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SettingThis study was conducted in approximately 40-year-old naturally regenerated P. densiflora stands in Wola National Experimental Forest in Gyeongnam province in South Korea (35°12′ N, 128°10′ E; Table 1). The productivity of this forest is low, with a dominant tree height of 10 m at 20 years of age. Over the last 10 years, the mean annual precipitation was 1490 mm, of which one third fell during summer (July–August), and the mean temperature was 13.1 °C. The vegetation growing season generally lasts for approximately 200 days, extending from early April to October. The soil texture is a silt loam originating from sandstone and shale (clay 13.0 ± 0.8%, silt 44.1 ± 1.3%, sand 42.9 ± 1.6%; n = 18). The given texture results in volumetric water contents at 13.4 ± 0.7% (m3 m−3) at permanent wilting point (1500 kPa) and 40.7 ± 1.2% at field capacity (10 kPa)55. The understory is covered with Lespedeza spp., Quercus variabilis, Q. serrata, Smilax china, and Lindera glauca.In 2010, we selected two adjacent P. densiflora stands approximately 100 m apart from each other (180 m and 195 m above sea level, on slopes of 15° and 33°, both stands face south). Following a completely randomized design, we established nine plots (10 × 10 m2 with a 5 m untreated buffer) within each stand, of which three were randomly assigned to annual NPK fertilization, three to PK fertilization, and the rest to a control treatment without fertilization. The fertilizer, composed of urea, fused superphosphate and potassium chloride (N3P4K1) or P4K1 was added manually by deposition on the forest floor for 3 years in April 2011, April 2012, and March 2013. Over these 3 years, the NPK plots received 33.9 g N, 45 g P, and 11.1 g K m−2, while the PK plots received 45 g P and 11.1 g K m−2.Tree and stand measurementsThe standing biomass of trees was estimated using a combination of site-specific allometric equations based on destructive harvesting56 and repeated measurements of the dimensions of all trees in each plot (5–18 trees plot−1). The stem diameter at 1.2 m (D) was measured for all trees in each plot for which D was ≥ 6 cm. Selecting a representative tree in size for each plot within the 4 × 4 m2 center of the plot, we measured the tree height (H) and crown base for the representative trees. Measurements were performed in April and September 2011, September 2012–2014, and October 2021. We observed no effect of fertilization on the relationship between D and H or between D and crown base, so we assumed no effect on the allometric functions for foliage or branch biomass. A dendrometer band (Series 5 Manual Band, Forestry Suppliers Inc., Jackson, MS, USA) was installed on 18 representative trees (one per plot) to monitor radial growth monthly.Three 0.25 m2 circular litter traps were installed 60 cm above the forest floor in each plot in April 2011. Litter was collected at 3-month intervals between June 2011 and March 2015. The litter from each trap was transported to the laboratory and then oven-dried at 65 °C for 48 h. All dried samples were separated into needles, bark, cones, branches, and miscellaneous components, and weighed separately.In September 2014, we estimated the biomass of understory vegetation, separately for woody plants and herbaceous plants. All woody plants  More

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