1
Biological and environmental legacies in study sites
2
Biological Legacies
3
Plant Species
4
A total of 4,289 individual plants belonging to 121 species were identified in the
5
reference ecosystem (BF), and 9,129 individual plants of 95 species were identified in the
6
four plantations. The number of species, Margalef index (d), Shannon-Wienner index (H),
7
Simpson index (D), and Pielou index (J) are listed in Table 1. The species number in the tree
8
layer was significantly higher in BF than in the four plantations, but species number did not
9
significantly differ in the herb layer. The values of d and H in the tree layer were significantly
10
higher in BF than in the four plantations, but these values did not significantly differ in the
11
herb layer. Indices D and J in the tree and herb layer were similar in BF and the four
12
plantations. All diversity indices in the shrub layer were significantly lower in MCP than in
13
the other forests.
14
Soil Seed Banks
15
A total of 6,010 seedlings belonging to 81 species were identified, and 63 species were
16
observed in the plantations. The number of species in the soil seed bank was larger in the
17
plantations than in BF. In addition, the values of diversity indices tended to be slightly higher
18
in EP than in the other plantations (Table 1).
19
The composition of soil seed banks was less complex in the four plantations than in BF
20
because, unlike BF, the plantation soil seed banks lacked vine species (Fig. 1a). The
21
proportion of herb species in the soil seed banks was much higher in the four plantations than
22
in BF, but the opposite was true for the proportions of shrub and tree species.
23
Soil Microbes
24
Soil microbes were more abundant in the 0-5 cm soil layer than in the 5-20 cm soil layer
25
in both the wet and dry season (Table 2). In the wet season, the contents of total PLFAs,
26
bacterial PLFAs, and fungal PLFAs in the 0-5 cm layer was lowest in MCP but did not differ
27
between the other plantations and BF. Fungal PLFAs in the 5-20 cm layer were significantly
28
higher in MLP than in BF, but bacterial PLFAs in the 5-20 cm layer did not significantly
29
differ among the five forests. In the dry season, total, bacterial, and fungal PLFAs were
30
significantly lower in BF than in the four plantations in the 0-5 cm layer but did not
31
significantly differ in the 5-20 cm layer.
32
The percentage of protozoa and microalgae in the 0-5 cm layer was much lower in MCP
33
than in the other four forests but the opposite was true in the 5-20 cm layer (Fig. 1b). The soil
34
microbial compositions of the other three plantations were similar to that of BF.
35
Soil Animals
36
Soil animals were represented by 32 orders in all five forests and by 25 in the four
37
plantations. The number of orders and values for d, H, D, and J were higher in BF than in the
38
plantations but tended to be similar among the plantations (Table 1).
39
Birds
40
A total of 59 bird species including non-forest species and forest-dependent species were
41
recorded, and 29 species were recorded in the plantations. Species number and d value were
42
highest in BF and lowest in MCP (Table 1). All diversity indices of the plantations other than
43
MCP were higher than those in BF.
44
Environmental legacies
45
Soil Age
46
We used one excavated soil profile to represent BF and a second profile to represent the
47
degraded hilly substrate of the plantations because the substrate was similar among the four
48
plantations. Soil C was older in the plantations than in BF except at 30-50 cm (Table 3). The
49
age of soil C tended to increase with depth.
50
Soil Pollen
51
A total of 64 genera were detected in the soil of the five forests, and 57 genera were
52
detected in the plantations. A high percentage of the pollen was from trees and shrubs (Fig. 2).
53
In contrast to the soil profiles from the plantations, the BF soil profile contained a high
54
percentage of Castanopsis/Lithocarpus, Gironniera, and Aporusa pollen. In addition, the
55
percentage of Poaceae in the BF soil profile was relatively small. The percentages of
56
Eucalyptus, Schima, and Mallotus pollen were high in the soil profiles of EP, MBP, and MLP.
57
The pollen of Pinus and Poaceae were abundant in the soil profiles from all four plantations,
58
and the percentage of Poaceae increased with soil depth. The number of pollen genera and the
59
d value were higher in EP and MBP than in BF and the other two plantations. The other
60
diversity indices tended to be similar among the five forests (Table 1).
61
Light Environment
62
The LAI averaged across all vegetation layers was significantly higher in BF than in the
63
plantations, and most of the difference was explained by the tree layer (Table 4). LAI of the
64
tree layer was significantly higher in BF than in the plantations. LAI of the shrub layer was
65
significantly lower in MBP than in BF. LAI of the herb layer was significantly higher in EP
66
and MBP than in BF
67
Soil Mineral Distribution
68
Fe content increased with soil depth in all five forests (Fig. 3). The contents of other
69
mineral elements fluctuated with the depth, especially in the profiles from the plantations. The
70
contents of K, Fe, and Ca were significantly higher in BF than in the plantations while the
71
contents of Mg and Na showed the opposite trend. The contents of all mineral elements were
72
relatively constant from 10 to 80 cm depth in the plantation soils.
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
Table 1. The number and diversity of plants, soil seed bank, soil animals, soil pollen, and
birds in the five forests.
97
Forest type
Indicator
Vegetation
Characteristic
Layer
No.
Tree
Shrub
Herb
Tree
Shrub
Herb
Tree
Shrub
Herb
Tree
Shrub
Herb
Tree
Shrub
Herb
d
H
D
J
Soil seed
bank
Soil
animals
Birds
Soil pollen
No.
0-10 cm
d
H
D
J
0-10 cm
0-10 cm
0-10 cm
0-10 cm
No.
0-15 cm
d
H
D
J
No.
d
H
D
J
No.
d
H
D
J
0-15 cm
0-15 cm
0-15 cm
0-15 cm
tree, shrub and herb
tree, shrub and herb
tree, shrub and herb
tree, shrub and herb
tree, shrub and herb
0-80cm
0-80cm
0-80cm
0-80cm
0-80cm
BF
EP
MLP
MBP
MCP
26.85±1.30a 11.44±0.84bc 7.67±0.55c 12.44±0.85b 11.11±0.45bc
9.50±0.88b 17.11±2.29a 15.89±1.43a 13.89±0.73ab 10.67±1.05b
7.15±1.00a
4.67±0.67a
4±0.29a
5.78±0.6a
4.44±0.24a
5.15±0.22a
2.71±0.2bc
1.93±0.14c
2.86±0.21b
2.51±0.13bc
2.50±0.18ab
3.11±0.4a
2.74±0.23ab 2.53±0.11ab
1.9±0.21b
1.91±0.23a
1.14±0.22a
1.09±0.1a
1.46±0.16a
1.67±0.14a
2.45±0.08a
1.93±0.08b
1.52±0.08c
1.9±0.08b
1.87±0.05b
1.71±0.07a
2.04±0.13a
2.03±0.08a
1.92±0.05a
1.3±0.12b
1.51±0.14a
0.89±0.17a
0.95±0.09a
1.33±0.14a
1.24±0.08a
0.84±0.02a
0.8±0.03ab
0.71±0.04b
0.8±0.02ab
0.8±0.01ab
0.78±0.02a
0.81±0.02a
0.81±0.02a
0.79±0.02a
0.57±0.05b
0.72±0.05a
0.47±0.09a
0.54±0.06a
0.68±0.06a
0.76±0.07a
0.75±0.02a
0.8±0.03a
0.76±0.04a
0.76±0.02a
0.78±0.01a
0.78±0.02a
0.74±0.01a
0.74±0.02a
0.73±0.02a
0.56±0.04b
0.82±0.04a
0.6±0.09a
0.7±0.05a
0.77±0.04a
0.84±0.05a
23
4.61
2.71
0.92
0.86
33
4.83
2.27
0.85
0.65
31
4
1.78
0.76
0.52
33
4.58
1.66
0.65
0.47
37
4.66
1.86
0.74
0.51
27
2.46
1.49
0.7
0.45
35
4.74
0.60
2.13
0.76
38
3.98
1.92
0.79
0.53
20
1.86
0.82
0.37
0.28
13
2.78
0.83
2.12
0.85
43
4.39
1.85
0.79
0.49
23
2.12
0.82
0.4
0.26
18
3.77
0.86
2.48
0.90
36
3.7
1.92
0.8
0.54
20
1.82
0.83
0.4
0.28
15
3.14
0.87
2.36
0.89
46
4.45
1.67
0.74
0.44
19
1.7
0.71
0.34
0.24
6
1.00
0.43
0.76
0.34
36
3.66
1.79
0.77
0.50
98
99
100
101
102
103
104
105
106
107
Note: values are means ± SE. Means within rows sharing the same letter are not significantly
different (P < 0.05). BF: Broad-leaved forest; EP: eucalyptus plantation; MBP: mixed
broad-leaved plantation; MLP: mixed legume plantation; MCP: mixed conifer plantation.
No.: number of species, d: Margalef index, H: Shannon-Wienner index, D: Simpson index,
J: Pielou index.
108
Table 2. The content of total, bacterial, and fungal PLFAs (ng/g) in two soil layers of the five forests.
109
110
Forest type
Season
Wet season
Layer
(cm)
0-5
5-20
Dry season
0-5
5-20
127
128
129
130
131
132
111
112
Category
Total PLFAs
Bacteria
Fungi
Total PLFAs
Bacteria
Fungi
Total PLFAs
Bacteria
Fungi
Total PLFAs
Bacteria
Fungi
113
BF
EP
MLP
MBP
MCP
5625.63±362.73a
4362.99±305.35a
427.97±32.47ab
1927.2±231.72ab
1343.79±196.21a
107.9±15.96b
5474.48±705.87b
4566.7±607.03b
423.66±80.24b
3802.58±702.2a
3055.09±607.31a
283.38±66.64a
5597.16±259.02a
4367.32±213.17a
504.68±41.55a
2520.08±149.08a
1823.38±120.45a
171.67±7.43ab
8980.62±204.38a
6694.51±297.4a
903.45±61.81a
2956.5±266.16a
2317.3±208.08a
201.84±12.89a
5893.8±317.95a
4372.69±250.76a
527.67±50.27a
2597.49±55.27a
1720.97±74.07a
195.18±6.14a
8486.83±105.72a
7175.11±83.36a
943.86±11.3a
3019.77±123.39a
2350.15±92.89a
255.82±8.32a
5006.11±124.95ab
3624.95±66.62ab
408.82±24.77ab
2058.51±159.72ab
1480.44±116.43a
129.78±6.74ab
9000.08±641.23a
7620.5±554.78a
979.69±80.2a
3024.34±174.93a
2416.43±185.04a
224.78±8.78a
114
4134.29±445.46b
115
2975.83±389.44b
116
334.33±39.41b
117
1652.44±241.53b
118
1330.45±87.29a
119
129.54±8.44ab
120
7825.1±707.62a
121
7736.54±483.73a
122
791.39±98.84a
123
3270.01±663.01a
124
1895.99±99.33a
261.7±55.74a125
126
Note: values are means ± SE. Means within rows sharing the same letter are not significantly different (P < 0.05).
BF: Broad-leaved forest; EP: eucalyptus plantation; MBP: mixed broad-leaved plantation; MLP: mixed legume plantation; MCP: mixed conifer
plantation.
Table 3. 14C dating (a BP) of soil profiles (0–80 cm depth) from the reference forest (BF)
and from the four plantations (one profile was assumed to represent all four plantations).
Forest type
Layer (cm)
BF
Plantation
0-5
5-10
10-15
15-20
769
474
852
20-30
755
894
30-40
1271
1248
40-50
1933
1634
50-60
2043
2300
60-70
2552
2800
70-80
2105
4074
Table 4. Leaf areas index (LAI) of tree layer, shrub layer, herb layer, and all layers in the
five forests.
Layer
All layers
Tree
Shrub
Herb
BF
5.59±0.12a
4.39±0.18a
0.66±0.07a
0.54±0.10b
Forest type
EP
2.77±0.16d
0.83±0.14d
0.64±0.13a
1.29±0.12a
MLP
4.57±0.09b
2.7±0.09b
0.97±0.12a
0.9±0.12ab
MBP
2.56±0.04d
1.27±0.15c
0.23±0.08b
1.07±0.13a
MCP
3.14±0.15c
1.48±0.12c
0.87±0.18a
0.79±0.19ab
Note: Values are means ± SE. Means within rows sharing the same letter are not significantly
different (P < 0.05). BF: Broad-leaved forest; EP: eucalyptus plantation; MBP: mixed
broad-leaved plantation; MLP: mixed legume plantation; MCP: mixed conifer plantation.
Fig. 1. The composition of soil seed banks and soil microbial communities in the five forests.
BF: broad-leaved forest, EP: eucalyptus plantation, MLP: mixed-legume plantation,
MBP: mixed broad-leaved species plantation, MCP: mixed-coniferous plantation.
a. Composition of the soil seed banks.
b. Composition of the soil microbial community including protozoa and microalgae at two
soil depths.
Other: protozoa and microalgae.
Fig. 2. Pollen diagram (0-80 cm soil depth) of the five forests. Percentages were calculated
based on the total number of pollen of all terrestrial seed plants.
Notes: + :< 0.5%. BF: broad-leaved forest, EP: eucalyptus plantation, MLP: mixed-legume
plantation, MBP: mixed broad-leaved species plantation, MCP: mixed-coniferous
plantation.
Fig. 3. The distribution of mineral elements with soil depth in the five forests.
BF: broad-leaved forest, EP: eucalyptus plantation, MLP: mixed-legume plantation, MBP:
mixed broad-leaved species plantation, MCP: mixed-coniferous plantation.
100
100
a
80
80
70
70
Vine
60
Tree
50
Shrub
40
Herb
50
20
20
10
10
EP
MLP
Forest type
MBP
MCP
Other
Fungi
40
30
BF
Fig.1
60
30
0
b
90
Percent
Percent
90
Becteria
0
BF
EP
MLP MBP MCP
0-5 cm layer
BF
EP
MLP MBP MCP
5-20 cm layer
Layer and forest type
Fig. 2
45
120
35
K
110
30
Fe
Al
40
100
25
35
90
30
80
20
25
70
15
20
60
Content (g/kg)
10
5
.6
50
15
40
25
10
5
Ca
Na
Mg
.5
20
4
15
3
Y Data
.4
.3
10
2
.2
1
5
.1
0
0
0.0
0~5
5~10 10~20 20~40 40~80
0~5
Depth (cm)
180
160
Mn
BF
EP
MLP
MBP
MCP
140
Content (mg/kg)
5~10 10~20 20~40 40~80
120
100
80
60
40
20
0
0~5
5~10 10~20 20~40 40~80
Depth (cm)
Fig.3
0~5
5~10 10~20 20~40 40~80