Abstract
The effects of black-locust invasion on plant forest diversity are still poorly investigated. Vascular plants are likely to be influenced by increasing nutrient availability associated with the nitrogen-fixing activity of black-locust, whereas it is not clear if, along with stand aging, black-locust formations regain forest species. The main aim of the present study was to test whether the increase of black-locust stand age promoted a plant variation in mature stands leading to assemblages similar to those of native forests. Therefore, plant richness and composition of stands dominated by native trees were compared with pure black-locust stands of different successional stages. Our study confirmed that the replacement of native forests by pure black-locust stands causes both plant richness loss and shifts in species composition. In black-locust stands plant communities are dominated by nitrophilous species and lack many of the oligothrophic and acidophilus species typical of native forests. Plant communities of native forests are more diverse with respect to pure black-locust stands, suggesting that black-locust invasion also causes a homogenization of the plant forest biota. We did not detect differences across the successional gradient of black-locust stands, and mature stands do not recover the diversity of plant species which are lost by the replacement of the native forests by black-locust. Accordingly some efforts in reducing the negative impacts of black-locust invasion on plant forest biota should be focused at least in those areas where conservation is among management priorities, such in the case of habitats included in the Habitat Directive (92/43 ECE).
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Acknowledgments
This work was partly conducted within the project QuiT (Inquinamento biologico in Toscana ed elaborazione di scenari futuri in relazione al cambiamento climatico) POR-FSE 2007–2013, Resp. Francesca Gherardi. Finally, we thank the Province of Pistoia for financial support.
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Appendix 1
Appendix 1
Species are listed in alphabetical order. Species frequency is expressed as the percentage of plots for each stand type in which the species occurred
Species | Frequency (%) | |||
---|---|---|---|---|
BL_Y | BL_A | BL_M | Native | |
Abies alba | 0 | 0 | 0 | 14 |
Acer campestre | 0 | 14 | 14 | 57 |
Acer pseudoplatanus a | 14 | 14 | 14 | 29 |
Agrostis capillaris bc | 14 | 29 | 0 | 0 |
Agrostis stolonifera | 0 | 0 | 14 | 0 |
Aira caryophyllea, subsp. caryophyllea | 0 | 0 | 0 | 14 |
Ajuga reptans | 0 | 0 | 0 | 29 |
Alliaria petiolata a | 29 | 14 | 14 | 29 |
Anemonoides nemorosa | 0 | 0 | 0 | 43 |
Anthoxanthum odoratum, subsp. odoratum | 0 | 0 | 0 | 14 |
Arabis sagittata b | 0 | 14 | 0 | 0 |
Arum italicum, subsp. italicum | 0 | 0 | 29 | 0 |
Asplenium adiantum-nigrum, subsp. adiantum-nigrum bc | 0 | 14 | 29 | 0 |
Brachypodium rupestre | 29 | 29 | 0 | 29 |
Brachypodium sylvaticum, subsp. sylvaticum | 0 | 14 | 0 | 0 |
Briza maxima b | 0 | 14 | 0 | 0 |
Bromus diandrus, subsp. maximus | 0 | 14 | 0 | 0 |
Bromus sterilis | 71 | 29 | 0 | 0 |
Calluna vulgaris | 0 | 0 | 0 | 14 |
Campanula rapunculus | 14 | 0 | 0 | 0 |
Campanula trachelium, subsp. trachelium | 0 | 0 | 0 | 14 |
Cardamine bulbifera | 0 | 0 | 0 | 14 |
Cardamine chelidonia | 0 | 14 | 0 | 0 |
Cardamine hirsuta | 0 | 0 | 29 | 0 |
Cardamine impatiens, subsp. impatiens a | 0 | 0 | 0 | 14 |
Carex sylvatica, subsp. sylvatica | 0 | 0 | 0 | 14 |
Castanea sativa | 14 | 29 | 71 | 86 |
Cephalanthera damasonium | 0 | 0 | 0 | 14 |
Chaerophyllum aureum a | 0 | 29 | 14 | 14 |
Chaerophyllum hirsutum a | 43 | 0 | 14 | 0 |
Clematis vitalba a | 43 | 43 | 14 | 0 |
Convolvulus althaeoides b | 0 | 0 | 14 | 0 |
Convolvulus arvensis | 14 | 14 | 0 | 0 |
Cornus mas | 14 | 0 | 0 | 14 |
Cornus sanguinea | 0 | 14 | 0 | 14 |
Corylus avellana a | 14 | 0 | 14 | 0 |
Crataegus monogyna b | 71 | 29 | 43 | 29 |
Crepis leontodontoides ca | 0 | 14 | 0 | 0 |
Crocus vernus | 0 | 0 | 0 | 29 |
Cruciata glabra | 0 | 0 | 0 | 29 |
Cytisus scoparius, subsp. scoparius bc | 0 | 0 | 0 | 57 |
Cytisus villosus | 0 | 0 | 0 | 14 |
Dactylis glomerata. | 0 | 14 | 0 | 0 |
Daphne laureola b | 0 | 0 | 0 | 29 |
Deschampsia flexuosa | 0 | 0 | 0 | 14 |
Digitalis lutea, subsp. australis | 0 | 0 | 0 | 57 |
Dryopteris filix | 0 | 0 | 0 | 14 |
Epipactis atrorubens | 0 | 0 | 0 | 14 |
Epipactis helleborine | 0 | 0 | 0 | 14 |
Erica arborea bc | 0 | 0 | 0 | 14 |
Erigeron sumatrensis a | 0 | 14 | 0 | 0 |
Euonymus europaeus | 14 | 0 | 0 | 0 |
Euphorbia amygdaloides, subsp. amygdaloides | 0 | 0 | 0 | 14 |
Euphorbia dulcis | 0 | 0 | 0 | 57 |
Fallopia dumetorum | 0 | 14 | 0 | 0 |
Festuca heterophylla | 29 | 29 | 0 | 57 |
Fragaria vesca, subsp. vesca | 0 | 14 | 0 | 0 |
Fraxinus ornus, subsp. ornus b | 43 | 29 | 57 | 43 |
Galium aparine | 14 | 57 | 14 | 14 |
Genista germanica | 0 | 0 | 0 | 14 |
Genista pilosa | 0 | 0 | 0 | 14 |
Genista tinctoria | 0 | 0 | 0 | 14 |
Geranium lucidum | 29 | 14 | 0 | 0 |
Geranium nodosum | 0 | 0 | 0 | 14 |
Geranium purpureum b | 14 | 29 | 14 | 14 |
Geranium robertianum | 0 | 14 | 0 | 14 |
Geranium rotundifolium b | 14 | 0 | 0 | 0 |
Geum urbanum a | 0 | 0 | 14 | 0 |
Hedera helix | 71 | 57 | 100 | 29 |
Helleborus bocconei, subsp. bocconei | 14 | 0 | 0 | 0 |
Hieracium murorum | 0 | 0 | 0 | 43 |
Holcus lanatus | 29 | 0 | 0 | 0 |
Holcus mollis bc | 0 | 0 | 0 | 14 |
Ilex aquifolium | 0 | 0 | 14 | 14 |
Juniperus communis | 0 | 0 | 0 | 14 |
Lactuca muralis | 14 | 0 | 43 | 14 |
Lamium album, subsp. album | 14 | 0 | 0 | 14 |
Lamium maculatum | 0 | 0 | 0 | 14 |
Lamium purpureum | 0 | 29 | 14 | 0 |
Lathyrus linifolius | 0 | 0 | 0 | 14 |
Laurus nobilis | 29 | 0 | 57 | 0 |
Lonicera caprifolium | 14 | 0 | 14 | 0 |
Lotus corniculatus | 0 | 0 | 0 | 14 |
Lunaria annua | 29 | 43 | 29 | 0 |
Luzula forsteri | 0 | 0 | 0 | 57 |
Luzula multiflora b | 0 | 14 | 0 | 0 |
Luzula pedemontana | 0 | 14 | 0 | 43 |
Medicago orbicularis | 0 | 14 | 0 | 0 |
Melica uniflora | 0 | 0 | 14 | 0 |
Melittis melissophyllum, subsp. melissophyllum b | 14 | 0 | 0 | 0 |
Moehringia trinervia a | 0 | 0 | 0 | 14 |
Molinia caerulea, subsp. arundinacea | 0 | 0 | 0 | 14 |
Muscari comosum | 0 | 29 | 0 | 0 |
Ornithogalum umbellatum | 0 | 14 | 0 | 0 |
Ostrya carpinifolia | 0 | 0 | 29 | 29 |
Physospermum cornubiense | 0 | 0 | 0 | 14 |
Phyteuma scorzonerifolium | 0 | 0 | 0 | 29 |
Pinus pinaster, subsp. pinaster | 0 | 0 | 0 | 14 |
Poa nemoralis, subsp. nemoralis b | 0 | 0 | 0 | 14 |
Poa sylvicola | 14 | 29 | 14 | 14 |
Poa trivialis a | 0 | 0 | 43 | 29 |
Polypodium interjectum c | 29 | 14 | 14 | 0 |
Polystichum aculeatum a | 0 | 29 | 14 | 0 |
Polystichum setiferum | 43 | 29 | 43 | 0 |
Potentilla micrantha | 14 | 0 | 0 | 14 |
Primula vulgaris, subsp. vulgaris | 0 | 0 | 0 | 14 |
Prunus avium, subsp. avium | 0 | 0 | 0 | 43 |
Prunus spinosa, subsp. spinosa | 0 | 0 | 0 | 14 |
Pteridium aquilinum, subsp. aquilinum bc | 0 | 0 | 0 | 71 |
Quercus cerris | 0 | 29 | 14 | 71 |
Quercus ilex | 14 | 0 | 0 | 0 |
Quercus pubescens, subsp. pubescens | 57 | 14 | 0 | 0 |
Ranunculus lanuginosus a | 0 | 14 | 29 | 0 |
Robinia pseudoacacia a | 100 | 100 | 100 | 14 |
Rorippa sylvestris, subsp. sylvestris | 0 | 0 | 0 | 14 |
Rosa canina | 0 | 0 | 0 | 14 |
Rubia peregrina | 29 | 29 | 29 | 0 |
Rubia peregrina, subsp. peregrina b | 0 | 0 | 0 | 14 |
Rubus caesius a | 0 | 14 | 0 | 0 |
Rubus hirtus a | 29 | 0 | 57 | 57 |
Rubus ulmifolius a | 43 | 100 | 57 | 0 |
Rumex crispus | 0 | 0 | 0 | 14 |
Rumex sanguineus a | 14 | 14 | 0 | 0 |
Ruscus aculeatus | 14 | 14 | 14 | 14 |
Sambucus nigra a | 57 | 43 | 86 | 0 |
Sanicula europaea | 0 | 0 | 0 | 29 |
Senecio ovatus a | 0 | 14 | 0 | 0 |
Serratula tinctoria, subsp. tinctoria | 0 | 0 | 0 | 14 |
Silene italica, subsp. italica | 0 | 14 | 0 | 14 |
Solidago virgaurea. | 0 | 0 | 0 | 43 |
Stachys officinali b | 0 | 0 | 0 | 14 |
Stellaria media, subsp. media | 14 | 71 | 57 | 29 |
Stellaria nemorum, subsp. montana a | 0 | 14 | 0 | 0 |
Tamus communis | 71 | 57 | 57 | 14 |
Teucrium scorodonia bc | 0 | 0 | 0 | 14 |
Torilis japonica a | 14 | 14 | 29 | 0 |
Trifolium repens | 0 | 0 | 0 | 14 |
Ulmus minor, subsp. minor | 0 | 0 | 0 | 14 |
Urtica dioica, subsp. dioica a | 14 | 29 | 0 | 0 |
Vaccinium myrtillus c | 0 | 0 | 0 | 43 |
Verbascum pulverulentum a | 0 | 14 | 0 | 0 |
Viburnum tinus, subsp. tinus b | 0 | 0 | 0 | 14 |
Viola alba, subsp. dehnhardtii | 14 | 29 | 43 | 0 |
Viola reichenbachiana | 14 | 0 | 14 | 14 |
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Benesperi, R., Giuliani, C., Zanetti, S. et al. Forest plant diversity is threatened by Robinia pseudoacacia (black-locust) invasion. Biodivers Conserv 21, 3555–3568 (2012). https://doi.org/10.1007/s10531-012-0380-5
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DOI: https://doi.org/10.1007/s10531-012-0380-5