ELE_1053_sm_appendixS1

advertisement
1
Appendix S1 Description of experiments and predictor characterization
2
3
Experimental methodology: Differences in experimental protocols are summarized in
4
Table S1. Further details on experiments and sites can be found in the literature for ARC
5
(Shaver et al. 1996; Gough et al. 2002a; Gough & Hobbie 2003), CDR (Tilman 1987;
6
Wedin & Tilman 1996), JRG (Shaw et al. 2002; Zavaleta et al. 2003), KBS (Huberty et
7
al. 1998), KNZ (Seastedt et al. 1991; Blair 1997; Turner et al. 1997; Collins et al.
8
1998b), NWT (Bowman et al. 1995; Theodose & Bowman 1997; Seastedt & Vaccaro
9
2001), SEV (Noy-Meir 1973; Kieft et al. 1998), and SGS (Lauenroth et al. 1978;
10
Milchunas & Lauenroth 1995).
11
12
Climatic, soil, and biogeochemical properties: We used data from either online sources
13
or the published literature to characterize climatic, soil, and biogeochemical properties.
14
Where possible, we used data that utilized similar methodologies at multiple sites over
15
the same time period. Site temperature was represented by the USDA Hardiness factor
16
(http://www.usna.usda.gov/Hardzone/ushzmap.html), which are categories (1-11) based
17
on the average annual minimum temperature from 1974-1986. Precipitation for all sites
18
was estimated by the annual rainfall averaged over a decade or more from individual
19
monitoring programs. Precipitation values were extracted from the site LTER websites
20
for the seven LTER sites (http://www.lternet.edu/), and from the literature for Jasper
21
Ridge (Dukes et al. 2005). Soil pH was available from the literature and unpublished
22
data sources for ARC (Giblin et al. 1991; Gough et al. 2002b; Hobbie et al. 2002), CDR
23
(Pastor et al. 1987), JRG (Bohannan, unpublished data), KBS (Johnson et al. 2003), KNZ
1
1
(Blair, unpublished data), NWT (Bowman 2001), SGS (Johnson et al. 2003), and SEV
2
(Johnson et al. 2003). All measurements were taken from surface soil layers relevant to
3
the local site (which varied in depth by site) using predominantly a 1:1 ratio of soil:water
4
to measure pH (18 of 23 communities). The remaining 5 sites (all ARC) used either a
5
1:10 ratio of soil:water (2 communities) or a 1:10 ratio of soil:0.01MCaCl2 (3
6
communities). Increases in soil:water ratios do not largely affect pH readings due to
7
internal buffering in the soil suspension in either acidic (from hydrolysis of Al or H) or
8
calcareous (from hydrolysis of basic cations) soils (Sparks 1996). Measurements of pH
9
using water versus 0.01M CaCl2 have been shown to be highly correlated (r > 0.9),
10
though these studies were with large sample sizes (Gascho et al. 1996). Data for net N
11
mineralization used more heterogeneous methods, and were assembled using unpublished
12
data and the literature for ARC (Giblin et al. 1991; Hobbie & Gough 2002), CDR (Wedin
13
& Tilman 1996), KBS (Robertson et al. 1997), KNZ (Blair, unpublished data), NWT
14
(Bowman 2001), SGS (Dodd et al. 2000), and SEV (Kieft et al. 1998). Mean values
15
were inputted for the two communities for which data was unavailable (JRG and one
16
community at ARC, dry heath tundra). Most studies used intact soil samples separated
17
from surrounding soil matrix (either with polyethylene bags or PVC sleeves), incubated
18
monthly or longer, and then extracted with KCl (17 of 23 communities). The remaining
19
4 sites used resin bags sampled monthly (2 communities) or laboratory incubations over
20
similar periods (2 communities). Studies have shown that N mineralization estimated
21
from intact samples may differ from estimates using other methodologies (Raison et al.
22
1987); unfortunately, this heterogeneity was out of our control. Soil bulk density was
23
measured similarly for all sites, with a large diameter core taken to minimize compaction
2
1
which was then dried to constant weight. These data were available from the literature
2
and unpublished data for ARC (Giblin et al. 1991; Hobbie & Gough 2002), CDR (Grigal
3
et al. 1974), JRG (Cleland, unpublished data), KBS (Johnson et al. 2003), KNZ (online
4
data, http://www.konza.ksu.edu/), NWT (Sherrod & Seastedt 2001), SGS (Dodd et al.
5
2000), and SEV (Kieft et al. 1998). We were unable to obtain information on the
6
methodology employed for CEC at every site and thus do not report these methods here.
7
8
Plant community properties: Community properties represent conditions in non N-
9
treated plots. Most community properties were calculated using the PDT-Net dataset,
10
including productivity, the proportional number of species that were rare, and the relative
11
abundance of different functional groups. Additional details of this dataset are available
12
in prior publications (Gough et al. 2000; Suding et al. 2005). Our estimate of species
13
richness (number per square meter) required adjustment for different methodologies
14
employed. When measurements were made on a 1 m2 basis or larger, actual field data
15
were used, corrected to 1 m2. When smaller measurements were made, we used nested
16
species area accumulation curves collected in the summer of 2003 to estimate the number
17
of species per square meter.
18
19
3
1
Table S1 Descriptions of experimental protocols. Experiments differed in the amount of
2
N annually added, form of N, time of year N was added, number of replicates, the
3
measure on which relative abundance was based (Response measure), sample area, and
4
the specific year(s) used in this analysis (from experiment initiation). Multiple values
5
refer respectively to the different experiments within a site (ordered by Experiments
6
column).
7
8
4
1
System and
location
Arctic tundra,
Toolik Lake,
AK (ARC)
Sand prairie
or old field,
Cedar Creek,
MN (CDR)
Annual
grassland,
Jasper Ridge,
CA (JRG)
Old field,
Kellogg
Biological
Station, MI
(KBS)
2
Tallgrass
prairie,
Konza
Prairie, KS
(KNZ)
Alpine dry
meadow
tundra, Niwot
Ridge, CO
(NWT)
Desert
grassland,
Sevilleta, NM
(SEV)
Upland
shortgrass
steppe, CO
(SGS)
Number of
Form of N Time added replicates
Response
measure
Sample
area (m2)
Yrs used
100
NH4-NO3
pellet
June
16, 16, 24,
32, 24
Cover (%)
1
14, 14, 5,
10, 10
95
NH4-NO3
pellet
May, June
6
Biomass
0.3
Ave 17-19*
70
Ca-NO3
pellet
Nov., Dec.
8
Cover
(# hits)
0.5
5
120
NH4-NO3
pellet
June
6
Biomass
1
Ave 10-12*
100
NH4-NO3
pellet
May-June
4-8
Cover (%)
1-10
5-13
100
Urea, NH4NO3 pellet
July
5, 4
Cover (hits)
1
10, 4
Desert grassland (1)
100
NH4-NO3
pellet
May, July
10
Biomass
(allometric
estimates)
4
10
Ungrazed, with and
without increased
water (2)
60
NH4-NO3
prilled
Early April 100 (2 true)
Cover (%)
0.1
4
Experiments
(# per site)
N added
(kg ha-1yr-1)
dry heath sag, moist
sag, moist nonacidic
and acidic tussock,
dry heath tundra (5)
native savanna, old
fields abandoned
from agriculture in
1957 or 1934 (3)
Coastal range
grassland, ungrazed
(1)
Old field abandoned
from agriculture
1989, untilled since
(1)
Upland: burned,
burned and watered.
Lowland: six
combinations of
burning, irrigation
and mowing (8) †
Dry and moist
meadow tundra (2)
3
* Species abundances were determined from three year averages in these studies because
4
the small area annually sampled produced high variability between years (the same area
5
was never sampled twice).
6
† For more detail on the eight experiments from Konza see prior publications (Seastedt et
7
al. 1991; Blair 1997; Turner et al. 1997; Collins et al. 1998a).
8
5
1
2
3
4
5
6
7
8
9
10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
Supplemental references
Blair J.M. (1997) Fire, N availability, and plant response in grasslands: A test of the
transient maxima hypothesis. Ecology, 78, 2359-2368
Bowman W.D., T. R. Seastedt (2001) Structure and Function of on Alpine Ecosystem:
Niwot Ridge, Colorado. Oxford University Press, Oxford.
Bowman W.D., Theodose T.A. & Fisk M.C. (1995) Physiological and production
responses of plant growth forms to increases in limiting resources in alpine
tundra: Implications for differential community response to environmental
change. Oecologia, 101, 217-227
Collins S.L., Knapp A.K., Briggs J.M., Blair J.M. & Steinauer E.M. (1998a) Modulation
of diversity by grazing and mowing in native tallgrass prairie. Science, 280, 745747
Collins S.L., Knapp A.K., Briggs J.M., Blair J.M. & Steinauer E.M. (1998b) Modulation
of diversity by grazing and mowing in native tallgrass prairie. Science, 280, 745747
Dodd M.B., Lauenroth W.K. & Burke I.C. (2000) Nitrogen availability through a coarsetextured soil profile in the shortgrass steppe. Soil Science Society Of America
Journal, 64, 391-398
Dukes J.S., Chiariello N.R., Cleland E.E., Moore L.A., Shaw M.R., Thayer S., Tobeck T.,
Mooney H.A. & Field C.B. (2005) Responses of grassland production to single
and multiple global environmental changes. Plos Biology, 3, 1829-1837
Gascho G.J., Parker M.B. & Gaines T.P. (1996) Reevaluation of suspension solutions for
soil pH. Communications In Soil Science And Plant Analysis, 27, 773-782
Giblin A.E., Nadelhoffer K.J., Shaver G.R., Laundre J.A. & McKerrow A.J. (1991)
Biogeochemical diversity along a riverside toposequence in arctic Alaska.
Ecological Monographs, 61, 415-435
Gough L. & Hobbie S.E. (2003) Responses of moist non-acidic arctic tundra to altered
environment: productivity, biomass, and species richness. Oikos, 102, 204-216
Gough L., Osenberg C.W., Gross K.L. & Collins S.L. (2000) Fertilization effects on
species density and primary productivity in herbaceous plant communities. Oikos,
89, 428-439
Gough L., Wookey P.A. & Shaver G.R. (2002a) Dry heath arctic tundra responses to
long-term nutrient and light manipulation. Arctic, Antarctic and Alpine Research,
34, 211-218
Gough L., Wookey P.A. & Shaver G.R. (2002b) Dry heath arctic tundra responses to
long-term nutrient and light manipulation. Arctic Antarctic And Alpine Research,
34, 211-218
Grigal D.F., Chamberlain L.M., Finney H.R., Wroblewski D.W. & Gross E.R. (1974)
Soils of the Cedar Creek Natural History Area. In. University of Minnesota
Agriculture Experiment Station, St. Paul
Hobbie S.E. & Gough L. (2002) Foliar and soil nutrients in tundra on glacial landscapes
of contrasting ages in northern Alaska. Oecologia, 131, 453-462
Hobbie S.E., Miley T.A. & Weiss M.S. (2002) Carbon and nitrogen cycling in soils from
acidic and nonacidic tundra with different glacial histories in Northern Alaska.
Ecosystems, 5, 761-774
6
1
2
3
4
5
6
7
8
9
10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
Huberty L.E., Gross K.L. & Miller C.J. (1998) Effects of nitrogen addition on
successional dynamics and species diversity in Michigan old-fields. Journal of
Ecology, 86, 794-803
Johnson N.C., Rowland D.L., Corkidi L., Egerton-Warburton L.M. & Allen E.B. (2003)
Nitrogen enrichment alters mycorrhizal allocation at five mesic to semiarid
grasslands. Ecology, 84, 1895-1908
Kieft T.L., White C.S., Loftin S.R., Aguilar R., Craig J.A. & Skaar D.A. (1998)
Temporal dynamics in soil carbon and nitrogen resources at a grassland-shrubland
ecotone. Ecology, 79, 671-683
Lauenroth W.K., Dodd J.L. & Simms P.L. (1978) The effects of water- and nitrogeninduced stresses on plant community structure in a semiarid grassland. Oecologia,
36, 211-222
Milchunas D.G. & Lauenroth W.K. (1995) Inertia in plant community structure -- state
changes after cessation of nutrient enrichment stress. Ecological Applications, 5,
452-458
Noy-Meir I. (1973) Desert ecosystems: environments and producers. Annual Review of
Ecology and Systematics, 4, 25-51
Pastor J., Stillwell M.A. & Tilman D. (1987) Nitrogen mineralization and nitrification in
four Minnnesota old fields. Oecologia, 71, 481-485
Raison R.J., Connell M.J. & Khanna P.K. (1987) Methodology for studying fluxes of soil
mineral-N in situ. Soil Biology and Biochemistry, 19, 521-530
Robertson G.P., Klingensmith K.M., Klug M.J., Paul E.A., Crum J.R. & Ellis B.G.
(1997) Soil resources, microbial activity, and primary production across an
agricultural ecosystem. Ecological Applications, 7, 158-170
Seastedt T.R., Briggs J.M. & Gibson D.J. (1991) Controls of nitrogen limitation in
tallgrass prairie. Oecologia, 87, 72-79
Seastedt T.R. & Vaccaro L. (2001) Plant species richness, productivity, and nitrogen and
phosphorus limitatino across a snowpack gradient in Alpine Tundra, Colorado,
USA. Arctic and Alpine Research, 33, 100-106
Shaver G.R., Laundre J.A., Giblin A.E. & Nadelhoffer K.J. (1996) Changes in live plant
biomass, primary production, and species composition along a riverside
toposequence in Arctic Alaska, U.S.A. Arctic and Alpine Research, 28, 363-379
Shaw M.R., Zavaleta E.S., Chiariello N., Cleland E.E., Mooney H.A. & Field C.B.
(2002) Grassland responses to global environmental changes suppressed by
elevated by CO2. Science, 298, 1987-1990
Sherrod S.K. & Seastedt T.R. (2001) Effects of the northern pocket gopher (Thomomys
talpoides) on alpine soil characteristics, Niwot Ridge, CO. Biogeochemistry, 55,
195-218
Sparks D.L. (1996) Methods of Soil Analysis: Part 3-Chemical Methods. Soil Science
Society of America, Madison.
Suding K.N., Collins S.L., Gough L., Clark C., Cleland E.E., Gross K.L., Milchunas D.G.
& Pennings S. (2005) Functional- and abundance-based mechanisms explain
diversity loss due to N fertilization. Proceedings Of The National Academy Of
Sciences Of The United States Of America, 102, 4387-4392
Theodose T.A. & Bowman W.D. (1997) Nutrient availability, plant abundance, and
species diversity in two alpine tundra communities. Ecology, 78, 1861-1872
7
1
2
3
4
5
6
7
8
9
10
11
Tilman D. (1987) Secondary succession and the pattern of plant dominance along
experimental nitrogen gradients. Ecological Monographs, 57, 189-214
Turner C.L., Blair J.M., Scartz R.J. & Neel J.C. (1997) Soil N and plant responses to fire,
topography, and supplemental N in tallgrass prairie. Ecology, 78, 1832-1843
Wedin D.A. & Tilman D. (1996) Influence of nitrogen loading and species composition
on the carbon balance of grasslands. Science, 274, 1720-1723
Zavaleta E.S., Shaw M.R., Chiariello N., Mooney H.A. & Field C.B. (2003) Addititve
effects of simulated changes, elevated CO2, and nitrogen deposition on grassland
diversity. PNAS, 100, 7650-7654
8
Download