Regeneration Traits of sinensis Pers. and Aphananthe aspera (Thunb.) Planch. in a Created Urban Tree Plantation approximately 20 years after construction

Keizo TABATA* and Yukihiro MORIMOTO**

Abstract： The groundbreaking example of creation of a tree plantation in an urban area is the “Inochi-No-Mori” project. The main goal of vegetation creation in Inochi-No-Mori is a broad-leaved forest dominated by Celtis sinensis and Aphananthe aspera. C. sinensis and A. aspera forests are thought to be the native vegetation of the Kyoto basin. For creation of C. sinensis and A. aspera plantations in urban areas, the promotion of natural regeneration of these two woody species is a necessity. To achieve this goal, an understanding of the growth characteristics of seedlings of these two species is required. To clarify the growth characteristics, recruitment, mortality and relative height growth rates of seedlings of C. sinensis and A. aspera in Inochi- No-Mori, we set up 163 quadrats (163 m2) on the forest floor of this tree plantation. As a result, there were no significance differences in mortality rates between two species. Recruitment rates of A. aspera seedlings, were relatively low. But low mortality and high growth rates were observed in this species. Although there were high recruitment rates in the C. sinensis seedlings, mortality rates were high and growth rates were relatively low. In a created urban tree plantation, the regeneration processes of the closely related species C. sinensis and A. aspera differed. Key Words： created urban tree plantation, growth characteristic, woody seedling

succession. The main goal of vegetation creation in INTRODUCTION Inochi-No-Mori is a deciduous broad-leaved forest dominated by Celtis sinensis Pers. and Modern urban areas are not appropriate spaces Aphananthe aspera (Thunb.) Planch. (Morimoto for most wildlife to inhabit because the top priority and Natuhara, 2005). Through the analysis of for these areas is convenience for humans. Urban fossil pollen, it is estimated that a deciduous ecosystems possess poor biological diversity. In this broad-leaved forest dominated by C. sinensis and situation, creation of near natural tree plantations A. aspera covered the Kyoto basin approximately and improvement of ecosystems in urban areas are 8,500–5,000 years ago (Takahara, 1998). Therefore, needed (Natuhara, 1998). The groundbreaking C. sinensis and A. aspera forests are thought to be example of creation of a tree plantation in an the native vegetation of the Kyoto basin (Shidei, urban area is the “Inochi-No-Mori” project 1993; Morimoto and Natuhara, 2005). Further, (Morimoto and Natuhara, 2005). At the site of a because C. sinensis and A. aspera are deciduous former freight station in Kyoto city where the trees and grow to a large size, they form a diverse vegetation had once been lost, soil has been city landscape in every season (Sakamoto, 1988). replenished and several mature trees have been These forests are suitable for recreational and planted sparsely during construction. Since then, relaxation places of city residents because C. human interference has been minimized as much sinensis and A. aspera forests have a bright as possible. Creation of a near natural tree atmosphere (Sakamoto, 1988). C. sinensis and A. plantation is the intended result of vegetation aspera bear sap (Okamoto and Kitajima, * Faculty of Agriculture, Kindai University * * Faculty of Bioenvironmental science, Kyotogakuen University

Journal of Environmental Information Science 45-5 1 freight train station of Japan National Railways on this site. The main target for vegetation creation at Inochi-No-Mori is Tadasu-No-Mori Forest in Shimogamo-Jinja shrine, one of the oldest shrine in Kyoto. The vegetation of Kyoto City Tadasu-No-Mori Forest is a deciduous ・Umekoji-Park broad-leaved forest dominated by A. aspera and C. sinensis. A secondary target for vegetation ・Kyoto City creation of Inochi-No-Mori is the deciduous broad-leaved forest dominated Quercus serrata 0 10 20m Murray that was once located in the hills around ● Celtis sinensis 〇 Aphananthe aspera Kyoto basin. When Inochi-No-Mori was created in 0 5 10km ● Others 1996, several forest stands with different densities Figure 2 Spatial distribution of planted trees of trees and with different tree species were in Inochi-No-Mori established (Morimoto and Natuhara, 2005). plantation and seedlings that were ≥15 cm in Mainly soil from the tunnel construction of the height. No study has been done in a created tree subway in Kyoto city was deposited in this site. plantation at a more advanced stage of vegetative Soil pH of the topsoil (10–15 cm) was measured at Umekoji-Park succession focusing on seedlings at earlier stages 175 points in Inochi-No-Mori on August, 2012 ● Inocｈi-No-Mori of growth. (Tabata, 2012 unpublished data). The mean value We investigated mortality, growth, and (± standard deviation) of pH of the soil was 6.1 recruitment of seedlings of C. sinensis and A. (±0.6). aspera at the early stages of growth in A survey on all planted trees with a diameter at Inochi-No-Mori about 20 years after initiation. In breast height of 5 cm or larger in Inochi-No-Mori addition, the influences of the initial size of was conducted in 2015 (Figure 2). The total basal 0 500 1000m seedlings, light environment on the forest floor, area of all planted trees was 24.0 m2/ha. Q. serrata Figure 1 Location of study site and the potential supply of seeds were examined. Murray had the largest basal area (6.1 m2/ha); Q. All maps were created based on the Fundamental Geospatial Data provided by the Geospatial Thus, the purpose of this study is to clarify glauca Thunb. had the second largest basal area 1)2) Information Authority of Japan . regeneration processes of C. sinensis and A. aspera (3.6 m2/ha); and Zelkova serrata (Thunb.) Makino 1988) which provide food to wild birds such as gray a necessity. To achieve this goal, an understanding in the created tree plantation in an urban area. had the third largest basal area (2.7 m2/ha). Basal starlings (Okamoto and Kitajima, 1988). C. of the growth characteristics of seedlings of these area of C. sinensis was 1.9 m2/ha. Basal area of A. sinensis and A. aspera improve biodiversity in two species is required. There are previous studies 1. METHODS aspera was 1.3 m2/ha. There were a total of 381 urban areas. Thus, creation of C. sinensis and A. of the growth characteristics of seedlings of these individual planted trees (630.0 individuals/ha). Q. aspera plantations in urban areas is very two species (Higa et al., 2006; Higa et al., 2012; 1.1 Study site serrata had the largest number with 101 beneficial. Panetta, 2001). However, these studies were The study site, Inochi-No-Mori, is located in individuals. There were 22 individuals of C. In creation of natural habitats, strategies must conducted in riparian forests (Higa et al., 2006) or Umekoji General Park (Figure 1) (34º59’N, sinensis and 21 individuals of A. aspera. be adopted that take advantage of the natural on experimental farms (Higa et al., 2012; Panetta, 135º44’E; 11.65 ha) in Shimogyo ward in Kyoto city. 1.2 Establishment of quadrat resilience of these biological communities 2001). They are not studies in urban forests that Kyoto City had a mean annual air temperature of We set up 1-m × 1-m quadrats at 5-m intervals (Committee on Ecosystem Management, The possess specific functions and structures 15.9ºC (28.2ºC in the highest; 4.6ºC in the lowest), in the entire area of Inochi-No-Mori in 2012. A Ecological Society of Japan, 2005). Further, in (Sakamoto, 1999). Studies in mature urban forests and an average annual precipitation of 1,491.3 total of 163 quadrats were established. creation of near natural tree plantations in urban and plantations have been described by Sakamoto mm over the past 30 years (1981–2010)3). This 1.3 Measurement of growth condition of areas, minimal human interference and the et al. (1987), Tabata and Morimoto (2012), and area belongs to the warm temperate evergreen seedlings facilitation of natural regeneration are desirable. Tabata et al. (2015). There is only one study broad-leaved forest zone with a warmth index of We identified individuals of all woody seedlings For creation of C. sinensis and A. aspera conducted in a created urban tree plantation 130.7ºC・months and a coolness index of –0.4 ºC・ that emerged in the 163 quadrats. Species names plantations in urban areas, the promotion of (Tabata and Morimoto, 2012); however, this study months3). Inochi-No-Mori is 6048 m2. It was and heights of seedlings were recorded. Surveys natural regeneration of these two woody species is focused on the early development stage of the tree established in April, 1996. Previously, there was a were conducted in 2012, 2013, 2014, and 2015.

2 Journal of Environmental Information Science 45-5 freight train station of Japan National Railways on this site. The main target for vegetation creation at Inochi-No-Mori is Tadasu-No-Mori Forest in Shimogamo-Jinja shrine, one of the oldest shrine in Kyoto. The vegetation of Tadasu-No-Mori Forest is a deciduous broad-leaved forest dominated by A. aspera and C. sinensis. A secondary target for vegetation creation of Inochi-No-Mori is the deciduous broad-leaved forest dominated Quercus serrata 0 10 20m Murray that was once located in the hills around ● Celtis sinensis 〇 Aphananthe aspera Kyoto basin. When Inochi-No-Mori was created in ● Others 1996, several forest stands with different densities Figure 2 Spatial distribution of planted trees of trees and with different tree species were in Inochi-No-Mori established (Morimoto and Natuhara, 2005). plantation and seedlings that were ≥15 cm in Mainly soil from the tunnel construction of the height. No study has been done in a created tree subway in Kyoto city was deposited in this site. plantation at a more advanced stage of vegetative Soil pH of the topsoil (10–15 cm) was measured at succession focusing on seedlings at earlier stages 175 points in Inochi-No-Mori on August, 2012 of growth. (Tabata, 2012 unpublished data). The mean value We investigated mortality, growth, and (± standard deviation) of pH of the soil was 6.1 recruitment of seedlings of C. sinensis and A. (±0.6). aspera at the early stages of growth in A survey on all planted trees with a diameter at Inochi-No-Mori about 20 years after initiation. In breast height of 5 cm or larger in Inochi-No-Mori addition, the influences of the initial size of was conducted in 2015 (Figure 2). The total basal seedlings, light environment on the forest floor, area of all planted trees was 24.0 m2/ha. Q. serrata and the potential supply of seeds were examined. Murray had the largest basal area (6.1 m2/ha); Q. Thus, the purpose of this study is to clarify glauca Thunb. had the second largest basal area regeneration processes of C. sinensis and A. aspera (3.6 m2/ha); and Zelkova serrata (Thunb.) Makino in the created tree plantation in an urban area. had the third largest basal area (2.7 m2/ha). Basal area of C. sinensis was 1.9 m2/ha. Basal area of A. 1. METHODS aspera was 1.3 m2/ha. There were a total of 381 individual planted trees (630.0 individuals/ha). Q. 1.1 Study site serrata had the largest number with 101 The study site, Inochi-No-Mori, is located in individuals. There were 22 individuals of C. Umekoji General Park (Figure 1) (34º59’N, sinensis and 21 individuals of A. aspera. 135º44’E; 11.65 ha) in Shimogyo ward in Kyoto city. 1.2 Establishment of quadrat Kyoto City had a mean annual air temperature of We set up 1-m × 1-m quadrats at 5-m intervals 15.9ºC (28.2ºC in the highest; 4.6ºC in the lowest), in the entire area of Inochi-No-Mori in 2012. A and an average annual precipitation of 1,491.3 total of 163 quadrats were established. mm over the past 30 years (1981–2010)3). This 1.3 Measurement of growth condition of area belongs to the warm temperate evergreen seedlings broad-leaved forest zone with a warmth index of We identified individuals of all woody seedlings 130.7ºC・months and a coolness index of –0.4 ºC・ that emerged in the 163 quadrats. Species names months3). Inochi-No-Mori is 6048 m2. It was and heights of seedlings were recorded. Surveys established in April, 1996. Previously, there was a were conducted in 2012, 2013, 2014, and 2015.

Journal of Environmental Information Science 45-5 3 Measurements were carried out from autumn calculated. Table 2 Individual density of seedlings equal rate of C. sinensis was 0.12 cm/cm/year. The through winter in each year. The values of the diffuse site factor were to or less than 50 cm in height average value differed significantly between the 1.4 Evaluation of the light conditions in the categorized into three classes by the tertile method Number of seedlings/100m2 two species (p < 0.01) (Table 4). Specie Year 2012 2013 2014 2015 forest floor for all the values in a total of 489 quadrats over 3 Quercus glauca 286.5 231.3 307.4 296.9 For tree height class, height relative growth rate The measurement of the light quantity was years in 2013, 2014, and 2015 ( I < the first tertile, Quercus serrata 116.6 63.8 135.6 233.1 of A. aspera was 0.21 cm/cm/year in the smaller Celtis sinensis 149.7 141.1 115.3 228.8 carried out in each 163 quadrat. Hemispherical the first tertile ≤ II < the second tertile, III ≥ the Ligustrum lucidum 54.0 36.8 77.3 147.9 than 15-cm height class and was 0.13 cm/cm/year Aphananthe aspera 54.0 62.6 84.7 123.9 photographs were taken at the about 50 cm height second tertile). Cinnamomum camphora 4.9 3.7 11.7 41.1 in the 15–50-cm height class. Mean values of A. from the ground in the center of each quadrat by a As with each height class, the mean values of Acer palmatum 17.2 16.6 22.7 32.5 aspera were significantly higher than those of C. Others 82.8 81.0 101.8 139.3 digital camera (Nikon Coolpix 950) fitted with a the height relative growth rates and mortality Total 765.6 636.8 856.4 1243.6 sinensis (p < 0.01 in the smaller than 15-cm height fish-eye lens (Nikon FC-E8). Photographs were rates were calculated for each species and for each (Table 1). There was no significant difference class, p < 0.05 in the 15–50-cm height class) (Table taken on a cloudy day or in the evening when solar diffuse site factor class. among values in 2013, 2014, and 2015. The first 4). When heights of seedlings and the light elevation was low in each of the 163 quadrats once For recruitment rates, the influence of the tertile of all measurements over 3 years was 12.2 environment were considered in conjunction, the a year from June to early November in 2013, 2014, potential supply of seeds was also examined. As an and the second tertile was 14.0. mean height relative growth rate for A. aspera was and 2015. Diffuse site factor values weighted the indicator of the seed supply potential, the number 2.2 Population densities of seedlings 0.27 cm/cm/year in the smaller than 15-cm height brightness of the zenith direction (Anderson, 1971) of planted trees of the same species as the Individual density of seedlings equal to or less class and in the smaller than 12.2% diffuse site were calculated from hemispherical photographs seedlings located around the quadrats was than 50 cm in height was the highest for Q. glauca factor class. The mean value of A. aspera was 0.17 by analysis software4). For diffuse site factors, examined. Numbers of planted trees of the same seedlings since 2012 to 2015 (Table 2). Individual cm/cm/year in the 15–50-cm height class and in significant differences were examined among the species as the seedlings within a circular range of density of Q. serrata seedlings was the second the 12.2%–14.0% diffuse site factor class. Mean three years, 2013, 2014, and 2015, using Tukey’s 10-m or 20-m radius centered in each quadrat highest in 2014 and in 2015 (Table 2). Individual values of A. aspera were significant higher than multiple comparison test. were counted for each species. The average density of C. sinensis seedlings was the third those of C. sinensis in these categories (p < 0.05) 1.5 Data analysis recruitment rates over the three periods from 2012 highest in 2014 and in 2015. Individual density of (Table 4). From the measurements of each seedling in 2012 to 2013, from 2013 to 2014, and from 2014 to 2015 A. aspera seedlings was the fifth highest in 2015 2.5 Recruitment rates of seedlings and 2013, 2013 and 2014, and 2014 and 2015, were calculated. Number of planted trees of the (Table 2). The total recruitment rate of A. aspera was 39.7 using equation 1, the height relative growth rate same species as the seedlings was categorized into 2.3 Mortality rates of seedlings seedlings /100 m2/year and the recruitment rate of during each period from 2012 to 2013, from 2013 one of three classes (I = 0, 1 ≤ II ≤ 3, III ≥ 4). As The total mortality rate of A. aspera was C. sinensis was 68.7 seedlings/100 m2/year (Table to 2014, and from 2014 to 2015 was calculated. described above for the planted trees classes, the 25.0%/year and the mortality rate of C. sinensis 5). There was a significant difference between two Height relative growth rate (cm/cm/year) mean values of the recruitment rates were was 30.2%/year (Table 3). Overall mortality rates species (p < 0.01).

= [Ln(Height end of period) − Ln(Height initial)] (1) calculated for each species and for each diffuse site of C. sinensis tended to be higher than the Considering the relationship between the supply

(Height end of period and Height initial represent the factor class. mortality rates of A. aspera. However, there was potential of the seeds, the mean value of the height at the end and beginning of the period, For the mortality rates, significant differences in no significant difference between the two species recruitment rate of C. sinensis was 85.8 respectively.) proportions were examined between the two in any size category (Table 3). seedlings/100 m2/year when planted trees of the The initial size of the seedlings and the light species in the same category using Fisher’s exact 2.4 Growth rates of seedlings same species as the seedlings were not within a environment, diffuse site factors were examined as test. Statistical analyses were performed using R Total height relative growth rate of A. aspera radius of 10 m. The mean recruitment rate of C. factors affecting the growth and mortality of version 3.2.55). was 0.17 cm/cm/year and height relative growth sinensis was significantly higher than that of A. seedlings. We used the height in 2012 for the Table 3 Mortality rate in each height class and in each diffuse site factor class period from 2012 to 2013, the height in 2013 for 2. RESULTS Diffuse Site Factor (%) < 12.2 12.2 ≤ < 14.0 14.0 ≤ Total the period from 2013 to 2014, and the height in Height Mortality Mortality Mortality Mortality 2014 for the period from 2014 to 2015 as the initial 2.1 Light environment (cm) Species N Rate (%/yr) N Rate (%/yr) N Rate (%/yr) N Rate (%/yr) sizes. The mean values of the diffuse site factor were Celtis sinensis 89 41.6 132 25.8 266 34.6 487 33.5 Each seedling was categorized into one of two 12.7% in 2013, 13.2% in 2014, and 13.0% in 2015 < 15 Aphananthe aspera 30 36.7 63 36.5 75 29.3 168 33.3 p nns nns nns nns sizes classes (I < 15 cm, 15 cm ≤ II ≤ 50 cm) Table 1 Light environment at 50 cm above Celtis sinensis 34 26.5 34 08.8 107 23.4 175 21.1 according to the height at the beginning of period. the ground in Inochi-No-Mori 15 ≤ ≤ 50 Aphananthe aspera 35 20.0 61 13.1 64 17.2 160 16.3 For each species and height class, the mean value Diffuse Site Factor (%) p nns nns nns nns Year N mean S.D. Min Max Celtis sinensis 123 37.4 166 22.3 373 31.4 662 30.2 of the height relative growth rate and mortality 2013 163 12.7 2.3 5.2 17.8 Total Aphananthe aspera 65 27.7 124 25.0 139 23.7 328 25.0 rate over the three periods from 2012 to 2013, 2014 163 13.2 2.3 6.4 21.6 p nns nns nns nns from 2013 to 2014, and from 2014 to 2015 was 2015 163 13.0 1.9 6.4 17.4 （ns：p ≥ 0.05)

4 Journal of Environmental Information Science 45-5 Table 2 Individual density of seedlings equal rate of C. sinensis was 0.12 cm/cm/year. The to or less than 50 cm in height average value differed significantly between the Number of seedlings/100m2 two species (p < 0.01) (Table 4). Specie Year 2012 2013 2014 2015 Quercus glauca 286.5 231.3 307.4 296.9 For tree height class, height relative growth rate Quercus serrata 116.6 63.8 135.6 233.1 of A. aspera was 0.21 cm/cm/year in the smaller Celtis sinensis 149.7 141.1 115.3 228.8 Ligustrum lucidum 54.0 36.8 77.3 147.9 than 15-cm height class and was 0.13 cm/cm/year Aphananthe aspera 54.0 62.6 84.7 123.9 Cinnamomum camphora 4.9 3.7 11.7 41.1 in the 15–50-cm height class. Mean values of A. Acer palmatum 17.2 16.6 22.7 32.5 aspera were significantly higher than those of C. Others 82.8 81.0 101.8 139.3 Total 765.6 636.8 856.4 1243.6 sinensis (p < 0.01 in the smaller than 15-cm height (Table 1). There was no significant difference class, p < 0.05 in the 15–50-cm height class) (Table among values in 2013, 2014, and 2015. The first 4). When heights of seedlings and the light tertile of all measurements over 3 years was 12.2 environment were considered in conjunction, the and the second tertile was 14.0. mean height relative growth rate for A. aspera was 2.2 Population densities of seedlings 0.27 cm/cm/year in the smaller than 15-cm height Individual density of seedlings equal to or less class and in the smaller than 12.2% diffuse site than 50 cm in height was the highest for Q. glauca factor class. The mean value of A. aspera was 0.17 seedlings since 2012 to 2015 (Table 2). Individual cm/cm/year in the 15–50-cm height class and in density of Q. serrata seedlings was the second the 12.2%–14.0% diffuse site factor class. Mean highest in 2014 and in 2015 (Table 2). Individual values of A. aspera were significant higher than density of C. sinensis seedlings was the third those of C. sinensis in these categories (p < 0.05) highest in 2014 and in 2015. Individual density of (Table 4). A. aspera seedlings was the fifth highest in 2015 2.5 Recruitment rates of seedlings (Table 2). The total recruitment rate of A. aspera was 39.7 2.3 Mortality rates of seedlings seedlings /100 m2/year and the recruitment rate of The total mortality rate of A. aspera was C. sinensis was 68.7 seedlings/100 m2/year (Table 25.0%/year and the mortality rate of C. sinensis 5). There was a significant difference between two was 30.2%/year (Table 3). Overall mortality rates species (p < 0.01). of C. sinensis tended to be higher than the Considering the relationship between the supply mortality rates of A. aspera. However, there was potential of the seeds, the mean value of the no significant difference between the two species recruitment rate of C. sinensis was 85.8 in any size category (Table 3). seedlings/100 m2/year when planted trees of the 2.4 Growth rates of seedlings same species as the seedlings were not within a Total height relative growth rate of A. aspera radius of 10 m. The mean recruitment rate of C. was 0.17 cm/cm/year and height relative growth sinensis was significantly higher than that of A. Table 3 Mortality rate in each height class and in each diffuse site factor class Diffuse Site Factor (%) < 12.2 12.2 ≤ < 14.0 14.0 ≤ Total Height Mortality Mortality Mortality Mortality (cm) Species N Rate (%/yr) N Rate (%/yr) N Rate (%/yr) N Rate (%/yr) Celtis sinensis 89 41.6 132 25.8 266 34.6 487 33.5 < 15 Aphananthe aspera 30 36.7 63 36.5 75 29.3 168 33.3 p nns nns nns nns Celtis sinensis 34 26.5 34 08.8 107 23.4 175 21.1 15 ≤ ≤ 50 Aphananthe aspera 35 20.0 61 13.1 64 17.2 160 16.3 p nns nns nns nns Celtis sinensis 123 37.4 166 22.3 373 31.4 662 30.2 Total Aphananthe aspera 65 27.7 124 25.0 139 23.7 328 25.0 p nns nns nns nns （ns：p ≥ 0.05)

Journal of Environmental Information Science 45-5 5 Table 4 Height relative growth rate in each height class and in each diffuse site factor class larger than 14.0% diffuse site factor class in Diffuse Site Factor (%) 3. DISCUSSION Inochi-No-Mori (Table 3). In Tadasu-No-Mori < 12.2 12.2 ≤ < 14.0 14.0 ≤ Total Height Height Height Height Forest, the mean height relative growth rate for C. Relative Relative Relative Relative Overall, A. aspera seedlings displayed higher sinensis seedlings was 0.37 cm/cm/year and that of Growth Rate Growth Rate Growth Rate Growth Rate growth rates than C. sinensis (Table 4). Seedlings A. aspera seedlings was 0.61 cm/cm/year in the (cm/cm/yr) (cm/cm/yr) (cm/cm/yr) (cm/cm/yr) Height (cm) Species N mean S.D. N mean S.D. N mean S.D. N mean S.D. that germinate from large-sized seeds have strong larger than 13.7% diffuse site factor class (Tabata Celtis sinensis 51 0.09 0.11 92 0.14 0.15 156 0.15 0.22 299 0.14 0.18 resistance to various stresses, such as shade et al, 2015). In contrast, in Inochi-No-Mori, the < 15 Aphananthe aspera 18 0.27 0.29 38 0.17 0.22 51 0.23 0.29 107 0.21 0.26 p * ns ns ** condition (Seiwa and Kikuzawa, 1989). The dry mean height relative growth rate for C. sinensis Celtis sinensis 23 0.04 0.06 29 0.07 0.09 76 0.09 0.14 128 0.08 0.12 weight of C. sinensis seed is 40 g/1000 seeds―80 seedlings was 0.13 cm/cm/year and that for A. 15 ≤ ≤ 50 Aphananthe aspera 26 0.10 0.13 53 0.17 0.25 48 0.12 0.21 127 0.13 0.22 p ns * ns * g/1000 seeds and that of A. aspera is 110 g/1000 aspera seedlings was 0.17 cm/cm/year in the larger Celtis sinensis 74 0.08 0.10 121 0.12 0.14 232 0.13 0.20 427 0.12 0.17 seeds―200 g/1000 seeds (Katsuta et. al, 1998). It is than 14.0% diffuse site factor class (Table 4). Total Aphananthe aspera 44 0.17 0.23 91 0.17 0.24 99 0.17 0.25 234 0.17 0.24 p * ns ns ** possible that differences in the seed weight of the Under similar light conditions, the growth rates of （**：p < 0.01, *：p < 0.05, ns：p ≥ 0.05) two species are related to the differences in growth seedlings of both species in Tadasu-No-Mori Forest 2 aspera in this case (p < 0.001) (Table 5). When four than that of A. aspera (28.9 seedlings/100 m /year) rate. were higher than those in Inochi-No-Mori. As or more planted trees of the same species as the (p < 0.001) (Table 5). C. sinensis had higher recruitment rates than A. mentioned above, the soil in Inochi-No-Mori is seedlings were within a radius of 10 m, the mean Considering the supply potential of the seeds aspera (Table 5). In this research site, mainly generated from subway construction. recruitment rate of C. sinensis (92.8 seedlings/100 and the light environment in conjunction, when Inochi-No-Mori, the number of indivisuals of Differences in the growth rates of seedlings 2 m /year) was significantly higher than that of A. 1–3 planted trees of the same species as the planted C. sinensis are almost the same as that between these two sites may be influenced by soil 2 aspera (38.1 seedlings/100 m /year) (p < 0.05) seedlings were within a radius of 20 m and in each of A. aspera (Figure 2). To promote the properties. It is necessary to examine these factors (Table 5). diffuse site factor class (<12.2%, 12.2%–14.0%, regeneration of A. aspera in created tree in further studies. When 1–3 planted trees of the same species as ≥14.0%), recruitment rates of C. sinensis were plantations in urban areas, it is necessary to In this study, we examined the initial growth the seedlings were within a radius of 20 m, the higher than those of A. aspera. There were more mother trees of A. aspera. Both C. sinensis characteristics of C. sinensis and A. aspera mean recruitment rate of C. sinensis (84.4 significant differences between the two species in and A. aspera bear well every year. However, seedlings. For the effective establishment of C. 2 seedlings/100 m /year) was significantly higher each category (p < 0.05) (Table 5). the amount of fruit per individual of these species sinensis and A. aspera plantation in urban areas, Table 5 Recruitment rate in each class of number of planted trees of the same species as seedlings within is unknown (Silvics of Japan Editorial Board, growth patterns of saplings of these species during the area in each diffuse site factor class 2009). Fruit amount per individual of these species later growth stages must be investigated in future Number Diffuse Site Factor (%) might influence differences in recruitment. As studies. of Total described above, because of the light seed weight of planted < 12.2 12.2 ≤ < 14.0 14.0 ≤ The trees of Recruitment Recruitment Recruitment Recruitment C. sinensis, their seeds may have been more CONCLUSIONS radius the same Rate Rate Rate Rate widely dispersed by birds. For effective planning of of the species (N/100m2/yr) (N/100m2/yr) (N/100m2/yr) (N/100m2/yr) area planting to create plantations in urban areas, we There were significance differences in the within Quadrat Quadrat Quadrat Quadrat Species mean S.D. mean S.D. mean S.D. mean S.D. the area Number Number Number Number focused on the relationship between the number of recruitment and growth rates of seedlings between Celtis sinensis 78 42.3 105.1 67 97.0 311.4 74 121.6 200.9 219 85.8 218.9 planted trees around quadrats and the number of C. sinensis and A. aspera. Although recruitment 0 Aphananthe aspera 76 21.1 63.9 81 25.9 62.8 71 54.9 84.2 228 33.3 71.7 p ns ns * *** emerged seedlings. In addition, in consideration of rates for C. sinensis seedlings were high, growth Celtis sinensis 60 40.0 115.3 79 41.8 124.7 62 43.5 98.5 201 41.8 113.8 10 m 1 ≤ ≤ 3 Aphananthe aspera 66 28.8 142.3 77 50.6 111.9 76 57.9 130.9 219 46.6 128.2 the view of visitors in this site, seed traps were not rates were relatively low. In contrast, recruitment p ns ns ns ns set in this study. However, for further study, rates of A. aspera seedlings were relatively low, Celtis sinensis 26 61.5 174.5 23 126.1 191.2 20 95.0 166.9 69 92.8 177.7 4 ≤ Aphananthe aspera 22 9.1 29.4 11 72.7 155.5 9 66.7 132.3 42 38.1 103.5 investigation of the seed supply directly by setting whereas growth rates were higher for this species. p ns ns ns * seed traps is needed. In a created urban tree plantation, the Celtis sinensis 24 41.7 97.4 23 143.5 482.3 16 43.8 72.7 63 79.4 299.5 0 Aphananthe aspera 14 0.0 0.0 25 24.0 59.7 33 54.5 75.4 72 33.3 65.0 In a previous study conducted in regeneration processes of the closely related p * ns ns ns Celtis sinensis 46 50.0 120.6 43 74.4 169.2 46 128.3 164.2 135 84.4 155.0 Tadasu-No-Mori Forest, an urban forest in Kyoto species C. sinensis and A. aspera differed. 20 m 1 ≤ ≤ 3 Aphananthe aspera 56 10.7 31.2 60 18.3 56.7 43 67.4 119.0 159 28.9 76.6 city, the mortality rate for C. sinensis seedlings p * * * *** Celtis sinensis 94 42.6 128.3 103 60.2 145.1 94 74.5 175.3 291 59.1 150.7 was 35.0%/year and that for A. aspera seedlings NOTES 4 ≤ Aphananthe aspera 94 33.0 129.8 84 60.7 120.3 80 52.5 120.1 258 48.1 123.9 was 20.0%/year in the larger than 13.7% diffuse 1）Geospatial Information Authority of Japan p ns ns ns ns Celtis sinensis 164 44.5 121.5 169 75.1 226.2 156 87.2 166.1 489 68.7 177.8 site factor class (Tabata et al, 2015). The mortality , 01/24/2017 referred. Total Aphananthe aspera 164 22.6 100.5 169 40.2 95.9 156 57.1 111.4 489 39.7 103.3 rate for C. sinensis seedling was 31.4%/year and 2）Tani, K. (01/17/2017 updated) Geographic information p ns ns ns ** （***：p < 0.001, **：p < 0.01, *：p < 0.05, ns：p ≥ 0.05) that for A. aspera seedlings was 23.7%/year in the analysis support system MANDARA

6 Journal of Environmental Information Science 45-5 larger than 14.0% diffuse site factor class in 3. DISCUSSION Inochi-No-Mori (Table 3). In Tadasu-No-Mori Forest, the mean height relative growth rate for C. Overall, A. aspera seedlings displayed higher sinensis seedlings was 0.37 cm/cm/year and that of growth rates than C. sinensis (Table 4). Seedlings A. aspera seedlings was 0.61 cm/cm/year in the that germinate from large-sized seeds have strong larger than 13.7% diffuse site factor class (Tabata resistance to various stresses, such as shade et al, 2015). In contrast, in Inochi-No-Mori, the condition (Seiwa and Kikuzawa, 1989). The dry mean height relative growth rate for C. sinensis weight of C. sinensis seed is 40 g/1000 seeds―80 seedlings was 0.13 cm/cm/year and that for A. g/1000 seeds and that of A. aspera is 110 g/1000 aspera seedlings was 0.17 cm/cm/year in the larger seeds―200 g/1000 seeds (Katsuta et. al, 1998). It is than 14.0% diffuse site factor class (Table 4). possible that differences in the seed weight of the Under similar light conditions, the growth rates of two species are related to the differences in growth seedlings of both species in Tadasu-No-Mori Forest rate. were higher than those in Inochi-No-Mori. As C. sinensis had higher recruitment rates than A. mentioned above, the soil in Inochi-No-Mori is aspera (Table 5). In this research site, mainly generated from subway construction. Inochi-No-Mori, the number of indivisuals of Differences in the growth rates of seedlings planted C. sinensis are almost the same as that between these two sites may be influenced by soil of A. aspera (Figure 2). To promote the properties. It is necessary to examine these factors regeneration of A. aspera in created tree in further studies. plantations in urban areas, it is necessary to plant In this study, we examined the initial growth more mother trees of A. aspera. Both C. sinensis characteristics of C. sinensis and A. aspera and A. aspera bear fruit well every year. However, seedlings. For the effective establishment of C. the amount of fruit per individual of these species sinensis and A. aspera plantation in urban areas, is unknown (Silvics of Japan Editorial Board, growth patterns of saplings of these species during 2009). Fruit amount per individual of these species later growth stages must be investigated in future might influence differences in recruitment. As studies. described above, because of the light seed weight of C. sinensis, their seeds may have been more CONCLUSIONS widely dispersed by birds. For effective planning of planting to create plantations in urban areas, we There were significance differences in the focused on the relationship between the number of recruitment and growth rates of seedlings between planted trees around quadrats and the number of C. sinensis and A. aspera. Although recruitment emerged seedlings. In addition, in consideration of rates for C. sinensis seedlings were high, growth the view of visitors in this site, seed traps were not rates were relatively low. In contrast, recruitment set in this study. However, for further study, rates of A. aspera seedlings were relatively low, investigation of the seed supply directly by setting whereas growth rates were higher for this species. seed traps is needed. In a created urban tree plantation, the In a previous study conducted in regeneration processes of the closely related Tadasu-No-Mori Forest, an urban forest in Kyoto species C. sinensis and A. aspera differed. city, the mortality rate for C. sinensis seedlings was 35.0%/year and that for A. aspera seedlings NOTES was 20.0%/year in the larger than 13.7% diffuse 1）Geospatial Information Authority of Japan site factor class (Tabata et al, 2015). The mortality , 01/24/2017 referred. rate for C. sinensis seedling was 31.4%/year and 2）Tani, K. (01/17/2017 updated) Geographic information that for A. aspera seedlings was 23.7%/year in the analysis support system MANDARA

Journal of Environmental Information Science 45-5 7 , 01/24/2017 referred. Panetta, F.D. (2001) Seedling emergence and seed longevity of Insight into Pro-Environmental Behaviors and People’s Perceptions in Bangkok, 3）Japan Meteorological Agency. Climatic statistics. the tree weeds Celtis sinensis and Cinnamomum camphora. Thailand , Weed Research, No.41，83～95. Sarunnoud PHUPHISITH*, Kiyo KURISU* and Keisuke HANAKI* 03/10/2015 referred. Sakamoto, K., Kobayashi, T. and Ikeuchi, Z. (1987) A case study 4）Takenaka, A. (03/06/2009 updated) CanopOn2 a software for on the structure of the stand in the forest surrounding Abstract: Fostering pro-environmental behaviors (PEBs) is a critical step towards achieving an analysis of hemispherical. Shimogamo Shrine, Kyoto. Journal of The Japanese sustainable societies, and has become a great challenge for both developed and developing , Institute of Landscape Architecture, No.48 ， 275 ～ 280. nations. However, understanding of PEBs from the perspective of developing countries is scant. 08/27/2015 referred. (Written in Japanese with English summary) Considering this limitation, the present study aimed to provide some insights into PEBs and 5）R Development Core Team (04/14/2016 updated) Sakamoto, K. (1988) Study on remnant forms and changes of reveal how they are perceived and performed by citizens in a developing society. A survey was conducted in Bangkok and its vicinity in Thailand. Our findings indicate that the majority of R: a language and environment for statistical computing Ulmaceae woods and trees in an urban area. Bulletin of respondents engaged to some extent in PEBs. Higher degree of perception, older generation and ： Revegetation Research, No.2，1～129. (Written in Japanese female were key predictors for most of the environmental attitudes. In addition, an in-depth R Foundation for Statistical Computing, Vienna. , with English summary) comparison to the literature highlighted similarities and dissimilarities in PEB domains 05/10/2016 referred. Sakamoto, K. (1999) Management and conservation of urban performed by citizens of different societies. The importance impacts of specific features of PEB domains and external factors were pointed out and discussed. forests. In: Okada, M., Ohasawa, M. and Suzuki, M., ed., Key words: pro-environmental behaviors, practice rate, perception, developing country REFERENCES “ Eco-technology for environmental conservation and Anderson, M.C. (1971) Radiation and crop structure. In: Sestak, creation”, p.32～p.41, Maruzen, Tokyo. (Written in Japanese) Z., Catsky, T. and Jarvis, P.G, ed., “Plant Photosynthetic Seiwa, K. and Kikuzawa, K. 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Nakanishiya Shuppan，Kyoto, 296 pp. (Written in Japanese) become one of the ultimate goals in dealing with These external factors include existing regulations (Written in Japanese with English summary) Silvics of Japan Editorial Board (2009) Silvics of Japan 1. environmental difficulties. Collaboration of all or policy provisions, availability of infrastructure Higa, M., Ishikawa, S. and Miyake, N. (2006) Ecological Nihon-Ringyo-Chosakai, Tokyo, 762pp. (Written in Japanese) sectors, from the individual to international levels, and culture and tradition. These society-oriented characteristics of Ulmus parvifolia, Celtis sinensis var. Tabata, K. and Morimoto, Y. (2012) Growth characteristics of is vital to accomplish this aim. Among these variables, which vary from one place to another, japonica and Aphananthe aspera on high and dry habitats of woody seedlings emerged in a restored urban forest for five stakeholders, the residential sector contributes to may facilitate or prevent people from committing to gravelly bars in rivers. Vegetation Science, No.23, 89～103. years since construction. Journal of The Japanese Institute environmental issues by consumption patterns and PEBs. (Written in Japanese with English summary) of Landscape Architecture No.75, 431 ～ 434. (Written in lifestyles. Encouraging people to change behaviors Furthermore, beyond the society level, cross- Higa, M., Moriyama, T. and Ishikawa, S. (2012) Effects of Japanese with English summary) by adopting environmental-friendly actions, so- national impacts on environmental attitudes and complete submergence on seedling growth and survival of Tabata, K., Hashimoto, H. and Morimoto, Y. (2015) Growth called ‘pro-environmental behaviors’ (PEBs), is one behaviors have been explored by many researchers five riparian tree species in the warm-temperate regions of characteristics of seedlings of Celtis sinensis, Aphananthe essential measure towards evolving into a (Dunlap & Mertig, 1995; Francis, 1998; Hunter et Japan. Journal of Forest Research, No.17, 129～136. aspera, Zelkova serrata and Cinnamomum camphora, sustainable society. al., 2004; Dalton, 2005; Gelissen, 2007; Franzen & Katsuta, M., Mori, T. and Yokoyama, T. (1998) Tree seeds in dominant species in the canopy layer in a large-scale mature Previous studies have revealed that engaging in Meyer, 2010; Pirani & Secondi, 2010; Pisano & del Japan (broad-leaved trees). Japan Forest Tree Breeding urban forest. Journal of The Japanese Institute of Landscape PEBs is a result of various factors. Many studies Carmen Hidalgo, 2014 and Pisano & Lubell, 2015). Association, Tokyo, 410pp. (Written in Japanese) Architecture, No.78, 663～666. (Written in Japanese with have shown the important impact of individual- This study area comprises issues on the level of Morimoto Y, and Natuhara Y (2005) Living forest: bio-city in English summary) oriented, namely psychological, factors (e.g. norms, country development, such as economics, education, theory and practice. Kyoto University Press, Kyoto, 398 pp. Takahara, H. (1998) Vegetation history in the Kinki region. In: values and attitudes) and socio-demographic factors and the situation of environmental degradation. It (Written in Japanese) Yasuda, Y. and Miyoshi, N., ed., “Atras of Vegetation history (e.g. gender, age and education) (Stern et al., 1999; has been claimed that people in wealthier countries Natuhara, Y. (1998) Design of woodlands in urban areas. in the Japanese Islands” p.114 ～p.137, Asakura-Shoten, Zelezny et al., 2000; Gatersleben et al., 2002; display more environmental concern (Franzen & Journal of Urban Living and Health Association, No.42，113 Tokyo. (Written in Japanese) Kollmuss & Agyeman, 2002; Diamantopoulos et al., Meyer, 2010) and engage more in environmental- ～133. (Written in Japanese with English summary) 2003; Tanner & Kast, 2003; Gilg et al., 2005; Gilg & friendly actions (Dalton, 2005 and Pisano & Lubell, Okamoto, M. and Kitajima, A. (1988) Growth and ripening Barr, 2006; Steg & Vlek, 2009; Whitmarsh & 2015) than people in poorer nations. Theoretical patterns in the fruits of several indigenous bird-dispersed O’Neill, 2010 and Tabi, 2013). These factors can explanations for differences in environmentalism cultivated in Nagai Botanic Garden. Bulletin of the personally motivate people to act on environmental issues and country-level variables have been Osaka Museum of Natural History, No.42, 1～13. (Written in concerns. In addition, a number of studies have discussed in other studies (Spaargaren & Mol, 1992;

Japanese with English summary) * Department of Urban Engineering, Graduate school of Engineering, The University of Tokyo, Tokyo, Japan

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