1
Supplementary Materials: Appendix 1. Studies used in our meta-analysis (see Appendix 2 for full references)
2
3
Table S1. Study sites for N enrichment experiments on aboveground biomass.
Country
Site
Argentina
Australia
Greece
Italy
Netherlands
Switzerland
United
Kingdom
USA
Venezuala
4
5
6
Latitude
Longitude
Altitude
Flooding
Pampa
-36.25
-58.27
13
W Victoria
Thessaloniki A
Thessaloniki B
Fabriano
Amerongse
Bennekomst A
Bennekomst B
Gerendal A
-37.80
40.80
40.80
43.33
51.98
52.02
52.02
50.85
143.02
21.38
21.38
13.00
5.43
5.60
5.60
5.90
Gerendal B
50.85
Utrecht A
Utrecht B
Alp
N Addition
g N m-2y-1
MAP
MAT
Bioclimatic
Zone
25
936
15.3
Grassland
Semmartin et al. (2007)
244
1340
1340
350
5
34
34
102
5
15
15
3.5
10
20
20
10
602
805
805
945
780
778
778
800
13.3
10.3
10.3
12.9
9.5
9.8
9.8
9.7
Morgan (2007)
Mamolos et al. (2005)
Mamolos et al. (2005)
Bonanomi et al. (2003)
Beltman et al. (2007)
van der Hoek et al. (2004)
van der Hoek et al. (2004)
Bobbink (1991)
5.90
102
10
800
9.7
52.07
52.07
9.65
5.58
5.58
46.53
11
11
2000
793
793
1200
9.4
9.4
2.8
Sourhope
55.48
-2.23
350
10
10
0.5, 1, 2.5,
5
12
Grassland
Grassland
Grassland
Grassland
Grassland
Grassland
Grassland
Temperate
forest
Temperate
forest
Grassland
Grassland
Boreal forest
1117
6.2
Cole et al. (2008)
CPER
40.83
-104.72
1650
24
312
8.7
Temperate
forest
Grassland
Irvine.Ranch
Jasper Ridge A
Konza
Niwot Ridge A
Niwot Ridge B
Sedgewick
IVIC
33.62
37.67
39.08
40.06
40.06
34.69
10.50
-117.76
-122.37
-96.58
-105.62
-105.62
-120.04
-66.90
23
120
378
3258
3258
500
1200
10
5
10
2, 4, 6
25
10
20
311
640
874
900
900
513
1063
16.5
13.6
12.4
-3.0
-3.0
13.7
17.9
Grassland
Grassland
Grassland
Boreal forest
Boreal forest
Grassland
Grassland
Reference
Bobbink (1991)
Aerts et al. (2003)
Aerts et al. (2003)
Bassin et al. (2007)
Paschke et al. (2000), Morgan et al. (2004), LeCain et al.
(2006)
Harpole et al. (2007)
Higgins et al. (2002), Henry et al. (2006)
Seastedt et al. (1991)
Bowman et al. (1993), Bowman et al. (2006)
Bowman et al. (1993), Bowman et al. (2006)
Harpole and Tilman (2007)
Barger et al. (2003)
7
8
Table S2. Study sites for N enrichment experiments on belowground biomass.
Country
Site
Argentina
Netherlands
Flooding.Pampa
USA
Venezuala
(b).
9
10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
Latitude
Longitude
Altitude
N addition
g N m-2y-1
MAP
MAT
-36.25
-58.27
13
25
936
15.3
Gerendal A
50.85
5.90
102
10
800
Gerendal B
50.85
5.90
102
10
CPER
Irvine.Ranch
Jasper Ridge
Niwot Ridge A
Niwot Ridge B
IVIC
40.83
33.62
37.67
40.06
40.06
10.50
-104.72
-117.76
-122.37
-105.62
-105.62
-66.90
1650
23
120
3258
3258
1200
10
10
7
25
25
20
Bioclimatic
Zone
Reference
Grassland
Semmartin et al. (2007)
9.7
Temperate forest
Bobbink (1991)
800
9.7
Temperate forest
Bobbink (1991)
312
311
640
900
900
1063
8.7
16.5
13.6
-3.0
-3.0
17.9
Grassland
Grassland
Grassland
Boreal forest
Boreal forest
Grassland
Paschke et al. (2000) Morgan et al. (2004), LeCain et al. (2006)
Harpole et al. (2007)
Higgins et al. (2002), Henry et al. (2006)
Bowman et al. (1993), Bowman et al. (2006)
Bowman et al. (1993), Bowman et al. (2006)
Barger et al. (2003)
30
31
Table S3. Study sites for CO2 enrichment experiments on aboveground biomass.
Country
Site
Canada
Germany
Farnham
Frottmaninger
Giessen
Rapolano
Warawatu
Uppsala
Bidmer
Jura
45.28
48.22
50.60
43.28
-40.23
59.80
46.58
47.55
72.98
11.60
8.70
11.58
175.27
17.63
8.38
7.57
CPER
Jasper Ridge A
Jasper Ridge B
40.83
37.67
37.40
Manhatten
39.12
Italy
New Zealand
Sweeden
Switzerland
USA
32
33
34
35
36
37
Latitude
Longitude
CO2
(ppm)
MAP
MAT
82
500
217
350
29
10
2470
520
625
660
440
580
475
700
680
600
1101
920
645
729
874
527
1782
809
6.0
10.5
9.4
13.2
12.9
5.5
-1.0
9.7
Temperate
forest
Grassland
Grassland
Grassland
Grassland
Grassland
Boreal forest
Grassland
Potvin and Vasseur (1997)
Winkler and Herbst (2003)
Kammann et al. (2005)
Marchi et al. (2004)
Edwards et al. (2001)
Marissink et al. (2002), Sindhoj et al. (2004)
Schappi and Körner (1996)
Leadley et al. (1999), Niklaus and Körner (2004)
-104.72
-122.37
-122.22
1760
120
224
720
680
716
321
640
632
8.0
13.6
13.6
Grassland
Grassland
Grassland
Paschke et al. (2000), Morgan et al. (2004), LeCain et al. (2006)
Higgins et al. (2002), Henry et al. (2006)
Higgins et al. (2002), Henry et al. (2006)
-96.35
324
708
840
12.4
Grassland
Ham et al. (1995), Owensby et al. (1999), Jastrow et al. (2000)
Bioclimatic
Zone
Reference
Table S4. Study sites for CO2 enrichment experiments on belowground biomass.
Country
Site
Latitude
Longitude
Italy
Sweden
Switzerland
Rapolano
Uppsala
Bidmer
43.28
59.80
46.58
11.58
17.63
8.38
350
16
2470
Jura
CPER A
CPER B
Jasper Ridge A
Jasper Ridge B
Manhatten
47.45
40.68
40.83
37.67
37.40
39.12
7.56
-104.75
-104.78
-122.37
-122.22
-96.35
520
1607
1760
120
224
324
USA
Altitude
Altitude
CO2
(ppm)
MAP
MAT
Bioclimatic
Zone
Reference
580
700
680
650
729
527
1782
13.2
5.5
-1.0
Marchi et al. (2004)
Marissink et al. (2002), Sindhoj et al. (2004)
Schappi and Körner (1996)
1078
383
320
640
632
840
9.3
8.8
8.0
13.6
13.6
12.4
Grassland
Grassland
Boreal forest
Temperate
forest
Grassland
Grassland
Grassland
Grassland
Grassland
720
720
680
716
708
Leadley et al. (1999), Niklaus and Körner (2004)
Paschke et al. (2000), Morgan et al. (2004), LeCain et al. (2006)
Paschke et al. (2000), Morgan et al. (2004), LeCain et al. (2006)
Higgins et al. (2002), Henry et al. (2006)
Higgins et al. (2002), Henry et al. (2006)
Ham et al. (1995), Owensby et al. (1999), Jastrow et al. (2000)
38
Appendix 2.
39
Studies used in meta-analysis
40
41
42
43
44
45
46
47
48
49
50
51
52
53
54
55
56
57
58
59
60
61
62
63
64
65
66
67
68
69
70
71
72
73
74
75
76
77
78
79
Aerts R., de Caluwe H. & Beltman B. (2003). Is the relation between nutrient supply and
biodiversity co-determined by the type of nutrient limitation? Oikos, 101, 489498.
Barger N.N., D'Antonio C.M., Ghneim T., Brink K. & Cuevas E. (2002). Nutrient
limitation to primary productivity in a secondary savanna in Venezuela.
Biotropica, 34, 493-501.
Bassin S., Volk M., Suter M., Buchmann N. & Fuhrer J. (2007). Nitrogen deposition but
not ozone affects productivity and community composition of subalpine grassland
after 3 yr of treatment. New Phytologist, 175, 523-534.
Beltman B., Willems J.H. & Gusewell S. (2007). Flood events overrule fertiliser effects
on biomass production and species richness in riverine grasslands. Journal of
Vegetation Science, 18, 625-634.
Bobbink R. (1991). Effects of Nutrient Enrichment in Dutch Chalk Grassland. Journal of
Applied Ecology, 28, 28-41.
Bonanomi G., Caporaso S. & Allegrezza M. (2006). Short-term effects of nitrogen
enrichment, litter removal and cutting on a Mediterranean grassland. Acta
Oecologica-International Journal of Ecology, 30, 419-425.
Bowman W.D., Gartner J.R., Holland K. & Wiedermann M. (2006). Nitrogen critical
loads for alpine vegetation and terrestrial ecosystem response: Are we there yet?
Ecological Applications, 16, 1183-1193.
Bowman W.D., Theodose T.A., Schardt J.C. & Conant R.T. (1993). Constraints of
Nutrient Availability on Primary Production in 2 Alpine Tundra Communities.
Ecology, 74, 2085-2097.
Cole L., Buckland S.M. & Bardgett R.D. (2008). Influence of disturbance and nitrogen
addition on plant and soil animal diversity in grassland. Soil Biology &
Biochemistry, 40, 505-514.
Edwards G.R., Clark H. & Newton P.C.D. (2001). The effects of elevated CO2 on seed
production and seedling recruitment in a sheep-grazed pasture. Oecologia, 127,
383-394.
Ham J.M., Owensby C.E., Coyne P.I. & Bremer D.J. (1995). Fluxes of Co2 and WaterVapor from a Prairie Ecosystem Exposed to Ambient and Elevated Atmospheric
Co2. Agricultural and Forest Meteorology, 77, 73-93.
Harpole W.S., Potts D.L. & Suding K.N. (2007). Ecosystem responses to water and
nitrogen amendment in a California grassland. Global Change Biology, 13, 23412348.
Harpole W.S. & Tilman D. (2007). Grassland species loss resulting from reduced niche
dimension. Nature, 446, 791-793.
Henry H.A.L., Chiariello N.R., Vitousek P.M., Mooney H.A. & Field C.B. (2006).
Interactive effects of fire, elevated carbon dioxide, nitrogen deposition, and
precipitation on a California annual grassland. Ecosystems, 9, 1066-1075.
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81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
100
101
102
103
104
105
106
107
108
109
110
111
112
113
114
115
116
117
118
119
120
121
122
123
124
125
Higgins P.A.T., Jackson R.B., Des Rosiers J.M. & Field C.B. (2002). Root production
and demography in a california annual grassland under elevated atmospheric
carbon dioxide. Global Change Biology, 8, 841-850.
Jastrow J.D., Miller R.M. & Owensby C.E. (2000). Long-term effects of elevated
atmospheric CO2 on below-ground biomass and transformations to soil organic
matter in grassland. Plant and Soil, 224, 85-97.
Kammann C., Grunhage L., Gruters U., Janze S. & Jager H.J. (2005). Response of
aboveground grassland biomass and soil moisture to moderate long-term CO2
enrichment. Basic and Applied Ecology, 6, 351-365.
Leadley P.W., Niklaus P.A., Stocker R. & Körner C. (1999). A field study of the effects
of elevated CO2 on plant biomass and community structure in a calcareous
grassland. Oecologia, 118, 39-49.
LeCain D.R., Morgan J.A., Milchunas D.G., Mosier A.R., Nelson J.A. & Smith D.P.
(2006). Root biomass of individual species, and root size characteristics after five
years of CO2 enrichment on native shortgrass steppe. Plant and Soil, 279, 219228.
Mamolos A.P., Vasilikos C.V. & Veresoglou D.S. (2005). Vegetation in contrasting soil
water sites of upland herbaceous grasslands and N : P ratios as indicators of
nutrient limitation. Plant and Soil, 270, 355-369.
Marchi S., Tognetti R., Vaccari F.P., Lanini M., Kaligaric M., Miglietta F. & Raschi A.
(2004). Physiological and morphological responses of grassland species to
elevated atmospheric CO2 concentrations in FACE-systems and natural CO2
springs. Functional Plant Biology, 31, 181-194.
Marissink M. & Hansson M. (2002). Floristic composition of a Swedish semi-natural
grassland during six years of elevated atmospheric CO2. Journal of Vegetation
Science, 13, 733-742.
Morgan J.A., Mosier A.R., Milchunas D.G., LeCain D.R., Nelson J.A. & Parton W.J.
(2004). CO2 enhances productivity, alters species composition, and reduces
digestibility of shortgrass steppe vegetation. Ecological Applications, 14, 208219.
Morgan J.W. (2007). Relationship between fire frequency and nitrogen limitation on
foliage production in a native grassland community in Victoria, Australia.
Rangeland Journal, 29, 101-105.
Niklaus P.A. & Körner C. (2004). Synthesis of a six-year study of calcareous grassland
responses to in situ CO2 enrichment. Ecological Monographs, 74, 491-511.
Owensby C.E., Ham J.M., Knapp A.K. & Auen L.M. (1999). Biomass production and
species composition change in a tallgrass prairie ecosystem after long-term
exposure to elevated atmospheric CO2. Global Change Biology, 5, 497-506.
Paschke M.W., McLendon T. & Redente E.F. (2000). Nitrogen availability and old-field
succession in a shortgrass steppe. Ecosystems, 3, 144-158.
Potvin C. & Vasseur L. (1997). Long-term CO2 enrichment of a pasture community:
Species richness, dominance, and succession. Ecology, 78, 666-677.
Schappi B. & Körner C. (1996). Growth responses of an alpine grassland to elevated
CO2. Oecologia, 105, 43-52.
Seastedt T.R., Briggs J.M. & Gibson D.J. (1991). Controls of Nitrogen Limitation in
Tallgrass Prairie. Oecologia, 87, 72-79.
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128
129
130
131
132
133
134
135
136
137
138
139
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141
142
143
144
145
146
147
148
149
150
151
Semmartin M., Oyarzabal M., Loreti J. & Oesterheld M. (2007). Controls of primary
productivity and nutrient cycling in a temperate grassland with year-round
production. Austral Ecology, 32, 416-428.
Sindhoj E., Andren O., Katterer T., Marissink M. & Pettersson R. (2004). Root biomass
dynamics in a semi-natural grassland exposed to elevated atmospheric CO2 for
five years. Acta Agriculturae Scandinavica Section B-Soil and Plant Science, 54,
50-59.
van der Hoek D., van Mierlo A. & van Groenendael J.M. (2004). Nutrient limitation and
nutrient-driven shifts in plant species composition in a species-rich fen meadow.
Journal of Vegetation Science, 15, 389-396.
Winkler J.B. & Herbst M. (2004). Do plants of a semi-natural grassland community
benefit from long-term CO2 enrichment? Basic and Applied Ecology, 5, 131-143.
152
Appendix 3 - Search terms used.
153
Search terms
154
experiment
deposition
deposition
deposition
deposition
deposition
nitrogen
nitrogen
nitrogen
nitrogen
nitrogen
grass*
prairie
savannah
meadow
pasture
experiment
addition
addition
addition
addition
addition
nitrogen
nitrogen
nitrogen
nitrogen
nitrogen
grass*
prairie
savannah
meadow
pasture
experiment
enrichment
enrichment
enrichment
enrichment
enrichment
nitrogen
nitrogen
nitrogen
nitrogen
nitrogen
grass*
prairie
savannah
meadow
pasture
experiment
fertilisation
fertilisation
fertilisation
fertilisation
fertilisation
nitrogen
nitrogen
nitrogen
nitrogen
nitrogen
grass*
prairie
savannah
meadow
pasture
experiment
pollution
pollution
pollution
pollution
pollution
nitrogen
nitrogen
nitrogen
nitrogen
nitrogen
grass*
prairie
savannah
meadow
pasture
function
deposition
deposition
deposition
deposition
deposition
nitrogen
nitrogen
nitrogen
nitrogen
nitrogen
grass*
prairie
savannah
meadow
pasture
function
addition
addition
addition
addition
addition
nitrogen
nitrogen
nitrogen
nitrogen
nitrogen
grass*
prairie
savannah
meadow
pasture
function
enrichment
enrichment
enrichment
enrichment
enrichment
nitrogen
nitrogen
nitrogen
nitrogen
nitrogen
grass*
prairie
savannah
meadow
pasture
function
fertilisation
fertilisation
fertilisation
fertilisation
fertilisation
nitrogen
nitrogen
nitrogen
nitrogen
nitrogen
grass*
prairie
savannah
meadow
pasture
function
pollution
pollution
pollution
pollution
pollution
nitrogen
nitrogen
nitrogen
nitrogen
nitrogen
grass*
prairie
savannah
meadow
pasture
Search terms
155
experiment
deposition
deposition
deposition
deposition
deposition
carbon dioxide
carbon dioxide
carbon dioxide
carbon dioxide
carbon dioxide
grass*
prairie
savannah
meadow
pasture
experiment
addition
addition
addition
addition
addition
carbon dioxide
carbon dioxide
carbon dioxide
carbon dioxide
carbon dioxide
grass*
prairie
savannah
meadow
pasture
experiment
enrichment
enrichment
enrichment
enrichment
enrichment
carbon dioxide
carbon dioxide
carbon dioxide
carbon dioxide
carbon dioxide
grass*
prairie
savannah
meadow
pasture
experiment
fertilisation
fertilisation
fertilisation
fertilisation
fertilisation
carbon dioxide
carbon dioxide
carbon dioxide
carbon dioxide
carbon dioxide
grass*
prairie
savannah
meadow
pasture
experiment
pollution
pollution
pollution
pollution
pollution
carbon dioxide
carbon dioxide
carbon dioxide
carbon dioxide
carbon dioxide
grass*
prairie
savannah
meadow
pasture
function
deposition
deposition
deposition
deposition
deposition
carbon dioxide
carbon dioxide
carbon dioxide
carbon dioxide
carbon dioxide
grass*
prairie
savannah
meadow
pasture
function
addition
addition
addition
addition
addition
carbon dioxide
carbon dioxide
carbon dioxide
carbon dioxide
carbon dioxide
grass*
prairie
savannah
meadow
pasture
function
enrichment
enrichment
enrichment
enrichment
enrichment
carbon dioxide
carbon dioxide
carbon dioxide
carbon dioxide
carbon dioxide
grass*
prairie
savannah
meadow
pasture
function
fertilisation
fertilisation
fertilisation
fertilisation
fertilisation
carbon dioxide
carbon dioxide
carbon dioxide
carbon dioxide
carbon dioxide
grass*
prairie
savannah
meadow
pasture
function
pollution
pollution
pollution
pollution
pollution
carbon dioxide
carbon dioxide
carbon dioxide
carbon dioxide
carbon dioxide
grass*
prairie
savannah
meadow
pasture
156
N.B. As the original intention of the project was to analyze a wider range of ecosystem
157
functions broad search terms were employed. Articles cited in studies which matched our
158
criteria and were added to our dataset were also searched to see if additional data could
159
be obtained.
160