Table 1. Mire types of the pristine, drained and restored treatments prior to the start of restoration in 2003, in the nine study mires in Central Finland and Northern Karelia, and their approximate coordinates. N = number of sampling locations representing the given mire type (total = 162 sampling locations).
Region/mire Treatment Mire types1 (N) Coordinates
Central Finland
Kiemanneva Pristine IR (1), RiNR (3), TR (2) 63°23´N, 25°16´E
Drained muIR (3), TKg (3)
Restored muIR (3), muRaR(3)
Väljänneva Pristine LkN (2), RaR(1), SR (3) 63°19´N, 25°18´E
Drained muIR (1), muRaR (3), muTR (1), TKg (1)
Restored muIR (1), muKeR (3), muRaR (2)
Southern Kulhanvuori Pristine IR (1), SR (1), TR (4) 62°34´N, 24°57´E
Drained muIR (1), TKg (5)
Restored muIR (3), TKg (3)
Northern Kulhanvuori Pristine LkR (6) 62°35´N, 24°57´E
Drained muRaR (1), TKg (5)
Restored muRaR (1), TKg (5)
Northern Karelia
Ristisuo Pristine LkR (1), RaR (5) 62°56´N, 31°20´E
Drained muIR (3), muKgR (1), muRaR (2)
Restored muIR (4), muPsR (1), muRaR (1)
Juurikkasuo Pristine LkR (1), RaR (5) 62°56´N, 31°26´E
Drained muIR (2), muKgR (1), TKg (2), VT (1)
Restored muPsR (1), TKg (5)
Rapalahdensuo Pristine LkR (4), RaR (2) 62°54´N, 29°30´E
Drained muIR (2), muLkR (1), muRaR (1), TKg (2)
Restored muIR (4), TKg (2)
Tiaissuo Pristine LkR (3), RaR (3) 62°56´N, 29°24´E
Drained muIR (4), TKg (2)
Restored muIR (4), muRaR (2)
Heinäsuo Pristine LkN (1), RaR (5) 62°54´N, 31°28´E
Restored-a muIR (3), muLkR (2), muRaR (1)
Restored-b LkR (1), muIR (4), muLkR (1)
1 Mire type abbreviations are according to Eurola et al. (1995), and English translations are according to Raunio et al. (2008): IR = Dwarf shrub pine bogs, LkN = Low-sedge bogs & fens, LkR = Low-sedge pine fens, muIR = Transforming Dwarf shrub pine bogs, muKeR = Transforming Ridge-hollow pine bogs, muKgR = Transforming Thin-peated pine mires, muLkR = Transforming Low-sedge pine fens, muPsR = Transforming Carex globularis pine mires, muRaR = Transforming Sphagnum fuscum bogs, muTR = Transforming Eriophorum vaginatum pine bogs, RiNR = Flark pine fens, RaR = Sphagnum fuscum bogs, SR = Tall-sedge pine fens, TKg = Transformed drained mires, TR = Eriophorum vaginatum pine bogs, VT = Sub-xeric heath forests.
Table 2. Mean (x± SE) tree-stand and sapling characteristics, microsite-type coverage, mire-surface coverage and water-table levels of the sampling locations in pristine, drained and restored mires. Differences in variables among the treatments were tested with Kruskal-Wallis rank sum test (H, df = 2; treatments not sharing the same letter differed significantly according to a posteriori test with critical α = 0.05). We adjusted the original p-values (p adj. in the Table) to control false discovery rate in multiple testing using the method in Benjamini and Yekutieli (2001). h = height, d1.3 = diameter at breast height (1.3 m from the mire-surface level or 1.3 m from the butt end of logs), and N = number of sampling locations. Data on living and dead trees are from the 100 m2 tree-sampling plots, data on tree saplings and microsite types are from the 25 m2 sapling squares, data on mire-surface coverage are from the 1 m2 vegetation squares, and data on water-table depth are from the ground-water wells.
Variable Treatment Test statistics
Pristine (N = 54) Drained (N = 48) Restored (N = 60)
x SE   x SE   x SE   H p adj.
Living trees (h > 1.5 m):
Total number of stems 5.8 1.0 a 23.1 2.5 b 9.2 1.2 a 60.47 <0.0001
   No. of pines 5.8 1.0 a 14.9 1.3 b 8.0 1.2 a 42.03 <0.0001
   No. of birches 0.1 0.0 a 7.4 2.3 b 1.2 0.4 a 42.95 <0.0001
   No. of stems d1.3 < 7 cm 4.7 0.8 a 13.8 2.3 b 7.1 1.1 a 17.65 0.0007
   No. of stems d1.3 7–20 cm 1.0 0.3 a 8.2 0.8 b 1.8 0.3 a 67.34 <0.0001
   No. of stems d1.3 > 20 cm 0.0 0.0 a 1.1 0.2 b 0.3 0.1 a 32.44 <0.0001
   No. of stems h 1.5–3 m 4.1 0.7   3.5 0.8   3.5 0.8   2.05 1.0000
   No. of stems h 3–8 m 1.7 0.3 a 12.5 2.0 b 4.9 0.7 c 40.24 <0.0001
   No. of stems h > 8 m 0.0 0.0 a 7.0 1.2 b 0.8 0.4 a 68.29 <0.0001
Number of species 0.9 0.1 a 1.8 0.1 b 1.0 0.1 a 53.10 <0.0001
Dead trees:
Total number of snags 0.7 0.2   1.1 0.2   0.9 0.3   1.49 1.0000
   No. of snags d1.3 < 7 cm 0.6 0.1   0.9 0.2   0.8 0.3   0.96 1.0000
   No. of snags d1.3 7–20 cm 0.1 0.1   0.2 0.1   0.1 0.0   1.16 1.0000
Total number of logs 0.0 0.0 a 0.4 0.2 a 0.6 0.2 a 10.24 0.0354
   No. of logs d1.3 < 7 cm 0.0 0.0 a 0.4 0.2 a 0.6 0.2 a 11.15 0.0235
   No. of logs d1.3 7–20 cm 0.0 0.0   0.1 0.0   0.1 0.0   1.57 1.0000
   No. of logs d1.3 > 20 cm 0.0 0.0   0.0 0.0   0.0 0.0   1.70 1.0000
Tree saplings (h 50–150 cm):
Total number of saplings 1.9 0.3   2.6 0.6   1.7 0.3   1.02 1.0000
   No. of pines 1.9 0.3 a 1.4 0.4 b 1.2 0.2 ab 8.60 0.0736
   No. of birches 0.0 0.0 a 0.9 0.4 b 0.3 0.1 ab 17.14 0.0014
Number of species 0.7 0.1   0.9 0.1   0.8 0.1   1.31 1.0000
Microsite types (%):
   Hummock 46.1 3.6 a 96.6 2.0 b 66.9 4.1 c 71.47 <0.0001
   Lawn 37.8 4.0 a 1.1 0.7 b 24.8 3.8 c 58.68 <0.0001
   Flark 15.5 3.6 a 0.0 0.0 b 6.3 1.7 a 27.05 <0.0001
Mire-surface coverage (%):
   Water 0.0 0.0 a 0.0 0.0 a 2.4 1.0 a 14.20 0.0052
   Litter 2.4 1.2 a 12.1 3.1 ab 15.5 2.4 b 27.30 <0.0001
   Sphagnum spp. 90.0 1.7 a 30.7 3.8 b 46.0 3.9 b 83.39 <0.0001
   Other mosses 3.1 0.6 a 38.4 3.9 b 22.7 2.8 c 64.53 <0.0001
   Herbs, sedges and grasses 14.0 1.1 a 8.2 1.0 b 20.9 2.2 a 22.51 <0.0001
   Low dwarf shrubs 9.6 1.2 ab 16.6 2.1 a 8.6 1.3 b 8.92 0.0655
   Tall dwarf shrubs 3.7 0.6 a 6.2 0.8 ab 9.5 1.0 b 19.39 0.0007
Water-table depth (cm below the mire-surface) 15.1 1.3 a 38.0 1.5 b 16.0 1.3 a 74.34 <0.0001
1

Fig. 1. NMDS ordination plots of the tree-stand variables presenting (A) sampling locations of the pristine (white dots), drained (black dots) and restored (grey dots) mires and (B) tree and sapling variables within the sampling locations. The dispersion ellipses in plot A indicate 1 SD of the weighted average of the site scores of pristine (solid line), drained (dotted line) and restored (dashed line) mires. The arrows in plot A indicate the environmental variables fitted to the ordination space such that only variables with highly significant p-values are shown (p < 0.001; the direction of the arrow indicates the direction of the gradient, and the length of the arrow indicates the strength of the correlation). View larger in new window/tab.

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Fig. 2. NMDS ordination plots of the floristic data presenting (A) sampling locations of the pristine (white dots), drained (black dots) and restored (grey dots) mires and (B) moss, lichen and vascular plant species within the sampling locations (note the differences in the scales of axes between plots A and B). The dispersion ellipses in plot A indicate 1 SD of the weighted average of the site scores of pristine (solid line), drained (dotted line) and restored (dashed line) mires. The arrows in plot A indicate the environmental variables fitted to the ordination space such that only variables with highly significant p-values are shown (p < 0.001). The species names indicated in plot B are such that for overlapping labels, priority is given to the most abundant species and the rest are indicated with “+”. After this, 14 frequent species (occurring in > 9% of sampling locations) remained without labels. These species are located as follows: Dicranum polysetum ca. 0.7 units right from the origin, Vaccinium uliginosum, Chamaedaphne calyculata, Polytrichum strictum Menzies ex Brid., Vaccinium oxycoccos, Betula nana, Sphagnum magellanicum Brid. and Andromeda polifolia within ca. 0.3 units from the origin, Carex rostrata Stokes ca. 1.0 units left from the origin, Pinus sylvestris L., Calluna vulgaris, Mylia anomala (Hook.) Gray, Drosera rotundifolia and Sphagnum rubellum within ca. 0.5–1.0 units toward the ca. lower left corner from the origin. For species abbreviations, see Appendix 1 (abbreviations represent the first three letters of the genus name and the first three letters of the species name). View larger in new window/tab.

Table 3. Mean (x, range 0–1) and total (fr) occurrence rates of ant workers and queens, and the mean number (x̄ ± SE, standard error given in column fr) of mire ant species and all ant species in the sampling locations representing different mire types in pristine and drained mires (mire types with less than four sampling locations in pristine and drained mires, and all sampling locations in restored mires were omitted). In the table, mire types within both pristine and the transforming and transformed mires are ordered by increasing growing stock from left to right (mire types to the left tend to have lower growing stock than types to the right). Differences in the occurrence rate (fr) of ant species among the mire types were tested with log-likelihood ratio test (G2, df = 6), and differences in the numbers of species were tested with Kruskal-Wallis rank sum test (H, df = 6; a posteriori test was unable to locate differences among mire types at α = 0.05 for the variable “number of mire ant species”). We adjusted the original p-values (p adj. in the Table) to control false discovery rate in multiple testing using the method in Benjamini and Yekutieli (2001). N = number of sampling locations (total N = 89).
Species Mire type1 Test statistics
Pristine Transforming and transformed
LkR
(N = 15)		 RaR
(N = 21) 		SR
(N = 4)		 TR
(N = 6) 		muRaR
(N = 7) 		muIR
(N = 16)		 TKg
(N = 20)
x fr x fr x fr x fr x fr x fr x fr G2 p adj.
Ant workers
Formica picea 0.5 7 0.3 7 0.8 3 0.3 2 0.0 0 0.0 0 0.1 1 27.30 0.0010
Formica uralensis 0.1 2 0.0 0 0.0 0 0.2 1 0.3 2 0.2 3 0.2 3 8.66 0.7269
Myrmica scabrinodis 0.6 9 0.9 18 0.8 3 0.7 4 0.9 6 0.6 10 0.4 7 13.90 0.1592
Lasius platythorax 0.3 4 0.4 9 0.5 2 0.5 3 0.3 2 0.5 8 0.5 10 3.25 1.0000
Formica sanguinea 0.1 2 0.4 9 0.3 1 0.0 0 0.4 3 0.1 1 0.1 1 16.86 0.0579
Leptothorax acervorum 0.2 3 0.4 9 0.5 2 0.2 1 0.1 1 0.1 1 0.1 2 11.43 0.3491
Myrmica rubra 0.1 2 0.1 2 0.3 1 0.0 0 0.0 0 0.1 1 0.2 3 4.43 1.0000
Myrmica ruginodis 0.3 5 0.4 9 0.0 0 0.3 2 0.9 6 0.9 15 0.9 18 36.74 < 0.0001
Camponotus herculeanus 0.1 1 0.0 0 0.0 0 0.2 1 0.1 1 0.8 12 0.7 14 51.34 < 0.0001
Ant queens
Myrmica scabrinodis 0.2 3 0.3 7 1.0 4 0.5 3 0.1 1 0.3 4 0.0 0 25.74 0.0017
Myrmica ruginodis 0.1 2 0.0 0 0.0 0 0.0 0 0.3 2 0.6 9 0.5 9 29.93 < 0.0001
Mean number of species H p adj.
Mire ants 1.3 0.2 1.2 0.1 1.8 0.3 1.3 0.4 1.1 0.1 0.9 0.2 0.6 0.1 20.45 0.0158
All ants 2.4 0.3 3.0 0.3 3.3 0.5 2.5 0.8 3.3 0.4 3.5 0.3 3.6 0.3 8.82 0.7269
1Mire type abbreviations are according to Eurola et al. (1995), and English translations are according to Raunio et al. (2008): LkR = Low-sedge pine fens, RaR = Sphagnum fuscum bogs, SR = Tall-sedge pine fens, TR = Eriophorum vaginatum pine bogs, muRaR = Transforming Sphagnum fuscum bogs, muIR = Transforming Dwarf shrub pine bogs, TKg = Transformed drained mires.
Table 4. Mean (x, range 0–1) and total (fr) occurrence rates of ant workers and queens, and the mean number (x̄ ± SE, standard error given in the column fr) of mire ant species and all ant species in pristine, drained and restored mires. Differences in the occurrence rate (fr) of ant species among the treatments were tested with log-likelihood ratio test (G2, df = 2), and differences in the number of species were tested with Kruskal-Wallis rank sum test (H, df = 2; treatments not sharing the same letter differed significantly from each other according to a posteriori test with critical α = 0.05). We adjusted the original p-values (p adj. in the Table) to control false discovery rate in multiple testing using the method in Benjamini and Yekutieli (2001). N = number of sampling locations (total N = 162).
Species Treatment Test statistics
Pristine (N = 54) Drained (N = 48) Restored (N = 60)
x fr   x fr   x fr   G2 p adj.
Ant workers:
   Formica picea 0.44 24   0.02 1   0.02 1   48.63 < 0.0001
   Formica uralensis 0.07 4   0.19 9   0.08 5   3.76 0.6328
   Myrmica scabrinodis 0.74 40   0.52 25   0.67 40   5.49 0.2918
   Lasius platythorax 0.35 19   0.50 24   0.60 36   7.13 0.1431
   Formica sanguinea 0.22 12   0.13 6   0.23 14   2.44 1.0000
   Leptothorax acervorum 0.30 16   0.15 7   0.20 12   3.52 0.6513
   Myrmica rubra 0.09 5   0.13 6   0.10 6   0.30 1.0000
   Myrmica ruginodis 0.37 20   0.92 44   0.72 43   37.34 < 0.0001
   Camponotus herculeanus 0.06 3   0.67 32   0.30 18   47.23 < 0.0001
Ant queens:
   Myrmica scabrinodis 0.41 22   0.10 5   0.18 11   14.25 0.0061
   Myrmica ruginodis 0.04 2   0.42 20   0.28 17   24.99 < 0.0001
   Camponotus herculeanus 0.11 6   0.02 1   0.20 12   9.68 0.0450
Mean number of species: H p adj.
Mire ants 1.3 0.1 a 0.8 0.1 b 0.8 0.1 b 22.59 < 0.0001
All ants 2.7 0.2 a 3.6 0.2 b 3.1 0.1 ab 10.23 0.0390
3

Fig. 3. Statistical responses of individual ant species to mire treatment (A = mire specialist species, B = generalist species, C = forest species). * = statistically significant (p < 0.05) responses (see Table 5).

Table 5. Generalised linear mixed model results of ant mire specialists, generalists and forest species. Model estimates, standard errors (SE) and p-values of terms retained in the models are given. Treatment, Sphagnum cover, hummock cover and number of trees (> 3 m) were a priori chosen as important environmental variables for the ant occurrences and were retained in all final models. Other variables were subject to model selection and their values are only given when they remained in the final model. Statistically significant p-values (< 0.05) are in bold. The intercept represents prediction in the pristine mire treatment. Est. = Estimate. * = mire specialist species, ** = generalist species, *** = forest species, all according to our a priori evaluations. View in new window/tab.
4

Fig. 4. Statistical responses (model coefficients ± SE, see Table 5) of individual ant species to environmental variables: A = Sphagnum moss cover, B = number of tall trees (> 3 m), C = hummock cover. * = statistically significant (p < 0.05) responses. Species were listed a priori (based on expert opinion and the literature: Krogerus 1960; Vepsäläinen et al. 2000; Punttila and Kilpeläinen 2009) from the most mire-associated (mire specialist) species at the top of each plot to species with the strongest forest affinities (forest species) at the bottom of each plot; generalist species are located in-between.

5

Fig. 5. NMDS ordination plots of the ant data presenting (A) sampling locations of the pristine (white dots), drained (black dots) and restored (grey dots) mires and (B) ant species within the sampling locations. The dispersion ellipses in plot A indicate 1 SD of the weighted average of the site scores of pristine (solid line), drained (dotted line) and restored (dashed line) mires. The arrows in plot A indicate the environmental variables fitted to the ordination space such that only variables with highly significant p-values are shown (p < 0.001). For species abbreviations, see Appendix 2 (abbreviations represent the first three letters of the genus name and the first three letters of the species name). View larger in new window/tab.