Yellow birch
23%
1 000 000
Sugar maple
100 000
Density (per ha)
Striped maple 1%
Pin cherry 2%
Density: 549 trees per ha
(> 10 cm DBH)
Balsam fir 3%
Red spruce 4%
Sugar maple
Paper birch 5%
35%
Beech
Yellow birch
10 000
1000
100
10
Seedling Sapling 2 to 10 11 to 20 21 to 70
cm DBH cm DBH cm DBH
Beech
27%
300
70
DBH > 10 cm
DBH > 10 cm
60
Density (per ha)
Density (per ha)
250
200
150
100
50
40
30
20
10
0
0
Low
Middle
(546-625 m)
Low
Upper
Beech
Yellow birch
Biomass
Foliage and fruit 332
Branch
Stem
Root
Paper birch
Red spruce
Balsam fir
Pin cherry
Upper
(709-791 m)
Striped maple
Production And Decomposition
Foliage and fruit
322
3856
Branch
254
9094
Stem
266
Root
2826
Total Living: 16 108 g m-2
Ecological Climatology © 2008 G. Bonan
Middle
(546-625 m) (625-709 m)
(625-709 m) (709-791 m)
Sugar maple
Figure 21.1
50
Decomposition
160
680
Net Primary Production: 1002 g m-2 yr-1
Net Ecosystem Production: 322 g m-2 yr-1
Gross Primary
Production
Consumers
Grazing
Autotrophic
Respiration
Tree
Shrub
Leaf
Leaf
Stem
Stem
Root
Root
Heterotrophic
Respiration
Litter
Litterfall
Humus
Decomposers
Respiration
Figure 21.2
Ecological Climatology © 2008 G. Bonan
Stand Structure
Quaking Aspen
Black Spruce
Age: 67 years
Height: 20 m
Density: 980 ha-1
Mean diameter: 20 cm
LAI: 3.3 m2 m-2
Jack Pine
Age: 115 years
Height: 7 m
Density: 5900 ha-1
Mean diameter: 7 cm
LAI: 5.6 m2 m-2
Age: 65 years
Height: 13 m
Density: 1190 ha-1
Mean diameter: 13 cm
LAI: 2.4 m2 m-2
Living Biomass
Foliage 94
Branch
Detritus
Forest floor
Mineral soil
Branch
675
Foliage 103
968
Branch
Stem
3633
Stem
Understory 72
Standing dead
505
897
8271
Stem
Foliage
Understory 51
Moss 60
Standing dead
616
1943
3599
Total Living:
9334 g C m-2
Total Detritus:
6510 g C m-2
Total Ecosystem: 15 844 g C m-2
Mineral soil 39 036
Total Living:
4924 g C m-2
Total Detritus:
39 652 g C m-2
Total Ecosystem: 44 576 g C m-2
413
2576
Lichen
Standing dead
347 Understory 16
567
Forest floor
1456
Mineral soil
1420
Total Living:
3455 g Cm-2
Total Detritus:
3443 g C m-2
Total Ecosystem: 6898 g C m-2
Figure 21.3
Ecological Climatology © 2008 G. Bonan
Tree Production (g m-2 yr-1)
Tree Production (g m-2 yr-1)
2500
2000
1500
1000
500
0
1200
1100
1000
900
800
700
0
500
1000
1500
Elevation (meters)
2000
Low
(546-625 m)
Middle
Upper
(625-709 m) (709-791 m)
Figure 21.4
Ecological Climatology © 2008 G. Bonan
3.5
3.0
1600
New Foliage (kg)
Aboveground Net Primary
Production (g m-2 yr-1)
1800
1400
1200
1000
2.5
2.0
1.5
1.0
0.5
800
0.0
1984
1985
1986
Pre-Treatment
Treated
Treated
0
Untreated
5
10
15
20
25
Stem Diameter (cm)
Irrigated
Fertilized
Figure 21.5
Ecological Climatology © 2008 G. Bonan
Biomass (g m-2)
35 000
30 000
Total
25 000
Forest Floor And Soil
20 000
15 000
10 000
Tree And Understory
5000
0
0
5
10
15
20
25
30
35
Leaf Area Index (m2 m-2)
7
1400
LAI
6
1200
5
1000
4
800
3
600
NPP
2
400
1
200
0
0
0
5
10
15
20
25
30
Aboveground NPP (g m-2 yr-1)
Stand Age (years)
35
Stand Age (years)
Figure 21.6
Ecological Climatology © 2008 G. Bonan
Aboveground NPP
Total NPP
Belowground
392 g C m-2 yr-1
10%
Understory
7%
Understory
15%
Quaking Aspen
Belowground
Wood
50%
Foliage
21%
Aboveground
90%
Wood
7%
Understory Moss
7%
8%
286 g C m-2 yr-1
Foliage
25%
Aboveground
58%
Foliage
5%
Belowground
222 g C m-2 yr-1
Jack Pine
47%
Aboveground
53%
166 g C m-2 yr-1
Wood
48%
Black Spruce
42%
352 g C m-2 yr-1
Foliage
32%
Wood
7%
Understory
117 g C m-2 yr-1
4%
Wood
51%
Foliage
2%
Wood
11%
Figure 21.7
Ecological Climatology © 2008 G. Bonan
Distance (km)
0
25
2000
75
1400 g C m-2 yr-1
1800
1600
Elevation (meters)
50
1400
100
125
1665 g C m-2 yr-1
150
2404 g C m-2 yr-1
175
200
879 g C m-2 yr-1
225
364 g C m-2 yr-1
302 g C m-2 yr-1
2500
2500
2500
2500
2500
2500
2000
2000
2000
2000
2000
2000
1500
1500
1500
1500
1500
1500
1000
1000
1000
1000
1000
1000
500
500
500
500
500
500
0
0
0
0
0
0
1200
Carbon Allocation (g C m-2 yr-1)
1000
Growth Respiration
Maintenance Respiration (Foliage)
Maintenance Respiration (Sapwood)
Aboveground NPP
Belowground Allocation
800
600
400
200
124.0
Figure 21.8
123.5
6
P
T
123.0
8
0
6
P
T
122.5
Longitude (W)
10
1000
8
0
6
P
T
122.0
Height (m)
Height (m)
Height (m)
Height (m)
0
1000
2000
3000
12
2000
10
1000
8
0
P
T
6
121.5
3000
12
2000
10
1000
8
0
P
T
6
Temperature (C)
T
10
12
Juniper woodland
Biomass: 1.1 kg m-2
LAI: 0.4
50
40
30
20
10
0
Precipitation (mm)
6
P
8
2000
3000
50
40
30
20
10
0
Temperature (C)
0
1000
12
Ponderosa pine
Biomass: 1.5 kg m-2
LAI: 0.9
Precipitation (mm)
8
10
3000
50
40
30
20
10
0
Temperature (C)
1000
2000
50
40
30
20
10
0
Mountain hemlock
Biomass: 37.0 kg m-2
LAI: 1.9
Precipitation (mm)
10
12
Precipitation (mm)
2000
3000
Temperature (C)
Height (m)
12
50
40
30
20
10
0
Precipitation (mm)
3000
Temperature (C)
Height (m)
Precipitation (mm)
50
40
30
20
10
0
Douglas fir and
Western hemlock
Biomass: 40.8 kg m-2
LAI: 8.6
Douglas fir
Biomass: 47.1 kg m-2
LAI: 5.3
Temperature (C)
Old growth Sitka spruce
and Western hemlock
Biomass: 71.1 kg m-2
LAI: 6.4
0
121.0
Ecological Climatology © 2008 G. Bonan
Calcium
Nitrogen
Precipitation
0.22 g m-2 yr-1
Precipitation
0.65 g m-2 yr-1
Translocation
0.51 g m-2 yr-1
Aboveground Biomass
Pool: 35.1 g m-2
Gain: 0.48 g m-2 yr-1
Litterfall
4.07 g m-2 yr-1
Throughfall
Stemflow
0.67 g m-2 yr-1
Root Litter
0.32 g m-2 yr-1
Soil Available
51.0 g m-2
Vegetation Uptake
6.22 g m-2 yr-1
Belowground Biomass
Pool: 18.1 g m-2
Gain: 0.42 g m-2 yr-1
Mineralization
4.24 g m-2 yr-1
Weathering
2.11 g m-2 yr-1
Mineral Soil: 960 g m-2
Rock: 6460 g m-2
Litterfall
5.42 g m-2 yr-1
Throughfall
Stemflow
0.27 g m-2 yr-1
0.66 g m-2 yr-1
Hydrologic Export
1.39 g m-2 yr-1
Forest Floor
Pool: 37.2 g m-2
Gain: 0.14 g m-2 yr-1
Root Exudate
0.35 g m-2 yr-1
Organically-bound Ca
Inorganic Ca
Denitrification
?
Translocation
3.98 g m-2 yr-1
Aboveground Biomass
Pool: 38.3 g m-2
Gain: 0.54 g m-2 yr-1
Belowground Biomass
Pool: 10.1 g m-2
Gain: 0.27 g m-2 yr-1
N Fixation
1.42 g m-2 yr-1
Root Litter
0.62 g m-2 yr-1
Root Exudate
0.01 g m-2 yr-1
0.08 g m-2 yr-1
Hydrologic Export
0.40 g m-2 yr-1
Forest Floor
Pool: 125.6 g m-2
Gain: 0.77 g m-2 yr-1
Mineralization
6.97 g m-2 yr-1
Soil Available
Vegetation Uptake
7.96 g m-2 yr-1
Organic N
Inorganic N
Figure 21.9
Ecological Climatology © 2008 G. Bonan
Chestnut Oak
Yellow Poplar
Foliage 350
Foliage 420
Branch
Bole
Branch
3770
14 380
Stump 1220
Total Biomass: 19 770 g m-2
Bole
3570
13 560
Stump 1170
Total Biomass: 18 640 g m-2
Oak-Hickory
Foliage 340
Branch
Bole
3140
11 910
Stump 1040
Total Biomass: 16 380 g m-2
Figure 21.10
Ecological Climatology © 2008 G. Bonan
Chestnut Oak
Yellow Poplar
Translocation
0.1 g m-2 yr-1
Translocation
0.5 g m-2 yr-1
Oak-Hickory
Calcium
Foliage
Pool: 5.8 g m-2
Branch and Bole
Pool: 88.7 g m-2
Gain: 2.5 g m-2 yr-1
Litterfall
4.5 g m-2 yr-1
Throughfall
Stemflow
1.2 g m-2 yr-1
Mortality
0.3 g m-2 yr-1
Foliage
Pool: 7.5 g m-2
Mortality
0.4 g m-2 yr-1
Branch and Bole
Pool: 51.7 g m-2
Gain: 1.1 g m-2 yr-1
Forest Floor
Pool: 31.8 g m-2
Vegetation Uptake
8.2 g m-2 yr-1
Translocation
0.6 g m-2 yr-1
Foliage
Pool: 7.0 g m-2
Litterfall
5.8 g m-2 yr-1
Throughfall
Stemflow
1.2 g m-2 yr-1
Forest Floor
Pool: 29.4 g m-2
Vegetation Uptake
8.1 g m-2 yr-1
Branch and Bole
Pool: 87.4 g m-2
Gain: 2.8 g m-2 yr-1
Throughfall
Stemflow
1.5 g m-2 yr-1
Mortality
0.7 g m-2 yr-1
Forest Floor
Pool: 41.7 g m-2
Vegetation Uptake
9.2 g m-2 yr-1
Precipitation
1.3 g m-2 yr-1
Precipitation
1.3 g m-2 yr-1
Precipitation
1.3 g m-2 yr-1
Litterfall
4.9 g m-2 yr-1
Nitrogen
Translocation
3.8 g m-2 yr-1
Foliage
Pool: 7.5 g m-2
Branch and Bole
Pool: 35.6 g m-2
Gain: 2.5 g m-2 yr-1
Litterfall
3.4 g m-2 yr-1
Throughfall
Stemflow
0.3 g m-2 yr-1
Mortality
0.1 g m-2 yr-1
Foliage
Pool: 7.8 g m-2
Mortality
0.2 g m-2 yr-1
Branch and Bole
Pool: 18.9 g m-2
Gain: 2.5 g m-2 yr-1
Forest Floor
Pool: 29.8 g m-2
Vegetation Uptake
6.2 g m-2 yr-1
Translocation
2.6 g m-2 yr-1
Translocation
3.9 g m-2 yr-1
Throughfall
Stemflow
0.3 g m-2 yr-1
Forest Floor
Pool: 18.7 g m-2
Vegetation Uptake
6.4 g m-2 yr-1
Foliage
Pool: 6.7 g m-2
Litterfall
3.6 g m-2 yr-1
Branch and Bole
Pool: 33.2 g m-2
Gain: 2.6 g m-2 yr-1
Litterfall
3.7 g m-2 yr-1
Throughfall
Stemflow
0.4 g m-2 yr-1
Mortality
0.3 g m-2 yr-1
Forest Floor
Pool: 33.4 g m-2
Vegetation Uptake
6.7 g m-2 yr-1
Figure 21.11
Ecological Climatology © 2008 G. Bonan
High Nitrogen
Uptake
High Nitrogen
Availability
High Foliage Nitrogen
Concentration
High Net Primary
Production
High Mineralization
Rate
Low Nitrogen
Uptake
Low Nitrogen
Availability
High Nitrogen
Return In Litter
Low Foliage Nitrogen
Concentration
Low Net Primary
Production
Low Mineralization
Rate
Low Nitrogen
Return In Litter
Figure 21.12
Ecological Climatology © 2008 G. Bonan
Sugar
Maple
Red Oak
900
Sugar
Maple
White Oak
800
Red Oak
700
White
Pine
600
Hemlock
500
Red
Pine
400
2
3
4
5
6
7
8
9
Nitrogen Mineralization (g N m-2 yr-1)
Aboveground Net Primary
Production (g m-2 yr-1)
1000
9
8
Sugar
Maple
Sugar
Maple
7
6
White
Oak
Red Oak
5
4
Red Oak
White Pine
3
2
1
0
30
35
40
P Return In Litter (g P m-2 yr-1)
N Return In Litter (g N m-2 yr-1)
Nitrogen Mineralization (g N m-2 yr-1)
5
Red Oak
4
Red Oak
3
Sugar
Maple
White Pine
Red Pine
Sugar
Maple
White Oak
2
Hemlock
1
0
Red Pine
Hemlock
45
50
55
60
Carbon:Nitrogen Ratio
65
70
0.5
Red Oak
0.4
Red Oak
Sugar
Maple
Sugar
Maple
White Oak
0.3
White Pine
Red Pine
0.2
Hemlock
0.1
0
2
3
4
5
6
Nitrogen Mineralization (g N
7
m-2
8
yr-1)
9
2
3
4
5
6
7
8
-2
-1
Nitrogen Mineralization (g N m yr )
9
Figure 21.13
Ecological Climatology © 2008 G. Bonan
Figure 21.14
Ecological Climatology © 2008 G. Bonan
Black Spruce
Biomass
Nitrogen
4990 g m-2
12.3 g N m-2
110 g
4.0 (0.72%)
Branch
788
Branch
3.2 (0.41%)
Branch
Trunk
5.1 (0.14%)
Trunk
70
Trunk
0.14
Litterfall
43
Litterfall
0.30
3643
Forest
Floor
7650
63 (0.82%)
1248
8.5 (0.68%)
1636
8.9 (0.54%)
Foliage
7
Branch
0.26
9357
209
6.1 (0.40%)
10 319
2200
15.6 (0.15%)
27 (1.23%)
6.54 g N m-2 yr-1
36
5.9
0.36
0.28
240
1.48
156
553 g m-2 yr-1
5.4 g N m-2 yr-1
4.6
269
4.6 (1.71%)
1519
1.89
286
69 (1.45%)
26.3 g N m-2
269
0.25
251
15.0 (0.15%)
4750
12 107 g m-2
0.36
562 g m-2 yr-1
5.7 (0.32%)
10 248
5.2
26
5.9 (2.06%)
1775
5.81 g N m-2 yr-1
234
86 (1.48%)
26.6 g N m-2
286
0.93
155
15.7 (0.17%)
5800
12 309 g m-2
0.16
197
469 g m-2 yr-1
7.3 (0.45%)
0.07
1.14
36
5.2 (2.22%)
1630
0.32
1.56 g N m-2 yr-1
133
57 (0.77%)
28.2 g N m-2
234
Figure 21.15
33
10.0 (0.07%)
7426
11 221 g m-2
Foliage
366 g m-2 yr-1
27.4 g N m-2
13 917
Balsam Poplar
0.53 g N m-2 yr-1
Foliage
16 801 g m-2
Quaking Aspen
N Uptake
yr-1
559
Forest Floor
Paper Birch
m-2
Foliage
Trunk
White Spruce
NPP
57
0.56
0.24
227
389
3.31
Ecological Climatology © 2008 G. Bonan
35
30
Black Spruce
25
20
Paper Birch
15
10
Quaking Aspen
Tree Production (g m-2 yr-1)
Forest Floor Thickness (cm)
40
White Spruce
Balsam Poplar
5
0
0
500
1000
1500
2000
2500
800
700
600
500
400
300
200
100
0
600
800
1000 1200 1400 1600 1800
Soil Degree-Days
18
16
14
12
10
8
6
4
2
0
500
Tree Production (g m-2 yr-1)
Annual Decomposition (%)
Soil Degree-Days
750
1000
1250
800
700
600
500
400
300
200
100
0
1500
0
7
6
5
4
3
2
1
0
50
Figure 21.16
75
100
125
2
3
4
N Mineralization (g
Litterfall Biomass/Litterfall N
N Requirement (g m-2 yr-1)
Soil Degree-Days
1
150
Forest Floor Biomass/Forest Floor N
5
m-2
6
7
yr-1)
170
160
150
140
130
120
110
100
50
75
100
125
150
Forest Floor Biomass/Forest Floor N
Ecological Climatology © 2008 G. Bonan
Net Ecosystem Production
(g C m-2 day-1)
Uptake
Release
8
6
4
2
0
-2
-4
-6
-8
Carbon Flux (g C m-2 day-1)
Uptake
Release
1992 1993 1994 1995 1996 1997 1998 1999
15
Respiration
10
5
0
-5
-10
GPP
-15
Uptake
Cumulative NEP (g C m-2)
1992 1993 1994 1995 1996 1997 1998 1999
Figure 21.17
200
0
-200
-400
-600
-800
-1000
-1200
-1400
-1600
-1800
1992 1993 1994 1995 1996 1997 1998 1999
Ecological Climatology © 2008 G. Bonan
0
1200
-50
1100
-100
1000
-150
900
GPP
-200
800
R
-250
NEP
Net Ecosystem Production
(g C m-2 yr-1)
Gross Primary Production,
Ecosystem Respiration
(g C m-2 yr-1)
1300
700
-300
1992 1993 1994 1995 1996 1997 1998 1999
Figure 21.18
Ecological Climatology © 2008 G. Bonan
Release
Uptake
Net Ecosystem Production
(g C m-2 day-1)
2.0
1.5
1.0
0.5
0.0
-0.5
-1.0
-1.5
-2.0
-2.5
1994
1995
1996
1997
Carbon Flux (g C m-2 day-1)
Uptake
Release
15
Respiration
10
5
0
-5
-10
GPP
-15
1994
1995
1996
1997
1994
1995
1996
1997
Cumulative NEP (g C m-2)
Release
120
100
80
60
40
20
0
Figure 21.19
Ecological Climatology © 2008 G. Bonan
100
900
80
850
60
800
40
750
20
700
0
650
600
550
-20
GPP
R
NEP
-40
-60
Net Ecosystem Production
(g C m-2 yr-1)
Gross Primary Production,
Ecosystem Respiration
(g C m-2 yr-1)
950
500
-80
1995 1996 1997 1998 1999 2000 2001 2002 2003 2004
Figure 21.20
Ecological Climatology © 2008 G. Bonan