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Table 1. Sizes and light environments of the sample trees (n = 5~6 for each species) and total

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number of individuals in the research plot and for each species. Species are listed in order of

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increasing distance of spatial distribution from forest edge (Fig. 3).

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Species

DBH (cm)

Height (m)

Canopy openness (%)

Total individuals

L. lucidum

4.13~7.83

4.43~9.04

5.4~26.4

58

Ci. japonicum

2.76~5.90

3.09~10.05

6.48~21.41

84

Q. glauca

2.45~6.78

3.30~10.66

5.51~17.12

166

T. gymnanthera

3.03~8.42

3.96~10.12

4.95~14.48

112

Ca. japonica

3.79~9.01

4.05~10.52

6.30~18.51

206

I. rotunda

2.31~6.23

3.43~10.15

6.88~24.15

55

All measurements were taken in 2019.

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357

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Table 2. Leaf functional traits (mean ± one standard error) of the study species.

Species

Thickness (mm)

LMA (g m-2)

Amax (µmol m-2 s-1)

Nmass (%)

L. lucidum

0.34 ± 0.01 bc

103.4 ± 3.9 b

8.58 ± 0.57 a

2.89 ± 0.09 a

Ci. japonicum

0.24 ±0.01 d

102.8 ± 4.5 b

4.76 ± 0.67 b

2.00 ± 0.11 b

Q. glauca

0.33 ± 0.01 c

101.9 ± 4.0 b

6.00 ± 0.61 b

1.97 ± 0.09 b

T. gymnanthera

0.45 ± 0.01 a

112.8 ± 4.7 ab

6.61 ± 0.72 b

1.12 ± 0.11 c

Ca. japonica

0.39 ± 0.01 b

125.4 ± 4.0 a

4.14 ± 0.58 b

1.35 ± 0.10 c

I. rotunda

0.26 ± 0.01 d

67.2 ± 4.9 c

8.85 ± 0.70 a

2.09 ± 0.12 b

Means with the same letter are not significantly different (ANOVA, P < 0.05)

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Figure captions

363

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Fig. 1. Aerial view and map of Nishinomiya Shrine in southeastern Hyogo Prefecture, Japan. The

365

evergreen broadleaved forest is located on the north and west sides of the shrine grounds. A 0.65 ha

366

contiguous research plot was established comprising 65 sub-plots (10 × 10 m). Of these tree locations

367

were mapped for the 40 × 100 m area (filled squares).

368

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Fig. 2. Stem map (all trees taller than 1.3 m) of the 40 × 100 m area of the shrine forest.

370

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Fig. 3. Box-whisker plot of the distance of individual trees of each species from the intensively

372

managed, western forest edge. Box length is the inter-quartile range. Lines and X inside boxes

373

represent median and mean distances, respectively. Bars represent 10 and 90 percentiles. Circles are

374

trees outside this range. Species are listed in order of increasing mean distance from forest edge.

375

Letter indicate significantly different means (P < 0.05, ANOVA and Tukey’s HSD).

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Fig.4. Spatial distribution pattern of each species in the shrine forest (solid lines). The horizontal line

378

indicates g(r) = 1. Dotted lines indicate 95% confidence interval for null hypothesis of for Poisson

379

random distribution. Positive and negative deviations outside the confidence intervals indicate

380

clustered and regular distribution patterns, respectively. Species order follows that of Fig. 3.

381

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Fig. 5. Spatial distribution pattern of L. lucidum in relation to each of the five native species (solid

383

lines). Dotted lines indicate 95% confidence interval for null hypothesis of for Poisson random

384

distribution. Positive and negative deviations outside the confidence intervals indicate sympatric and

385

exclusive distribution patterns, respectively. Species order follows that of Fig. 3.

386

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Fig. 6. Leaf traits in relation to canopy openness for L. lucidum and five native species. Data points

388

for Nmass, LMA and thickness are the mean of five leaves at each sampling position, while Amax is the

389

mean of ten measurements taken at light saturation for one leaf. All standard errors were less than 3%.

390

Lines indicate significant regressions (P < 0.05).

391

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Fig.7. Results of principle component analysis of leaf functional traits of L. lucidum and the five

393

native species. Arrows indicate correlation vectors showing correlation coefficients of each trait in

394

relation to PC1 and PC2.

19

Road

Figure

Shrine grounds

10m

10m

Houses

Fig. 1

X L. lucidum

□ Ci. japonicum

◊ Q. glauca

+ T. gymnanthera

Δ Ca. japonica

○ I. rotunda

Distance (m)

X Others

Distance (m)

Fig. 2

Distance from forest edge (m)

L. lucidum

Ca. japonica

Q. glauca

Ci. japonicum T. gymnanthera

I. rotunda

Fig. 3

L. lucidum

10

Ci. japonicum

yabuni

tou

10

Q. glauca

g(r)

T. gymnanthera

ara

mokk

10

Ca. japonica

I. rotunda

kuro

tsuba

10

r (m)

10

Fig. 4

Ci. japonicum

-1 0

10

-2

-3

L(r)

Q. glauca

-1 0

10

-1 0

-2

-2

-3

-3

Ca. japonica

-1 0

T. gymnanthera

10

I. rotunda

10

-1 0

-2

-2

-3

-3

r (m)

10

Fig. 5

X L. lucidum

□ Ci. japonicum

◊ Q. glauca

+ T. gymnanthera

0.4

LMA (g m2)

Leaf thickness (mm)

○ I. rotunda

200

0.6

0.2

100

16

10

20

Canopy openness (%)

30

12

10

20

30

Canopy openness (%)

Nmass (%)

Amax (µmol m-2 s-2)

Δ Ca. japonica

10

20

Canopy openness (%)

30

10

20

Canopy openness (%)

30

Fig. 6

PC 2

I. rotuna

L. lucidum

T. gymnanthera

Q. glauca

Amax

Thickness

Nmass

-4

-3

-2

-1

PC 1

LMA

-1

Ca. japonica

-2

Ci. japonicum

-3

-4

Fig. 7

...