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Fig. 1. Location of the study area in Poland and distribution of the relevés in Peucedano-Pinetum and Querco roboris-Pinetum habitats. View larger in new window/tab.

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Fig. 2. Section of the study area with the location of the relevés (AF – ancient forests; RF1, RF2 – recent forests; Table 1).

Table 1. Forest categories distinguished after cartographic analysis and field studies.
Forest category  
AF ancient forests defined as forests that have existed continuously since at least 1800; forest origin ascertained with the help of maps from 1800–1830; without a post-agricultural horizon in the soil
RF1 recent, post-agricultural forest regenerated over 200 years; forest origin ascertained with the help of maps from 1800–1830; with a distinct post-agricultural horizon in the soil
RF2 recent, post-agricultural forest regenerated in the last 80–180-year period; forest origin ascertained with the help of maps from 1885, 1928, 1950; with a post-agricultural horizon in the soil
Table 2. Variables included in the analyses (for species recorded in both forest types).
Trait Description Variable type Number of species with trait data available (max. = 50)
Max. height Maximum height of plant individual in cm continuous 49
Growth form Two classes: graminoids and herbs categorical 50
Life span Perennial/annual categorical 50
SLA Specific leaf area (mm2 mg–1) continuous 48
Seed longevity index Proportion (%) of short- and long-term persistent records on total (Bekker et al. 1998b) continuous 35
Seed weight Weight of 1000 dried seeds (g) continuous 39
Seed number Seed number per plant continuous 43
Dispersal type Two categories: long- and short-distance dispersal categorical 50
EA Ecological affinity: forest or open habitats categorical 50
Table 3. Percentage frequency of species and their indicator values computed using the phi coefficient for three pine forest categories: AF – ancient forests; RF1, RF2 – recent forests with different persistence; p – statistical significance; bold indicates max. values with the significance of the test between categories p ≤ 0.05.
Species name   AF RF1 RF2  p
No. of relevés 57 28 117
  frequency (%) phi value
herbs
Agrostis capillaris   8.8–0.025 3.6–0.124 12.00.082 0.476
Anthoxanthum odoratum   1.8–0.321 14.3–0.096 32.50.354 0.006
Athyrium filix-femina**   15.80.282 3.6–0.047 0.9–0.188 0.002
Calamagrostis arundinacea   38.60.244 17.9–0.065 16.2–0.166 0.010
Calamagrostis epigejos   7.00.030 10.70.102 4.3–0.074 0.351
Calluna vulgaris   98.20.452 78.60.145 47.9–0.493 <0.001
Chimaphila umbellata   5.3–0.256 3.6–0.251 32.50.378 <0.001
Convallaria majalis**   43.90.467 17.90.022 2.6–0.359 <0.001
Deschampsia flexuosa   77.2–0.145 78.6–0.097 90.60.159 0.076
Dryopteris carthusiana**   24.6–0.146 28.6–0.076 41.90.163 0.092
Festuca ovina   19.3–0.232 46.40.116 41.90.137 0.134
Hieracium lachenalii   3.5–0.100 0.0–0.175 11.10.185 0.066
Luzula pilosa**   42.10.251 25.0–0.003 17.1–0.199 0.007
Lycopodium annotinum*   10.5–0.046 10.7–0.036 14.5–0.057 0.664
Lycopodium clavatum   5.3–0.238 14.3–0.071 27.40.255 0.008
Melampyrum pratense**   96.50.135 96.40.117 86.3–0.189 0.127
Molinia caerulea   19.30.283 7.1–0.006 1.7–0.207 0.001
Monotropa hypopitys   1.8–0.169 10.70.034 12.00.133 0.168
Peucedanum oreoselinum   15.80.141 7.1–0.044 6.8–0.092 0.189
Pyrola chlorantha   0.0–0.199 3.6–0.094 12.80.250 0.005
Rumex acetosella   3.5–0.373 28.6–0.002 41.00.336 <0.001
Scorzonera humilis   29.80.349 0.0–0.225 6.0–0.183 <0.001
Solidago virgaurea*   10.50.003 3.6–0.131 12.00.060 0.733
Trientalis europaea*   33.30.342 7.1–0.118 6.8–0.216 <0.001
Vaccinium myrtillus**   100.00.097 96.4–0.058 97.4–0.052 0.294
Vaccinium uliginosum   14.00.313 0.0–0.126 0.0–0.183 <0.001
Vaccinium vitis-idaea**   100.00.331 92.90.179 68.4–0.423 <0.001
mosses and lichens
Cladonia arbuscula   24.6–0.228 46.40.054 48.70.165 0.079
Cladonia furcata   22.8–0.037 35.70.128 23.9–0.034 0.179
Cladonia rangiferina   22.80.033 14.3–0.090 21.40.016 0.901
Dicranum polysetum   98.20.011 100.00.088 97.4–0.052 0.723
Dicranum scoparium   38.60.112 28.6–0.025 27.4–0.080 0.222
Hylocomium splendens   91.20.300 71.40.008 60.7–0.268 0.001
Leucobryum glaucum   22.80.024 28.60.095 18.8–0.066 0.408
Pohlia nutans   17.50.352 0.0–0.141 0.0–0.206 <0.001
Polytrichum formosum   43.90.105 28.6–0.087 34.2–0.045 0.306
Polytrichum juniperinum   3.5–0.352 21.4–0.067 39.30.357 <0.001
Ptilium crista-castrensis   28.10.005 25.0–0.034 28.2–0.012 0.977
* – ancient forest species according to Dzwonko and Loster (2001)
** – according to Hermy et al. (1999) and Dzwonko and Loster (2001)
Table 4. Percentage frequency of species and their indicator values computed using the phi coefficient for three mixed oak-pine forest categories: AF – ancient forests; RF1, RF2 – recent forests with different persistence; p – statistical significance; bold indicates max. values with the significance of the test between categories p ≤ 0.05.
Species name   AF RF1 RF2  p
No. of relevés 30 13 51
  frequency (%) phi value
herbs
Agrostis capillaris   23.30.116 15.4–0.024 13.7–0.087 0.516
Anthoxanthum odoratum   10.0–0.301 15.4–0.178 45.10.376 0.003
Athyrium filix-femina**   33.30.225 30.80.138 9.8–0.248 0.120
Calamagrostis arundinacea   73.30.194 61.50.022 51.0–0.184 0.196
Calluna vulgaris   50.00.104 69.20.292 31.4–0.231 0.039
Carex digitata**   16.70.166 7.7–0.036 5.9–0.120 0.237
Chamaenerion angustifolium   16.70.135 7.7–0.054 7.8–0.089 0.380
Chimaphila umbellata   3.3–0.293 23.10.039 29.40.259 0.039
Convallaria majalis**   76.70.544 30.8–0.086 17.6–0.426 <0.001
Cytisus scoparius   3.3–0.254 7.7–0.143 27.50.328 0.014
Danthonia decumbens   10.0–0.014 7.7–0.054 11.80.038 1.000
Deschampsia flexuosa   83.3–0.013 76.9–0.102 86.30.061 0.805
Diphasiastrum complanatum   3.3–0.087 15.40.177 5.9–0.021 0.294
Dryopteris carthusiana**   70.00.059 46.2–0.223 68.60.058 0.764
Dryopteris filix-mas   13.30.088 0.0–0.200 9.80.008 0.713
Festuca ovina   26.70.072 23.10.010 19.6–0.066 0.826
Fragaria vesca   26.70.072 15.4–0.094 21.6–0.019 0.828
Galeopsis pubescens   10.00.194 7.70.086 0.0–0.182 0.277
Galium mollugo   10.0–0.146 7.7–0.156 11.10.216 0.108
Genista tinctoria   10.00.194 0.0–0.130 2.0–0.105 0.269
Hieracium lachenalii   10.0–0.077 7.7–0.102 17.60.121 0.604
Holcus mollis   0.0–0.213 0.0–0.188 15.70.332 0.021
Luzula pilosa**   90.00.314 76.90.094 54.9–0.342 0.008
Lycopodium annotinum*   26.7–0.047 23.1–0.082 33.30.081 0.759
Lycopodium clavatum   0.0–0.328 30.80.170 25.50.216 0.105
Maianthemum bifolium**   53.30.317 23.1–0.106 21.6–0.221 0.019
Melampyrum pratense**   83.3–0.135 100.00.212 90.20.027 0.230
Moehringia trinervia   13.30.012 0.0–0.234 15.70.095 0.585
Molinia caerulea   26.70.255 7.7–0.102 7.8–0.166 0.055
Mycelis muralis*   20.00.178 7.7–0.071 7.8–0.116 0.309
Orthilia secunda*   16.70.013 0.0–0.265 19.60.107 0.490
Oxalis acetosella**   53.30.198 38.5–0.010 31.4–0.166 0.154
Peucedanum oreoselinum   26.70.227 15.40.008 7.8–0.189 0.062
Poa augustifolia   10.00.067 0.0–0.175 7.80.016 0.856
Polygonatum odoratum*   33.30.384 7.7–0.092 3.9–0.042 0.003
Potentilla erecta   10.00.194 0.0–0.130 2.0–0.105 0.263
Pteridium aquilinum**   46.70.313 30.80.051 13.7–0.283 0.030
Rubus saxatilis   26.70.091 38.50.217 13.7–0.180 0.154
Rumex acetosella   13.3–0.194 7.7–0.238 37.30.301 0.024
Scorzonera humilis   20.00.282 7.7–0.016 2.0–0.212 0.029
Solidago virgaurea*   20.00.149 23.10.158 5.9–0.194 0.368
Trientalis europaea*   83.30.282 69.20.062 51.0–0.288 0.026
Vaccinium vitis-idaea**   86.70.150 100.00.323 66.7–0.299 0.028
Veronica officinalis   6.7–0.161 7.7–0.116 21.60.213 0.183
Viola reichenbachiana**   6.70.041 23.10.334 0.0–0.205 0.023
Viola riviniana*   16.70.201 7.7–0.016 3.9–0.154 0.191
mosses and lichens
Aulacomnium palustre   10.00.143 0.0–0.146 3.90.061 0.369
Cladonia arbuscula   0.0–0.213 15.40.125 11.80.131 0.345
Dicranum polysetum   83.30.110 69.2–0.093 74.50.052 0.455
Dicranum scoparium   26.70.035 23.1–0.018 23.5–0.022 0.946
Hylocomium splendens   90.0–0.037 84.6–0.125 94.10.091 0.679
Plagiomnium affine   23.30.116 7.7–0.143 15.7–0.036 0.507
Pleurozium schreberi   100.00.073 100.00.064 98.0–0.115 1.000
Pohlia nutans   10.00.037 23.10.247 3.9–0.154 0.084
Polytrichum commune   3.3–0.061 15.40.208 3.9–0.061 0.219
Polytrichum formosum   70.00.248 38.5–0.150 45.1–0.146 0.071
Polytrichum juniperinum   13.3–0.068 0.0–0.275 23.50.193 0.135
Ptilium crista-castrensis   40.0–0.080 38.5–0.080 51.00.109 0.460
Sciuro-hypnum oedipodium   23.30.320 7.7–0.036 2.0–0.233 0.026
* – ancient forest species according to Dzwonko and Loster (2001)
** – according to Hermy et al. (1999) and Dzwonko and Loster (2001)
Table 5. Categorical plant trait variables in pine and mixed oak-pine forests; p value from Fisher’s exact test; bold indicates the significant differences.
Trait Category Number of species (pine forest, n = 202) p value Number of species (mixed oak-pine forest, n = 84) p value
AFI non-AFI AFI non-AFI
Growth form graminoids 2 5 0.509 1 9 0.436
herbs 8 13 7 29
Life span perennial 10 17 0.643 8 35 0.556
annual 0 1 0 3
Dispersal type long-distance 3 18 0.037 3 30 0.031
short-distance 7 0 5 8
Ecological affinity forest 8 8 0.076 7 24 0.181
open habitat 2 10 1 14
Table 6. Plant trait variables of continuous type in pine and mixed oak-pine forests; p value from Mann-Whitney U-test; SD – standard deviation; bold indicates the significant differences.
Trait value Pine forest (n = 202) Mixed oak-pine forest (n = 84)
  AFI non-AFI p value AFI non-AFI p value
Maximum plant height (cm) Mean
(SD) 		61
(36.9) 		60
(43.8) 		0.801 62
(60.8) 		59
(35.1) 		0.222
Specific leaf area (mm2 mg–1) Mean
(SD) 		23
(15.4) 		29
(16.9) 		0.206 33
(12.1) 		27
(15.5) 		0.167
Seed bank longevity index (ratio) Mean
(SD) 		0.24
(0.27) 		0.22
(0.25) 		0.546 0.15
(0.20) 		0.25
(0.23) 		0.213
Seed weight (g) Mean
(SD) 		2.8
(5.86) 		1.01
(1.77) 		0.201 9.1
(10.64) 		1.4
(2.09) 		0.011
Seed number Mean
(SD) 		295.9
(410.1) 		1 955 502.6
(7 311 917.7) 		0.044 204
(408.3) 		1 113 042.2
(4 902 340.2) 		0.010