Abstract
The relative strengths of the effects of forest edges and recreational use on understorey vegetation were studied at sub-xeric boreal urban forest edges in the greater Helsinki region, Finland. The study was performed at northern, eastern, southern and western edges, and vegetation sample plots were placed on, next to and away from paths with different trampling intensities 0–107 m from the forest edges. We found that human trampling altered vegetation more than the effects of forest edges. Vegetation changed dramatically on paths and the effects of path edges were seen in seemingly untrampled vegetation at least up to 4 m from the path edge. However, our results suggested that the effect of the edge may penetrate up to 50 m into forest interiors. Changes in vegetation composition indicated that the effects of the edge were stronger at eastern, southern and western than at northern edges.
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Acknowledgements
We thank Hannu Rita for his help in planning the sampling design, Minna Malminvaara-Lämsä for assisting in the field, Bob O’Hara for statistical advice and Sisko Salminen for her help in modifying the figures. The cities of Helsinki, Vantaa and Espoo, and The Finnish Forest Research Institute allowed us to work in their forests. The study was partly funded by the University of Helsinki, the city of Helsinki, the Maj and Tor Nessling Foundation, The Metsämiesten Säätiö Foundation and Helsinki University Centre for Environment (HENVI).
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Appendices
Appendix A
Average covers and occurrences (=occ, i.e. the percentages of sample plots in which the plant species were found) of vascular plants, bryophytes and lichens at northern, eastern, southern and western sub-xeric urban forest edges. Values at northern edges represent distances up to 18 m from the forest edge
| NORTH |
| EAST |
| SOUTH |
| WEST |
| |||
|---|---|---|---|---|---|---|---|---|---|---|---|
n = 30 |
| n = 74 |
| n = 79 |
| n = 73 |
| ||||
cover % | occ | cover % | occ | cover % | occ | cover % | occ | ||||
Vascular plants | Abbreviation | eda | tr | ||||||||
Acer platanoides | Acer plat | I | I | 0.01 | 7 | 0.00 | 0 | 0.00 | 0 | 0.00 | 0 |
Agrostis capillaris | Agro capi | T | T | 0.27 | 3 | 0.00 | 0 | <0.01 | 1 | 0.00 | 0 |
Anemone nemorosa | Anem nemo | T | I | 0.00 | 0 | 0.07 | 3 | 0.00 | 0 | 0.00 | 0 |
Arctostaphylos uva-ursi | Arct uvau | T | I | 0.60 | 3 | 0.00 | 0 | 0.00 | 0 | 4.03 | 12 |
Betula spp. | Betu spp | I | I | 0.07 | 7 | <0.01 | 1 | 0.00 | 0 | 0.04 | 4 |
Calamagrostis arundinacea | Cala arun | T | I | 0.00 | 0 | 0.84 | 12 | 0.06 | 4 | 0.05 | 1 |
Calamagrostis epigejos | Cala epig | T | I | 0.17 | 3 | 0.00 | 0 | 0.05 | 3 | 0.34 | 5 |
Calamagrostis spp. | Cala spp | 0.00 | 0 | 0.01 | 1 | 0.00 | 0 | 0.00 | 0 | ||
Calluna vulgaris | Call vulg | T | I | 1.81 | 17 | 1.31 | 23 | 4.28 | 19 | 2.61 | 19 |
Carex spp. | Care spp. | 0.00 | 0 | 0.02 | 4 | 0.00 | 0 | 0.03 | 3 | ||
Convallaria majalis | Conv maja | T | T | 0.00 | 0 | 0.93 | 5 | 0.46 | 8 | 0.53 | 10 |
Deschampsia flexuosa | Desc flex | T | I | 0.03 | 3 | 9.61 | 57 | 0.87 | 11 | 0.85 | 18 |
Epilobium angustifolium | Epil angu | T | I | 0.13 | 7 | 0.00 | 0 | 0.00 | 0 | 0.00 | 0 |
Festuca ovina | Fest ovin | T | T | 1.43 | 3 | 0.03 | 1 | 0.43 | 4 | 0.00 | 0 |
Goodyera repens | Good repe | S | S | 0.02 | 3 | 0.00 | 0 | 0.00 | 0 | 0.00 | 0 |
Hieracium vulgata coll. | Hier vulg | 0.20 | 3 | 0.00 | 0 | 0.00 | 0 | 0.00 | 0 | ||
Linnea borealis | Linn bore | S | S | 0.00 | 0 | 0.01 | 1 | 0.57 | 6 | 0.04 | 7 |
Luzula pilosa | Luzu pilo | T | I | 0.21 | 20 | 0.38 | 19 | 0.15 | 6 | 0.41 | 8 |
Lycopodium spp. | Lyco spp. | 0.00 | 0 | <0.01 | 1 | 0.00 | 0 | 0.00 | 0 | ||
Maianthemum bifolium | Maia bifo | S | I | 0.00 | 0 | 0.25 | 9 | 0.04 | 4 | 0.07 | 4 |
Melampyrum pratense | Mela prat | T | I | 3.87 | 43 | 1.00 | 18 | 2.16 | 37 | 0.89 | 23 |
Melampyrum sylvaticum | Mela sylv | T | I | 0.17 | 3 | 0.23 | 7 | 0.44 | 5 | 0.86 | 5 |
Melampyrum spp. | Mela spp. | T | I | 2.83 | 20 | <0.01 | 1 | 0.04 | 3 | 0.02 | 3 |
Picea abies | Pice abie | I | S | 0.00 | 0 | 0.00 | 0 | 0.18 | 3 | 0.00 | 0 |
Pinus sylvestris | Pinu sylv | T | S | 0.07 | 13 | 0.02 | 12 | 0.12 | 24 | 0.03 | 16 |
Poa spp. | Poa spp. | T | T | 0.01 | 7 | <0.01 | 1 | 0.15 | 8 | 0.03 | 1 |
Populus tremula | Popu trem | T | I | 0.00 | 0 | 0.04 | 1 | 0.18 | 5 | 0.00 | 0 |
Pteridium aquilinum | Pter aqui | T | I | 0.00 | 0 | 0.00 | 0 | 0.47 | 3 | 0.41 | 3 |
Rumex spp. | Rume spp. | T | T | 0.00 | 0 | 0.00 | 0 | 0.00 | 0 | 0.02 | 3 |
Seedlings | Seedl. | 0.02 | 17 | 0.01 | 4 | 0.01 | 6 | <0.01 | 1 | ||
Solidago virgaurea | Soli virg | T | S | 0.00 | 0 | 0.04 | 3 | 0.00 | 0 | 0.00 | 0 |
Sorbus aucuparia | Sorb aucu | I | I | 0.30 | 40 | 0.25 | 31 | 0.08 | 14 | 0.06 | 16 |
Trientalis europaea | Trie euro | S | I | 0.17 | 3 | 0.02 | 4 | 0.22 | 9 | 0.27 | 11 |
Vaccinium myrtillus | Vacc myrt | S | S | 25.62 | 73 | 20.59 | 66 | 18.42 | 56 | 14.87 | 48 |
Vaccinium vitis-idaea | Vacc viti | S | S | 7.07 | 63 | 15.11 | 87 | 12.27 | 82 | 15.25 | 92 |
Viola spp. | Viola spp. | 0.00 | 0 | 0.03 | 1 | 0.00 | 0 | 0.00 | 0 | ||
Bryophytes and lichens | |||||||||||
Aulacomnium palustre | Aula palu | I | I | 0.00 | 0 | 0.24 | 9 | 0.00 | 0 | 0.00 | 0 |
Brachythecium spp. | Brac spp. | T | S | 1.16 | 20 | 1.60 | 28 | 0.39 | 11 | 0.27 | 10 |
Ceratodon purpureus | Cera purp | T | T | 0.00 | 0 | 0.00 | 0 | 0.66 | 3 | 0.00 | 0 |
Certraria islandica | Cetr isla | T | T | 0.05 | 13 | 0.12 | 3 | 0.03 | 3 | 0.01 | 5 |
Cladina rangiferina | Cladina rang | T | S | 0.00 | 0 | 0.00 | 0 | 0.00 | 0 | 0.04 | 4 |
Cladina spp. | Cladina spp. | 0.00 | 0 | 0.00 | 0 | 0.03 | 3 | 0.00 | 0 | ||
Cladonia spp. | Clad spp. | I | S | 0.05 | 7 | 0.09 | 14 | 0.08 | 11 | 0.09 | 21 |
Dicranella spp. | Dicra spp. | 0.00 | 0 | 0.00 | 0 | 0.00 | 0 | 0.03 | 1 | ||
Dicranum majus | Dicr maju | S | S | 0.00 | 0 | <0.01 | 1 | 0.00 | 0 | 0.00 | 0 |
Dicranum polysetum | Dicr poly | S | S | 12.17 | 83 | 10.93 | 80 | 12.91 | 76 | 23.40 | 95 |
Dicranum scoparium | Dicr scop | S | I | 1.14 | 13 | 0.51 | 41 | 0.73 | 43 | 1.38 | 38 |
Dicranum spurium | Dicr spur | 0.00 | 0 | 0.00 | 0 | 0.03 | 1 | 0.00 | 0 | ||
Hepaticae | Hepa | 0.00 | 0 | 0.06 | 7 | 0.02 | 3 | 0.01 | 1 | ||
Hylocomium splendens | Hylo sple | S | S | 2.77 | 23 | 2.07 | 22 | 0.71 | 14 | 1.89 | 16 |
Plagiothecium spp. | Plag spp. | I | 0.00 | 0 | <0.01 | 4 | <0.01 | 1 | 0.01 | 1 | |
Pleurozium schreberi | Pleu schr | S | S | 21.02 | 93 | 13.25 | 84 | 11.52 | 65 | 11.91 | 88 |
Pohlia nutans | Pohl nuta | T | T | 0.27 | 17 | 0.27 | 14 | 0.15 | 15 | 0.12 | 21 |
Polytrichum commune | Poly comm | T | I | 0.00 | 0 | 0.16 | 5 | 0.00 | 0 | 0.84 | 5 |
Polytrichum juniperinum | Poly juni | T | T | <0.01 | 3 | <0.01 | 3 | 0.01 | 1 | 0.00 | 0 |
Ptilidium ciliare | Ptil cili | 0.00 | 0 | 0.06 | 3 | 0.00 | 0 | 0.05 | 4 | ||
Ptilium crista-castrensis | Ptil cris | S | S | 0.00 | 0 | 0.00 | 0 | 0.03 | 1 | 0.00 | 0 |
Rhytidiadelphus squarrosus | Rhyt squa | T | T | 0.00 | 0 | 0.43 | 5 | 0.00 | 0 | 0.00 | 0 |
Sanionia uncinata | Sani unci | 0.34 | 13 | <0.01 | 1 | 0.00 | 0 | 0.00 | 0 | ||
Sphagnum spp. | Spha spp. | 0.00 | 0 | 0.07 | 4 | 1.16 | 3 | 0.32 | 4 | ||
Appendix B
Generalized linear mixed model results for the plant species or species groups investigated (see Table 2). Sample plots on, next to and away from the paths are included in the models. Coefficients and p-values of the variables investigated are presented. One-tailed p-values are given for variables, because predictions from the literature were obtained for plant species responses to trampling and edge effects before the analyses were performed. However, for tree volume (m3 ha−1) and conifer percentage (%) two-tailed p-values are given. Statistically significant p-values are indicated in boldface
Model | Intercepta | tree volumeb | conifer%c | pathdistd | wearclass Le | wearclass Q | |||||||
|---|---|---|---|---|---|---|---|---|---|---|---|---|---|
Coeff. | p | Coeff. | p | Coeff. | p | Coeff. | p | Coeff. | p | Coeff. | p | ||
Vaccinium myrtillus | cover | −1.148 | 0.046 | −0.002 | 0.043 | −0.003 | 0.447 | 0.422 | <0.001 | −0.295 | 0.137 | −0.261 | 0.146 |
occur. | 0.647 | 0.527 | −0.004 | 0.017 | 0.003 | 0.760 | 0.250 | 0.023 | −1.573 | <0.001 | −0.460 | 0.139 | |
Pleurozium schreberi | cover | −2.182 | <0.001 | −0.001 | 0.353 | <0.001 | 0.952 | 0.330 | <0.001 | −0.731 | 0.003 | −0.251 | 0.153 |
occur. | 2.808 | 0.081 | −0.005 | 0.052 | 0.005 | 0.710 | 0.967 | <0.001 | −1.483 | 0.015 | 0.175 | 0.392 | |
Dicranum polysetum | cover | −3.658 | <0.001 | −0.001 | 0.441 | 0.016 | 0.009 | 0.315 | <0.001 | −0.203 | 0.148 | 0.274 | 0.079 |
occur. | 1.471 | 0.170 | <0.001 | 0.981 | 0.016 | 0.138 | 1.816 | <0.001 | −1.790 | <0.001 | −1.321 | 0.005 | |
Dicranum scoparium | cover | −6.833 | <0.001 | 0.002 | 0.107 | 0.001 | 0.824 | −0.350 | 0.005 | 0.218 | 0.251 | 0.111 | 0.364 |
occur. | −0.436 | 0.534 | 0.001 | 0.697 | 0.001 | 0.843 | −0.322 | 0.004 | 0.628 | 0.038 | −0.595 | 0.039 | |
Vaccinium vitis-idaea | cover | −1.655 | <0.001 | <−0.001 | 0.845 | −0.007 | 0.014 | 0.383 | <0.001 | −0.181 | 0.131 | 0.142 | 0.196 |
occur. | 1.635 | 0.588 | 0.002 | 0.717 | 0.001 | 0.972 | 2.187 | 0.052 | −1.533 | 0.154 | 0.031 | 0.491 | |
Melampyrum species | cover | −4.077 | <0.001 | −0.002 | 0.111 | −0.005 | 0.381 | 0.270 | 0.001 | −1.227 | 0.001 | −0.139 | 0.346 |
occur. | −0.674 | 0.451 | 0.002 | 0.381 | −0.009 | 0.289 | 0.242 | 0.024 | −1.455 | <0.001 | 0.424 | 0.159 | |
Grasses | cover | −1.911 | <0.001 | −0.002 | 0.204 | −0.005 | 0.300 | 0.246 | 0.005 | 0.146 | 0.321 | −0.435 | 0.073 |
occur. | 1.229 | 0.229 | −0.001 | 0.664 | −0.010 | 0.217 | 0.092 | 0.249 | −1.330 | 0.004 | −0.648 | 0.091 | |
Tolerant species | cover | −5.989 | <0.001 | −0.001 | 0.562 | <−0.001 | 0.964 | 0.159 | 0.077 | −0.631 | 0.110 | −1.054 | 0.007 |
occur. | −0.411 | 0.704 | <0.001 | 0.880 | 0.006 | 0.484 | −0.236 | 0.035 | 0.271 | 0.241 | −0.222 | 0.278 | |
Edge favouring | cover | −4.891 | <0.001 | 0.001 | 0.738 | −0.016 | 0.038 | −0.250 | 0.009 | −0.849 | 0.039 | 1.144 | 0.030 |
species | occur. | 0.596 | 0.430 | −0.002 | 0.243 | −0.016 | 0.019 | −0.084 | 0.245 | 0.320 | 0.182 | 0.620 | 0.044 |
Model | wearclass C | aspect S f | aspect Wg | edgelin h | edgesq | edgecub | |||||||
Coeff. | p | Coeff. | p | Coeff. | p | Coeff. | p | Coeff. | p | Coeff. | p | ||
Vaccinium myrtillus | cover | −0.109 | 0.305 | 0.348 | 0.280 | 0.608 | 0.169 | −0.034 | 0.046 | −0.001 | 0.097 | <0.001 | 0.021 |
occur. | −0.174 | 0.315 | 0.674 | 0.243 | 1.544 | 0.059 | 0.001 | 0.483 | 0.001 | 0.050 | – | – | |
Pleurozium schreberi | cover | 0.032 | 0.440 | 0.557 | 0.147 | −0.326 | 0.255 | −0.048 | 0.007 | −0.001 | 0.025 | <0.001 | 0.003 |
occur. | −0.420 | 0.220 | −1.787 | 0.053 | −0.410 | 0.400 | −0.081 | 0.081 | −0.002 | 0.018 | <0.001 | 0.025 | |
Dicranum polysetum | cover | −0.083 | 0.330 | −0.285 | 0.264 | 0.299 | 0.234 | 0.016 | 0.154 | <−0.001 | 0.244 | <0.001 | 0.405 |
occur. | 0.310 | 0.261 | −2.079 | 0.032 | 3.062 | 0.113 | 0.034 | 0.194 | −0.003 | 0.002 | <0.001 | 0.076 | |
Dicranum scoparium | cover | −0.656 | 0.014 | 1.324 | 0.116 | 1.101 | 0.159 | −0.116 | 0.050 | −0.001 | 0.275 | <0.001 | 0.149 |
occur. | −0.272 | 0.189 | 0.328 | 0.244 | 0.087 | 0.428 | −0.038 | 0.005 | – | – | – | – | |
Vaccinium vitis-idaea | cover | 0.345 | 0.015 | −0.007 | 0.489 | −0.079 | 0.375 | −0.004 | 0.228 | – | – | – | – |
occur. | 0.027 | 0.491 | −0.608 | 0.353 | −0.134 | 0.468 | 0.029 | 0.303 | – | – | – | – | |
Melampyrum species | cover | −0.479 | 0.035 | 0.857 | 0.155 | −0.783 | 0.196 | 0.063 | 0.020 | <−0.001 | 0.417 | <−0.001 | 0.053 |
occur. | −0.583 | 0.052 | 0.584 | 0.246 | −0.083 | 0.461 | 0.076 | 0.014 | −0.001 | 0.196 | <−0.001 | 0.053 | |
Grasses | cover | 0.348 | 0.111 | −2.205 | 0.027 | −1.384 | 0.007 | −0.030 | 0.080 | <−0.001 | 0.266 | <−0.001 | 0.243 |
occur. | 0.331 | 0.186 | 0.085 | 0.470 | −1.782 | 0.040 | 0.019 | 0.333 | 0.001 | 0.182 | <−0.001 | 0.059 | |
Tolerant species | cover | 0.005 | 0.494 | 1.076 | 0.186 | 0.484 | 0.353 | −0.153 | 0.039 | −0.001 | 0.288 | <0.001 | 0.181 |
occur. | −0.538 | 0.055 | −1.139 | 0.137 | −0.367 | 0.362 | −0.050 | 0.171 | −0.007 | 0.053 | <−0.001 | 0.084 | |
Edge favouring species | cover | 0.774 | 0.103 | 0.382 | 0.352 | −0.770 | 0.264 | −0.029 | 0.039 | – | – | – | – |
occur. | 0.088 | 0.403 | 0.171 | 0.406 | −0.010 | 0.494 | −0.008 | 0.277 | 0.001 | 0.010 | – | – | |
Model | aspect S: edgelini | aspect W: edgelinj | aspect S: edgesq | aspect W: edgesq | aspect S: edgecub | aspect W: edgecub | |||||||
Coeff. | p | Coeff. | p | Coeff. | p | Coeff. | p | Coeff. | p | Coeff. | p | ||
Vaccinium myrtillus | cover | 0.038 | 0.069 | 0.061 | 0.046 | <−0.001 | 0.330 | −0.002 | 0.014 | <−0.001 | 0.171 | <−0.001 | 0.093 |
occur. | 0.031 | 0.065 | 0.006 | 0.401 | −0.001 | 0.086 | −0.004 | <0.001 | – | – | – | – | |
Pleurozium schreberi | cover | 0.030 | 0.097 | 0.056 | 0.017 | −0.001 | 0.136 | <0.001 | 0.239 | <0.001 | 0.411 | <−0.001 | 0.039 |
occur. | 0.077 | 0.139 | 0.198 | 0.302 | 0.002 | 0.080 | 0.013 | 0.318 | <−0.001 | 0.141 | <0.001 | 0.359 | |
Dicranum polysetum | cover | 0.017 | 0.196 | 0.016 | 0.220 | 0.001 | 0.131 | <0.001 | 0.314 | <−0.001 | 0.046 | −0.001 | 0.409 |
occur. | −0.003 | 0.471 | −0.113 | 0.158 | 0.003 | 0.002 | <0.001 | 0.494 | <−0.001 | 0.034 | <0.001 | 0.290 | |
Dicranum scoparium | cover | 0.062 | 0.205 | 0.133 | 0.037 | <0.001 | 0.427 | 0.001 | 0.091 | <−0.001 | 0.385 | <−0.001 | 0.056 |
occur. | 0.016 | 0.173 | 0.032 | 0.041 | – | – | – | – | – | – | – | – | |
Vaccinium vitis-idaea | cover | 0.010 | 0.066 | 0.015 | 0.026 | – | – | – | – | – | – | – | – |
occur. | −0.020 | 0.382 | −0.040 | 0.293 | – | – | – | – | – | – | – | – | |
Melampyrum species | cover | −0.081 | 0.014 | −0.085 | 0.029 | <−0.001 | 0.243 | 0.001 | 0.120 | <0.001 | 0.063 | <0.001 | 0.115 |
occur. | −0.087 | 0.020 | −0.105 | 0.015 | 0.001 | 0.200 | 0.001 | 0.252 | <0.001 | 0.096 | <0.001 | 0.069 | |
Grasses | cover | −0.058 | 0.233 | −0.013 | 0.376 | −0.001 | 0.283 | −0.001 | 0.172 | <0.001 | 0.274 | <0.001 | 0.060 |
occur. | 0.003 | 0.484 | −0.091 | 0.048 | −0.003 | 0.035 | −0.002 | 0.120 | <0.001 | 0.433 | <0.001 | 0.012 | |
Tolerant species | cover | 0.176 | 0.025 | 0.056 | 0.284 | 0.001 | 0.258 | <0.001 | 0.437 | <−0.001 | 0.074 | <−0.001 | 0.478 |
occur. | 0.074 | 0.109 | 0.015 | 0.400 | 0.007 | 0.049 | 0.007 | 0.057 | <0.001 | 0.122 | <0.001 | 0.082 | |
Edge favouring species | cover | −0.007 | 0.395 | 0.019 | 0.220 | – | – | – | – | – | – | – | – |
occur. | −0.027 | 0.062 | −0.005 | 0.391 | <−0.001 | 0.265 | −0.001 | 0.187 | – | – | – | – | |
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Hamberg, L., Fedrowitz, K., Lehvävirta, S. et al. Vegetation changes at sub-xeric urban forest edges in Finland – the effects of edge aspect and trampling. Urban Ecosyst 13, 583–603 (2010). https://doi.org/10.1007/s11252-010-0142-7
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DOI: https://doi.org/10.1007/s11252-010-0142-7