Abstract
Changes in the structure of plant communities may have much more impact on ecosystem carbon (C) cycling than any phenotypic responses to environmental changes. We studied these impacts via the response of plant litter quality, at the level of species and community, to persistent water-level (WL) drawdown in peatlands. We studied three sites with different nutrient regimes, and water-level manipulations at two time scales. The parameters used to characterize litter quality included extractable substances, cellulose, holocellulose, composition of hemicellulose (neutral sugars, uronic acids), Klason lignin, CuO oxidation phenolic products, and concentrations of C and several nutrients. The litters formed four chemically distinct groups: non-graminoid foliar litters, graminoids, mosses and woody litters. Direct effects of WL drawdown on litter quality at the species level were overruled by indirect effects via changes in litter type composition. The pristine conditions were characterized by Sphagnum moss and graminoid litters. Short-term (years) responses of the litter inputs to WL drawdown were small. In long-term (decades), total litter inputs increased, due to increased tree litter inputs. Simultaneously, the litter type composition and its chemical quality at the community level greatly changed. The changes that we documented will strongly affect soil properties and C cycle of peatlands.
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References
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Acknowledgements
This study was supported by the Academy of Finland projects 104425 and 106197. We thank Satu Repo, Tuija Hytönen, Minna Piippo and Päivi Paasela for their valuable help with the laboratory work, Timo Penttilä, Nigel Roulet and the other reviewers for helpful comments on the manuscript, Meeri Pearson for correcting the language, and Jan Květ for pre-submission review of the manuscript.
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Appendices
Appendix 1
NPE | AE | EE | WE | cel | ara | rha | xyl | man | gal | glu | gl.ac | gal.ac | hol | P1 | P2 | P3 | V1 | V2 | V3 | S1 | S2 | S3 | C1 | C2 | Kl.lig | Sol.lig | Kl.lig:N | |
PN | 142.1 (3.0) | 28.36 (1.66) | 67.38 (3.86) | 88.26 (2.77) | 251.7 (13.9) | 22.46 (1.66) | 4.84 (0.46) | 13.76 (1.52) | 45.36 (3.07) | 25.69 (1.29) | 51.99 (3.62) | 1.21 (0.39) | 33.65 (2.88) | 445.8 (6.3) | 1.26 (0.03) | 0.38 (0.01) | 1.26 (0.03) | 13.26 (0.59) | 1.69 (0.07) | 3.23 (0.41) | 0 (0) | 0 (0) | 0 (0) | 4.35 (0.17) | 1.88 (0.04) | 227.1 (6.6) | 14.0 (0.4) | 53.6 (2.7) |
PC | 24.9 (5.0) | 7.04 (2.48) | 12.61 (1.90) | 27.34 (1.62) | 339.2 (16.7) | 11.97 (0.48) | 1.78 (0.23) | 18.32 (0.86) | 111.26 (5.62) | 26.44 (1.63) | 36.00 (1.49) | 0 (0) | 12.62 (0.72) | 566.6 (7.4) | 0.17 (0.02) | 0.05 (0.005) | 0.08 (0.01) | 20.89 (4.02) | 1.96 (0.35) | 2.78 (0.43) | 0.004 (0.001) | 0.019 (0.003) | 0.009 (0.002) | 0.08 (0.01) | 0.50 (0.06) | 345.7 (11.9) | 11.1 (0.5) | 109.1 (10.2) |
PB1 | 61.9 (7.0) | 9.27 (1.10) | 16.85 (2.03) | 24.82 (1.29) | 193.3 (20.7) | 19.21 (1.96) | 4.09 (0.24) | 38.21 (4.03) | 45.49 (2.50) | 44.94 (2.87) | 50.18 (4.58) | 3.90 (1.85) | 16.83 (1.34) | 511.6 (15.1) | 1.04 (0.09) | 0.25 (0.02) | 0.26 (0.02) | 44.39 (2.79) | 5.34 (0.96) | 5.65 (0.44) | 0.008 (0.001) | 0.051 (0.006) | 0.020 (0.006) | 0.12 (0.02) | 2.58 (0.34) | 374.4 (11.0) | 14.0 (1.3) | 65.9 (5.7) |
PB2 | 33.3 (2.8) | 9.17 (0.78) | 16.47 (2.26) | 23.26 (2.26) | 254.7 (42.2) | 15.45 (1.58) | 2.76 (0.41) | 49.84 (4.43) | 44.54 (4.52) | 48.62 (3.33) | 35.77 (3.42) | 1.22 (0.62) | 13.52 (2.03) | 581.5 (15.1) | 1.93 (0.09) | 0.36 (0.05) | 0.32 (0.04) | 51.95 (6.30) | 7.32 (1.35) | 9.32 (0.27) | 0.005 (0.003) | 0.034 (0.022) | 0.003 (0.002) | 0.08 (0.01) | 1.81 (0.13) | 373.6 (12.3) | 12.3 (0.7) | 129.0 (6.7) |
BN | 88.9 (2.2) | 68.76 (4.63) | 95.08 (5.07) | 116.93 (2.32) | 101.6 (15.3) | 21.89 (0.77) | 17.70 (0.55) | 36.26 (2.16) | 4.08 (1.33) | 28.15 (0.93) | 34.75 (1.31) | 0.96 (0.49) | 60.38 (2.38) | 258.3 (6.9) | 0.28 (0.02) | 0.77 (0.04) | 0.49 (0.02) | 5.16 (0.28) | 0.81 (0.04) | 3.30 (0.19) | 0.146 (0.010) | 0.331 (0.026) | 1.31 (0.12) | 0.93 (0.03) | 0.56 (0.03) | 327.0 (4.5) | 16.6 (1.3) | 38.5 (4.6) |
BNB | 34.8 (3.5) | 23.89 (3.51) | 18.43 (3.87) | 30.73 (1.10) | 103.6 (8.8) | 20.32 (1.65) | 4.77 (0.36) | 88.04 (11.22) | 6.87 (1.10) | 17.95 (1.26) | 30.10 (3.79) | 1.08 (0.47) | 25.44 (1.80) | 538.4 (6.0) | 0.20 (0.04) | 0.05 (0.003) | 0.09 (0.01) | 10.42 (2.06) | 1.24 (0.28) | 1.60 (0.28) | 2.312 (0.613) | 3.755 (0.372) | 25.6 (1.9) | 0.12 (0.01) | 1.55 (0.11) | 488.1 (18.9) | 19.1 (0.9) | 62.1 (1.5) |
BP | 94.6 (11.5) | 41.48 (5.08) | 122.66 (3.11) | 137.88 (9.41) | 67.7 (3.2) | 23.96 (3.92) | 16.53 (3.00) | 38.76 (6.18) | 3.32 (0.55) | 27.98 (3.61) | 30.96 (3.85) | 0.93 (0.93) | 71.37 (10.83) | 320.9 (22.8) | 0.67 (0.04) | 0.68 (0.16) | 0.74 (0.04) | 3.74 (0.20) | 0.61 (0.03) | 2.44 (0.09) | 0.300 (0.015) | 0.670 (0.015) | 3.43 (0.11) | 3.33 (0.69) | 0.66 (0.07) | 257.2 (2.7) | 10.9 (0.5) | 32.4 (3.3) |
BPB | 16.4 (2.9) | 13.97 (1.76) | 9.47 (0.77) | 15.63 (1.64) | 270.8 (33.8) | 4.62 (0.21) | 3.85 (0.17) | 149.47 (4.43) | 7.43 (0.85) | 13.44 (1.10) | 22.62 (2.00) | 2.25 (0.13) | 19.26 (1.14) | 754.6 (3.0) | 0.19 (0.03) | 0.04 (0.01) | 0.07 (0.01) | 16.07 (0.82) | 1.45 (0.10) | 2.97 (0.13) | 5.150 (0.685) | 7.623 (0.676) | 47.0 (3.3) | 0.10 (0.03) | 1.32 (0.24) | 349.6 (12.9) | 18.0 (0.9) | 112.3 (10.1) |
CR | 38.0 (1.9) | 35.51 (1.27) | 45.71 (2.01) | 80.17 (2.57) | 303.9 (27.1) | 38.46 (3.70) | 1.96 (0.04) | 146.73 (6.62) | 2.54 (0.17) | 15.45 (0.48) | 38.13 (1.62) | 16.41 (1.56) | 11.65 (0.56) | 650.1 (10.7) | 1.57 (0.03) | 0.29 (0.01) | 0.69 (0.07) | 4.43 (0.43) | 0.55 (0.04) | 0.91 (0.05) | 1.980 (0.115) | 7.018 (0.329) | 13.05 (0.48) | 21.26 (1.19) | 13.41 (0.61) | 234.8 (4.7) | 20.1 (3.0) | 30.7 (1.7) |
CL | 31.4 (6.0) | 26.88 (1.33) | 25.54 (2.99) | 58.38 (8.70) | 333.1 (9.9) | 46.22 (5.78) | 2.78 (0.18) | 163.69 (4.89) | 3.08 (0.18) | 20.01 (1.89) | 35.38 (2.81) | 20.10 (0.47) | 14.90 (0.83) | 622.8 (19.4) | 3.27 (0.65) | 0.56 (0.07) | 2.26 (0.73) | 8.74 (1.07) | 0.87 (0.12) | 1.09 (0.09) | 3.232 (0.678) | 9.504 (1.045) | 17.54 (2.42) | 32.85 (5.99) | 11.86 (1.06) | 287.3 (5.8) | 18.5 (1.1) | 37.9 (3.4) |
EV | 24.5 (0.8) | 35.63 (1.80) | 44.97 (1.81) | 49.25 (1.77) | 368.6 (40.8) | 49.19 (2.96) | 1.78 (0.46) | 154.16 (6.54) | 2.91 (0.40) | 17.45 (0.78) | 41.99 (2.92) | 6.07 (2.15) | 13.80 (1.01) | 643.2 (9.1) | 0.85 (0.03) | 0.26 (0.02) | 0.66 (0.03) | 4.41 (0.28) | 0.60 (0.04) | 1.35 (0.03) | 1.577 (0.057) | 4.109 (0.292) | 14.71 (0.44) | 14.28 (0.28) | 13.38 (0.56) | 278.7 (2.4) | 21.4 (1.0) | 45.8 (1.7) |
EVB | 9.4 (1.9) | 64.98 (2.91) | 34.20 (2.27) | 63.90 (5.75) | 319.8 (24.0) | 79.08 (6.42) | 2.78 (0.29) | 147.87 (12.99) | 5.18 (0.47) | 34.47 (2.64) | 29.59 (2.85) | 5.77 (1.50) | 11.25 (1.05) | 634.9 (10.2) | 1.34 (0.06) | 0.25 (0.02) | 0.32 (0.02) | 14.59 (1.04) | 1.81 (0.16) | 2.98 (0.12) | 1.156 (0.126) | 1.724 (0.091) | 10.04 (0.46) | 9.21 (0.51) | 16.32 (0.82) | 287.3 (4.4) | 14.7 (0.7) | 61.2 (5.3) |
RC | 63.5 (3.4) | 53.62 (1.22) | 72.51 (8.25) | 139.98 (4.99) | 85.2 (17.6) | 23.60 (2.82) | 10.70 (1.24) | 15.98 (2.14) | 4.65 (0.77) | 44.54 (4.17) | 47.00 (5.09) | 3.43 (1.16) | 47.03 (4.49) | 199.0 (5.6) | 0.14 (0.03) | 0.02 (0.01) | 0.20 (0.02) | 1.15 (0.25) | 0.23 (0.04) | 1.40 (0.16) | 0.135 (0.023) | 0.288 (0.038) | 0.883 (0.148) | 0.52 (0.05) | 0.67 (0.13) | 232.3 (16.8) | 83.7 (2.8) | 15.2 (1.5) |
VU | 61.8 (3.1) | 42.56 (2.75) | 120.08 (8.64) | 163.15 (9.46) | 106.9 (13.9) | 21.31 (1.31) | 4.15 (0.25) | 40.46 (3.10) | 2.65 (0.19) | 20.49 (0.86) | 29.59 (2.18) | 9.88 (1.27) | 95.28 (7.16) | 284.8 (6.6) | 0.21 (0.02) | 0.10 (0.01) | 0.82 (0.08) | 4.40 (0.40) | 0.86 (0.04) | 5.54 (0.50) | 0.180 (0.004) | 0.296 (0.011) | 1.222 (0.106) | 3.40 (0.16) | 1.13 (0.07) | 349.2 (16.2) | 18.9 (0.8) | 45.7 (4.9) |
LP | 122.9 (3.1) | 42.46 (2.89) | 48.04 (3.30) | 104.37 (5.49) | 61.2 (4.8) | 25.90 (0.91) | 7.29 (0.15) | 25.33 (1.21) | 4.64 (0.19) | 30.69 (0.97) | 44.00 (0.76) | 1.63 (0.68) | 49.42 (1.00) | 296.2 (5.8) | 0.11 (0.02) | 0.24 (0.02) | 0.83 (0.08) | 0.78 (0.16) | 0.31 (0.01) | 1.08 (0.30) | 0.173 (0.010) | 0.242 (0.011) | 0.656 (0.116) | 3.48 (0.29) | 1.98 (0.16) | 400.8 (11.6) | 18.5 (0.4) | 45.9 (0.7) |
PLS | 41.8 (9.4) | 18.70 (11.97) | 20.70 (1.87) | 58.88 (7.42) | 121.4 (15.2) | 17.92 (0.33) | 38.16 (1.78) | 20.21 (0.83) | 85.21 (3.74) | 68.95 (2.88) | 60.16 (4.63) | 1.37 (1.37) | 67.86 (3.19) | 436.4 (42.4) | 0.42 (0.05) | 15.33 (0.79) | 1.75 (0.13) | 0.45 (0.22) | 0.16 (0.02) | 0.12 (0.04) | 0.023 (0.003) | 0.018 (0.006) | 0.068 (0.008) | 0.52 (0.02) | 0.06 (0.02) | 306.2 (15.0) | 16.7 (3.0) | 39.7 (2.2) |
SA | 36.6 (7.3) | 19.92 (4.22) | 26.84 (3.57) | 54.47 (5.49) | 140.3 (10.9) | 10.84 (0.42) | 54.76 (2.36) | 44.08 (2.17) | 30.30 (1.63) | 81.26 (3.27) | 57.54 (2.45) | 21.63 (1.02) | 76.06 (2.47) | 699.3 (17.3) | 2.47 (0.22) | 10.20 (0.89) | 2.11 (0.16) | 0.23 (0.04) | 0.06 (0.01) | 0.05 (0.01) | 0.022 (0.005) | 0.022 (0.005) | 0.077 (0.023) | 0.33 (0.02) | 0.06 (0.02) | 106.4 (10.2) | 60.0 (4.0) | 14.3 (1.9) |
SFA | 40.6 (11.4) | 13.59 (2.66) | 22.25 (4.01) | 40.92 (3.74) | 222.1 (31.1) | 8.98 (0.98) | 48.95 (3.27) | 39.03 (4.82) | 26.00 (2.19) | 74.78 (5.67) | 51.49 (4.14) | 14.89 (2.23) | 67.06 (5.79) | 708.5 (8.8) | 2.84 (0.24) | 10.40 (1.18) | 2.21 (0.27) | 0.16 (0.04) | 0.04 (0.003) | 0.04 (0.01) | 0.030 (0.006) | 0.027 (0.005) | 0.090 (0.016) | 0.39 (0.04) | 0.05 (0.01) | 109.9 (4.4) | 58.7 (2.5) | 16.1 (2.1) |
SP | 62.6 (11.1) | 20.94 (5.39) | 33.42 (8.29) | 47.74 (9.55) | 219.0 (27.8) | 3.77 (1.03) | 31.58 (0.79) | 31.19 (1.45) | 24.40 (1.78) | 49.00 (1.74) | 55.81 (2.00) | 11.21 (2.89) | 47.23 (0.64) | 612.7 (34.3) | 3.31 (0.20) | 12.91 (0.67) | 4.24 (0.44) | 0.25 (0.02) | 0.11 (0.02) | 0.06 (0.01) | 0.091 (0.021) | 0.057 (0.007) | 0.232 (0.013) | 0.56 (0.03) | 0.14 (0.01) | 219.0 (21.8) | 52.3 (3.8) | 32.4 (6.4) |
SR | 82.5 (3.4) | 26.27 (6.04) | 22.94 (10.21) | 68.75 (8.85) | 230.3 (21.3) | 5.56 (3.09) | 39.96 (2.80) | 33.52 (3.84) | 18.76 (0.63) | 54.59 (4.86) | 54.78 (3.81) | 16.05 (2.79) | 59.07 (3.55) | 630.3 (20.9) | 2.57 (0.48) | 12.22 (2.03) | 2.52 (0.30) | 0.38 (0.13) | 0.10 (0.01) | 0.08 (0.02) | 0.032 (0.016) | 0.030 (0.009) | 0.110 (0.041) | 0.41 (0.04) | 0.06 (0.005) | 139.2 (21.0) | 58.2 (5.6) | 22.0 (3.7) |
SMG | 46.3 (7.5) | 29.46 (6.70) | 27.99 (6.62) | 62.22 (9.02) | 226.6 (23.9) | 6.13 (0.39) | 36.34 (1.94) | 32.90 (1.76) | 19.95 (1.19) | 46.46 (2.12) | 58.71 (2.81) | 26.78 (1.33) | 51.23 (2.89) | 662.3 (28.6) | 2.71 (0.32) | 9.41 (3.27) | 1.16 (1.11) | 0.80 (0.11) | 0.23 (0.05) | 3.27 (1.56) | 0.030 (0.005) | 0.030 (0.00001) | 0.121 (0.016) | 0.49 (0.02) | 0.10 (0.02) | 132.2 (20.6) | 67.0 (2.6) | 16.6 (2.8) |
SB | 35.1 (9.9) | 20.78 (4.33) | 33.87 (1.82) | 55.71 (19.39) | 204.3 (2.1) | 8.44 (2.76) | 43.00 (0.56) | 47.51 (0.15) | 33.53 (2.48) | 79.72 (2.86) | 56.47 (0.20) | 7.58 (3.20) | 58.29 (5.29) | 655.4 (33.5) | 2.38 (0.28) | 10.55 (1.05) | 1.96 (0.11) | 0.14 (0.03) | 0.04 (0.002) | 0.03 (0.01) | 0.014 (0.001) | 0.010 (0.003) | 0.047 (0.004) | 0.30 (0.03) | 0.06 (0.04) | 150.2 (31.9) | 53.9 (0.1) | 22.8 (4.6) |
SFC | 54.9 (6.1) | 25.80 (6.13) | 34.33 (3.19) | 76.58 (17.84) | 226.2 (16.8) | 4.15 (1.84) | 41.81 (0.54) | 37.26 (2.25) | 19.90 (0.85) | 51.43 (1.76) | 56.31 (0.82) | 28.60 (1.74) | 57.79 (0.08) | 556.2 (29.6) | 2.77 (0.27) | 10.01 (2.20) | 3.02 (0.29) | 0.27 (0.05) | 0.09 (0.01) | 0.11 (0.05) | 0.029 (0.008) | 0.029 (0.004) | 0.105 (0.028) | 0.47 (0.08) | 0.11 (0.03) | 234.0 (33.8) | 48.7 (1.6) | 34.7 (4.5) |
SCU | 17.3 (4.1) | 28.24 (0.15) | 17.15 (0.44) | 83.98 (1.40) | 293.7 (14.7) | 10.21 (2.13) | 39.22 (1.45) | 48.51 (2.13) | 35.55 (0.10) | 91.59 (9.98) | 74.26 (4.44) | 9.87 (0.29) | 52.87 (0.39) | 709.0 (5.9) | 3.03 (0.37) | 10.09 (1.51) | 2.50 (0.40) | 0.10 (0.02) | 0.04 (0.0003) | 0.03 (0.01) | 0.019 (0.003) | 0.015 (0.003) | 0.054 (0.015) | 0.34 (0.05) | 0.08 (0.01) | 81.8 (30.2) | 76.9 (2.3) | 11.4 (1.9) |
Appendix 2
Tree diagram from cluster analysis showing differences between litter types based on their chemical composition. Pinus sylvestris branches 1, branches with diameter ≤1 cm; Pinus sylvestris branches 2, branches with diameter 1–2 cm
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Straková, P., Anttila, J., Spetz, P. et al. Litter quality and its response to water level drawdown in boreal peatlands at plant species and community level. Plant Soil 335, 501–520 (2010). https://doi.org/10.1007/s11104-010-0447-6
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DOI: https://doi.org/10.1007/s11104-010-0447-6