toussaint L et al (2023) A simulation model of functional habitat connectivity demonstrates the importance of species establishment in older forests. Ecological Modelling 481:110361. https://doi.org/10.1016/j.ecolmodel.2023.110361\u003c/li\u003e\n\u003cli\u003eJonsson BG, Svensson J, Mikusinski G, Manton M, Angelstam P (2019) European Union\u0026rsquo;s last intact forest landscapes are at a value chain crossroad between multiple use and intensified wood production. Forests 10:564. https://doi.org/10.3390/f10070564 \u003c/li\u003e\n\u003cli\u003eKeskitalo C, Naumov V, Ahlkrona E, J\u0026ouml;nsson C, Olsson B (2019) Nationella markt\u0026auml;ckedata till\u0026auml;ggsskikt produktivitet. Naturv\u0026aring;rdsverket, Stockholm\u003c/li\u003e\n\u003cli\u003eK\u0026ouml;rner C (2021) Plant ecology at high elevations. In \u003cem\u003eAlpine plant life: functional plant ecology of high mountain ecosystems\u003c/em\u003e (pp. 1-22). Cham: Springer International Publishing. https://doi.org/10.1007/978-3-030-59538-8_1\u003c/li\u003e\n\u003cli\u003eKyaschenko J, Strengbom J, Felton A, Aakala T, Staland H, Ranius T (2022) Increase in dead wood, large living trees and tree diversity, yet decrease in understory vegetation cover: The effect of three decades of biodiversity-oriented forest policy in Swedish forests. Journal of Environmental Management, 313:114993. https://doi.org/10.1016/j.jenvman.2022.114993\u003c/li\u003e\n\u003cli\u003eMikkonen N, Leikola N, Lehtom\u0026auml;ki J, Halme P, Moilanen A (2023) National high-resolution conservation prioritisation of boreal forests. Forest Ecology and Management 541:121079. https://doi.org/10.1016/j.foreco.2023.121079\u003c/li\u003e\n\u003cli\u003eMikusiński G, Orlikowska EH, Bubnicki JW, Jonsson BG, Svensson J (2021). Strengthening the network of high conservation value forests in boreal landscapes. Frontiers in Ecology and Evolution 8:595730. https://doi.org/10.3389/fevo.2020.595730\u003c/li\u003e\n\u003cli\u003eNaumburg B (ed.) (2024) Forests and forestry in Sweden. Stockholm, The Royal Swedish Academy of Agriculture and Forestry\u003c/li\u003e\n\u003cli\u003eNilsson M, Ahlkrona E, J\u0026ouml;nsson C, Allard A (2020) Regionala j\u0026auml;mf\u0026ouml;relser mellan Nationella Markt\u0026auml;ckedata och f\u0026auml;ltdata fr\u0026aring;n Riksskogstaxeringen och NILS. SLU, Ume\u0026aring; and Metria, Stockholm\u003c/li\u003e\n\u003cli\u003eOrlikowska EH, Roberge JM, Svensson J, Mikusiński G (2020) Hit or miss? Evaluating Natura 2000 sites for conservation of forest bird habitat in Sweden. Global Ecology and Conservation 22, e00939. https://doi.org/10.1016/j.gecco.2020.e00939\u003c/li\u003e\n\u003cli\u003e\u0026Ouml;stlund L, Zackrisson L, Axelsson, AL (1997) The history and transformation of a Scandinavian boreal forest landscape since the 19th century. Can. J. For. Res. 27:1198\u0026ndash;1206. https://doi.org/10.1139/x97-070\u003c/li\u003e\n\u003cli\u003ePotapov P et al (2008) Mapping the World\u0026apos;s Intact Forest Landscapes by Remote Sensing. Ecology and Society 13:51. https://www.jstor.org/stable/26267984\u003c/li\u003e\n\u003cli\u003ePressey RL, Cabeza M, Watts ME, Cowling RM, Wilson KA (2007) Conservation planning in a changing world. Trends in Ecology \u0026amp; Evolution 22:583\u0026ndash;592. https://doi.org/10.1016/j.tree.2007.10.001\u003c/li\u003e\n\u003cli\u003eRamberg E, Edman M, Granath G, Sj\u0026ouml;gren J, Strengbom J (2026) Prescribed burning for boreal forest restoration: Evaluating challenges and conservation outcomes. Ambio 55:608\u0026ndash;619. https://doi.org/10.1007/s13280-025-02248-z\u003c/li\u003e\n\u003cli\u003eSandstr\u0026ouml;m J, Edman M, Jonsson BG (2020) Rocky pine forests in the High Coast Region in Sweden: structure, dynamics and history. Nature Conservation 38:101\u0026ndash;130. https://doi.org/10.3897/natureconservation.38.34870\u003c/li\u003e\n\u003cli\u003eSCB (2025) Formellt skyddad skogsmark, frivilliga avs\u0026auml;ttningar, h\u0026auml;nsynsytor samt improduktiv skogsmark 2024. MI 41 Rapport 2025:02\u003c/li\u003e\n\u003cli\u003eSLU (2025) Forest Statistics 2025. Swedish University of Agricultural Sciences, Ume\u0026aring;\u003c/li\u003e\n\u003cli\u003eSvancara LK, Brannon JR, Scott M, Groves CR, Noss RF, Pressey RL (2005) Policy-driven versus evidence-based conservation: a review of political targets and biological needs. BioScience 55:989-995. https://doi.org/10.1641/0006-3568(2005)055[0989:PVECAR]2.0.CO;2\u003c/li\u003e\n\u003cli\u003eSwedish EPA (2016) A database of High Conservation Value Forests in Sweden. Available online at https://geodata.naturvardsverket.se/nedladdning/land/skogliga_vardekarnor_2016.zip. Updated with new data from the mountain region 2019-2020\u003c/li\u003e\n\u003cli\u003eSwedish EPA (2019) National landcover database. https:// www.naturvardsverket.se/verktyg-och-tjanster/kartor-och-karttjanster/ nationella-marktackedata\u003c/li\u003e\n\u003cli\u003eSwedish Forest Agency (2022) Levande skogar - F\u0026ouml;rdjupad utv\u0026auml;rdering 2023. Skogsstyrelsen Rapport 2022/12\u003c/li\u003e\n\u003cli\u003eSwedish Forest Agency (2025) Skogsv\u0026aring;rdslagstiftningen \u0026ndash; G\u0026auml;llande regler 1 april 2025. Skogsstyrelsen (in Swedish)\u003c/li\u003e\n\u003cli\u003eSvensson J, Andersson J, Sandstr\u0026ouml;m P, Mikusiński G, Jonsson BG (2019) Landscape trajectory of natural boreal forest loss as an impediment to green infrastructure Cons. Biol. 33:152-163. https://doi.org/10.1111/cobi.13148\u003c/li\u003e\n\u003cli\u003eSvensson J, Bubnicki JW, Jonsson BG, Andersson J, Mikusiński G (2020) Conservation significance of intact forest landscapes in the Scandinavian Mountains Green Belt. Landscape Ecology 35:2113\u0026ndash;2131. https://doi.org/10.1007/s10980-020-01088-4\u003c/li\u003e\n\u003cli\u003eSvensson J, Wallin I, Droste N (2026) Makten \u0026ouml;ver skogen. Rapport 3, Skogsstyrelsen,\u003c/li\u003e\n\u003cli\u003eUliczka H, Angelstam P (1999) Occurrence of epiphytic macrolichens in relation to tree species and age in managed boreal forest. Ecography, 22: 396-405. https://www.jstor.org/stable/3683231\u003c/li\u003e\n\u003cli\u003eVillard MA, Hach\u0026eacute; S (2012) Conifer plantations consistently act as barriers to movement in a deciduous forest songbird: A translocation experiment. Biological Conservation 155:33-37. https://doi.org/10.1016/j.biocon.2012.06.007\u003c/li\u003e\n\u003cli\u003eWang X, Svensson J, Jonsson BG, Singh NJ, Bubnicki JW, Lop\u0026eacute;z-Peinado A, Angelstam P, Mikusiński G, Ard\u0026ouml; J (2025). Where to restore: connectivity forest for spatial prioritization in forest landscape restoration. iScience 28:113263. https://doi.org/10.1016/j.isci.2025.113263\u003c/li\u003e\n\u003c/ol\u003e"}],"fulltextSource":"","fullText":"","funders":[],"hasAdminPriorityOnWorkflow":false,"hasManuscriptDocX":true,"hasOptedInToPreprint":true,"hasPassedJournalQc":"","hasAnyPriority":false,"hideJournal":false,"highlight":"","institution":"","isAcceptedByJournal":false,"isAuthorSuppliedPdf":false,"isDeskRejected":"","isHiddenFromSearch":false,"isInQc":false,"isInWorkflow":false,"isPdf":false,"isPdfUpToDate":true,"isWithdrawnOrRetracted":false,"journal":{"display":true,"email":"[email protected]","identity":"landscape-ecology","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":false,"externalIdentity":"land","sideBox":"Learn more about [Landscape Ecology](https://www.springer.com/journal/10980)","snPcode":"10980","submissionUrl":"https://submission.nature.com/new-submission/10980/3","title":"Landscape Ecology","twitterHandle":"","acdcEnabled":true,"dfaEnabled":true,"editorialSystem":"em","reportingPortfolio":"Springer Hybrid","inReviewEnabled":true,"inReviewRevisionsEnabled":false},"keywords":"Biodiversity, Connectivity, Landscape planning, Habitat loss, Fragmentation, Policy implementation","lastPublishedDoi":"10.21203/rs.3.rs-9379486/v1","lastPublishedDoiUrl":"https://doi.org/10.21203/rs.3.rs-9379486/v1","license":{"name":"CC BY 4.0","url":"https://creativecommons.org/licenses/by/4.0/"},"manuscriptAbstract":"\u003ch2\u003eContext:\u003c/h2\u003e \u003cp\u003eGreen infrastructure (GI) is an EU policy concept addressing functional connectivity to support biodiversity conservation and ecosystem services. In regions with significant forest fragmentation, establishing functional GI presents a challenge.\u003c/p\u003e\u003ch2\u003eObjectives\u003c/h2\u003e \u003cp\u003eWe evaluate the contribution of low-productive forests (LPF) in networks of existing High Conservation Value Forests (HCVF) in Sweden, and discuss conservation implications for GI across different forest types and ecoregions.\u003c/p\u003e\u003ch2\u003eMethods\u003c/h2\u003e \u003cp\u003eThree characteristics for LPF were assessed: area extent of different forest types, patch size distribution, and role in functional landscape connectivity across five ecoregions. The primary data sources were a database of known HCVF and the national land cover data that includes data on LPF.\u003c/p\u003e\u003ch2\u003eResults\u003c/h2\u003e \u003cp\u003eLPF are dominant in the mountain region, both for all forests and among HCVF classified as low-productive. For other ecoregions, the extent of LPF was lower. Pine forests dominate LPF in all regions below the mountain region. For pine forests outside the mountain region, the inclusion of LPF increased the area of functional HCVF habitat for forest generalist species by 5\u0026ndash;8% and for forest specialists by 6\u0026ndash;16%. For other forest types, there was minimal improvement in connectivity. Overall, most LPF occur as small fragments, particularly when considering individual forest types.\u003c/p\u003e\u003ch2\u003eConclusion\u003c/h2\u003e \u003cp\u003eLPF can strengthen GI in some regions and some forest types. However, they cannot replace the need for additional protection and restoration of productive forests to ensure functional connectivity. The role of LPFs as a contribution to conservation objectives is limited and related to species confined to LPFs.\u003c/p\u003e","manuscriptTitle":"Limited contribution of low-productive forests to green infrastructure along a temperate to boreal biogeographic gradient","msid":"","msnumber":"","nonDraftVersions":[{"code":1,"date":"2026-04-30 18:37:57","doi":"10.21203/rs.3.rs-9379486/v1","editorialEvents":[{"type":"communityComments","content":0},{"type":"reviewerAgreed","content":"217605055562635144924821357659940962332","date":"2026-05-18T15:31:03+00:00","index":"hide","fulltext":""},{"type":"reviewerAgreed","content":"226064936498669825590829421721607565564","date":"2026-05-18T06:42:54+00:00","index":"hide","fulltext":""},{"type":"reviewersInvited","content":"","date":"2026-04-22T15:57:23+00:00","index":"","fulltext":""},{"type":"editorAssigned","content":"","date":"2026-04-13T11:53:40+00:00","index":"","fulltext":""},{"type":"checksComplete","content":"","date":"2026-04-13T11:53:15+00:00","index":"","fulltext":""},{"type":"submitted","content":"Landscape Ecology","date":"2026-04-10T12:16:29+00:00","index":"","fulltext":""}],"status":"published","journal":{"display":true,"email":"[email protected]","identity":"landscape-ecology","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":false,"externalIdentity":"land","sideBox":"Learn more about [Landscape Ecology](https://www.springer.com/journal/10980)","snPcode":"10980","submissionUrl":"https://submission.nature.com/new-submission/10980/3","title":"Landscape Ecology","twitterHandle":"","acdcEnabled":true,"dfaEnabled":true,"editorialSystem":"em","reportingPortfolio":"Springer Hybrid","inReviewEnabled":true,"inReviewRevisionsEnabled":false}}],"origin":"","ownerIdentity":"f0b52a0d-d0ef-4e15-81fa-b6046b0b7df9","owner":[],"postedDate":"April 30th, 2026","published":true,"recentEditorialEvents":[{"type":"reviewerAgreed","content":"217605055562635144924821357659940962332","date":"2026-05-18T15:31:03+00:00","index":29,"fulltext":""},{"type":"reviewerAgreed","content":"226064936498669825590829421721607565564","date":"2026-05-18T06:42:54+00:00","index":28,"fulltext":""}],"rejectedJournal":[],"revision":"","amendment":"","status":"under-review","subjectAreas":[],"tags":[],"updatedAt":"2026-04-30T18:37:57+00:00","versionOfRecord":[],"versionCreatedAt":"2026-04-30 18:37:57","video":"","vorDoi":"","vorDoiUrl":"","workflowStages":[]},"version":"v1","identity":"rs-9379486","journalConfig":"researchsquare"},"__N_SSP":true},"page":"/article/[identity]/[[...version]]","query":{"redirect":"/article/rs-9379486","identity":"rs-9379486","version":["v1"]},"buildId":"XKTyCvWXoU3ODBz1xrDgd","isFallback":false,"isExperimentalCompile":false,"dynamicIds":[84888],"gssp":true,"scriptLoader":[]}