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The floodplain woods of Tuscany: towards a phytosociological synthesis
expand article infoMatilde Gennai, Antonio Gabellini§, Daniele Viciani, Roberto Venanzoni|, Lorella Dell’Olmo, Michele Giunti, Fabio Lucchesi, Francesco Monacci, Michele Mugnai, Bruno Foggi
‡ University of Florence, Florence, Italy
§ University of Siena, Siena, Italy
| University of Perugia, Perugia, Italy
¶ NEMO srl, Florence, Italy
Open Access

Abstract

The recent Landscape Plan of Tuscany Region pays particular attention towards the floodplain woods, for their conservation concern, landscape, and historical importance. The floodplain forest vegetation is defined as the natural and semi-natural wood vegetation located close to the rivers and submerged only during exceptional flooding. We gathered 180 both published and unpublished relevés coming from Tuscany and carried out in floodplain woods, where Alnus glutinosa, Fraxinus angustifolia subsp. oxycarpa, Quercus robur, Ulmus minor, Carpinus betulus, Populus nigra, P. alba and Salix alba were dominant, alone or in consociation with each other. On this dataset we performed a multivariate analysis, and the resulting groups were characterized by several points of view: floristic, ecological, chorological, by mean of diagnostic species, with the use of EIV. Moreover, their distribution in Tuscany was better defined. According to our results, six associations were found to be present in Tuscany. Thereby, one new association and three new subassociations were proposed. The syntaxonomic arrangement above the association level was discussed, with particular attention to the Italian Fraxinus angustifolia subsp. oxycarpa-rich communities. Finally, a comparison with the Annex I habitat types (sensu Directive 92/43/EEC) was carried out.

Keywords

Alnetea glutinosae, Alno glutinosae-Populetea nigrae, chorology, EIV, interdune woods, NMDS, Quercetalia pubescentis, syntaxonomy, swampy forests

Introduction

Floodplain forests are considered one of the most widespread forest communities in Europe (Schnitler et al. 2007) and represent an important biodiversity hotspot (Ward et al. 1999; Geilen et al. 2004). Unfortunately, they are severely endangered ecosystems, threatened by land-use changes. In fact, the shrinking of alluvial forests throughout Europe began in remote times and then strongly intensified in the last two centuries, due to agriculture and urbanization; this phenomenon strongly affected most areas of the floodplains, leading to the process of “insularization” and fragmentation of the remaining forests. Finally, the growing number of invasive alien species such as Robinia pseudoacacia L., Ailanthus altissima (L.) Swingle, Acer negundo L. and Amorpha fruticosa L. represents an additional risk today (Schnitler et al. 2007; Lazzaro et al. 2020; Viciani et al. 2020).

For these reasons, the recent landscape plan of the Tuscany Administrative Region identifies the conservation and management of "alluvial soils", as one of the main concerns for conservation (Marson 2016), in order to respect the Code of Cultural Heritage and Landscape, which was approved as a regional law in the 2015 by the Tuscany Regional Council within the new Landscape Plan. The plan integrates the three main components of the landscape: aesthetic-perceptive (aesthetic values), ecological (environmental values of the landscape) and structural features (relationships between cultural and natural aspects structured over time).

According to many authors (e.g. Ellenberg1988; Bernetti 2005; Douda et al. 2016), alluvial forests include the vegetation established at a specific distance from the rivers and submerged only during exceptional floods. The soil of such forests is generally deep, with the water level frequently reaching roots of plants. The vegetation that occurs in these ecological conditions can be defined as azonal (Géhu 2006), as it is linked to particular geomorphological and edaphic conditions present in many climatic types, rather than only in a specific climate (Bernetti 2005; Blasi 2010). Alluvial forests are widespread across the continent, covering large areas in central and northern Europe, while representing only relic communities in southern Europe and the Mediterranean. They are dominated by deciduous broadleaves: the black alder (Alnus glutinosa (L.) Gartner) in the marshy areas and the English oak (Quercus robur L.) in the typical alluvial plains; in our region we also found the elm (Ulmus minor Mill.) and southern ash (Fraxinus angustifolia Vahl subsp. oxycarpa (Willd.) Franco & Rocha Afonso). In suburban areas, where the influence of river dynamics is severely limited (e.g. dead meanders), poplar coenoses (Populus nigra L., P. alba L. and P. canescens (Aiton) Sm.), with elm, and relict stands of white willow (Salix alba L.) can be found. According to Biondi and Blasi (2015), Douda et al. (2016) and also recent Italian works (e.g. Spampinato et al. 2019), the European alluvial plain forests can be as attributed to two classes: Alnetea glutinosae Br.-Bl. and Tx. ex Westhoff et al., 1946, for the swampy woods, and Salici purpureae-Populetea nigrae Rivas-Martínez & Cantó ex Rivas-Martínez, Báscones, TE Díaz, Fernández-González et Loidi, 2001, for the periodically flooded riparian forests. On the contrary, Mucina et al. (2016) do not consider as an appropriate solution the use of the class Salici purpureae-Populetea nigrae and prefer to attribute part of the alluvial forests to Alno glutinosae-Populetea albae P. Fukarek et Fabijanić, 1968, because the concept of Salici purpureae-Populetea nigrae (as presented in the original diagnosis) also includes the Salicetalia purpureae order, that is the type of the Salicetea purpureae, i.e. a shrubby riparian class. We prefer to use Alno glutinosae-Populetea albae, in accordance with some recent Italian literature (e.g. Poldini et al., 2020), in order to include only riparian forest vegetation.

The syntaxonomic scheme below the class rank, especially at alliance level, is very intricate and a plethora of classification schemes have been proposed, often based on very few and local surveys.

With this study, we aim to contribute to the phytosociological knowledge of the alluvial forests of Tuscany, including the true swampy forests dominated by Alnus glutinosa and/or Fraxinus angustifolia subsp. oxycarpa, the suburban forests with Populus sp.pl., Salix alba and/or Ulmus minor, the hygrophilous and meso-hygrophilous woods with Quercus robur and the meso-hygrophilous forests dominated by Carpinus betulus.

Matherials and Methods

In order to define and identify in the field floodplain woods, we carried out several literature and field investigations which also led to the publication of an updated map for Tuscany (Gennai et al. 2020). The dataset was extracted from published and unpublished relevés stored in the database of the Laboratory of Phytogeography (Dept. of Biology, University of Florence). We firstly selected the relevés in which the tree species Alnus glutinosa, Fraxinus angustifolia subsp. oxycarpa, Quercus robur, Ulmus minor, Populus alba, P. nigra, P. cancescens, Salix alba and Carpinus betulus, alone or in association with each other, determined at least the 75% of the canopy cover. In Appendix I we provided references of literature used for the selected relevés. In Appendix II we reported the correspondence between numbers of relevés in original reference and numbers attributed in our tables. We excluded the strictly linear riparian formations. To the obtained data-set, 25 unpublished relevés, surveyed following the classical approach of the Zürich-Montpellier school and further updates (Braun-Blanquet 1964; Dengler et al. 2005, 2008; Biondi 2011), were added. These relevés were carried out in sites at a maximum altitude of 200 m a.s.l. and with maximum inclination of 2°, where the larger parts of the Tuscan floodplain woods are distributed (see Gennai et al. 2020). The final data-set comprised 180 relevés with a total of 425 species. We considered as sporadic the species occurring in less than four relevés and with cover-abundance values ≤ 1; such species were excluded from the numerical analyses, reducing the total number of species to 152. The Braun-Blanquet cover-abundance scale was transformed according to the ordinal scale proposed by Van der Maarel (1979) and Noest et al. (1989): r = 1, + = 2, 1 = 3, 2 = 5, 3 = 7, 4 = 8, 5 = 9.

The matrix was numerically classified by mean of Cluster Analysis using Ward’s method and by mean of NMDS analysis with the Bray-Curtis method as similarity measure; the analyses were performed using PAST software (Hammer et al. 2001).

The diagnostic species were statistically defined by the Phi coefficient of association (Chytrý et al. 2002) and its significance was calculated through a Fisher test. We considered a species as diagnostic of each group if phi > 0,20, with p < 0.01 (Douda et al. 2016).

A simplified high-level chorotype was attributed to each species, following Pignatti (2005).

The Ellenberg Indicators Values (Ellenberg et al. 1992) were attributed to species according to Pignatti (2005). We considered the following parameters: Light (L), Temperature (T), Continentality (C), Soil Moisture (U), Soil pH (R) and Nutrients (N). For each relevé, the total indicator value was calculated using the weighted averages of the presence/absence data of the species recorded in the relevé (except for sporadic species). To investigate possible different EIV characters of relevés and groups, the EIV vectors were passively projected onto NMDS ordination.

For the analysis of syntaxonomic data we referred to local and international literature separately cited in each paragraph. In particular, for the woods dominated by Fraxinus angustifolia subsp. oxycarpa we referred to the recent work by Poldini and Sburlino (2018), and we compared their outcomes with our results, through a NMDS analysis and a synoptic table.

The syntaxonomic scheme at higher ranks follows Biondi et al. (2014a, 2014b, 2015) and Biondi and Blasi (2015), with the exception of Salici purpureae-Populetea nigrae class, that was replaced by Alno glutinosae-Populetea albae in order to include only forest communities (Mucina et al. 2016; Poldini et al. 2020).

Vascular plant species names follow the Portal to the Flora of Italy (2020) and Pignatti (2017–2019).

In the descriptions of syntaxa, we indicated as diagnostic the characteristic and differential species reported by the original authors and the species highlighted by the phi analysis. In proper terms, the species indicated by the original authors as characteristics must be mainly considered as differentials (Géhu 2006).

In the syntaxonomic tables, we also indicated the diagnostic species as reported by the original authors of the syntaxa at the association level, and the phi values of the species (when phi> 20) resulting from the Phi analysis. The attribution of species to Alnetea, Phragmito-Magnocaricetea and Alno-Populetea follows Poldini and Sburlino (2018) and Poldini et al. (2020).

Results and Discussion

Both the cluster analysis dendrogram (Suppl. material 1: Figure S1) and NMDS results (Fig. 1) confirmed a clear separation between the relevés respectively dominated or co-dominated by Quercus robur and Carpinus betulus on one side, and Fraxinus angustifolia subsp. oxycarpa, Alnus glutinosa, Ulmus minor and Populus spp./Salix alba on the other side. The Cluster Analysis dendrogram (Suppl. material 1: Figure S1) shows that the first separation divides the mesohygrophylous woods dominated by Quercus robur from the other ones. As regards the NMDS ordination, he swampy forests with Alnus glutinosa (A) can be found in the first quadrant, while the linear woods of the interdune wet areas (locally named “lame”) with Fraxinus angustifolia subsp. oxycarpa (F) are placed in the second quadrant. The mesohygrophylous forests with Quercus robur (Q) and those dominated by Carpinus betulus (C) are in the third quadrant, while the thickets with Ulmus minor (U) and the woods with poplars and/or white willow (P) can be mainly found in the fourth quadrant.

Figure 1. 

NMDS ordination scatterplot of the relevés of the floodplain woods of Tuscany. A = Alnus glutinosa comm.; F = Fraxinus angustifolia subsp. oxycarpa comm.; P = Populus sp./Salix alba comm.; U = Ulmus minor comm.; C = Carpinus betulus comm.; Q = Quercus robur comm. The vectors resulting from EIV analysis are superimposed (L: Light, T: Temperature, C: Continentality, U: Soil Moisture, R: Soil pH, N: Nutrients).

The resulting EIV vectors projected onto NMDS ordination (Fig. 1) show that the woods dominated by Fraxinus angustifolia subsp. oxycarpa and Alnus glutinosa are positively influenced by soil moisture (U), light (L), continentality (C), and negatively influenced (but to a little extent), by temperature (T), while the woods dominated by Quercus robur and Carpinus betulus show an opposite trend. Nutrients (N) and soil pH (R) seem to affect the ordination to a lesser extent.

Statistical and floristic analyses allowed to recognize six different groups, interpreted as different vegetation types and analyzed through Phi analysis. Diagnostic species of these six groups, together with their frequencies, were reported in Table 1. Diagnostic species were indicated in light grey (phi > 20) and dark grey shading (phi > 30).

The results of the chorological analysis of the vegetation types are reported in Fig. 2.

The distribution in Tuscany of the six types of floodplain forests is shown in Fig. 3.

According to the results of our analysis and on the basis of the literature taken into account, we propose the following syntaxonomic scheme. In general terms, we confirmed the associations already reported for Tuscany, with the exception of some minor syntaxonomic formal changes, and the increase in the diagnostic species due to the increase in the number of relevés considered.

Table 1.

Results of Phi analysis. Synoptic table showing the fidelity coefficient of species for each community type. Vegetation types (A: Alnus glutinosa comm.; F: Fraxinus angustifolia subsp. oxycarpa comm.; Q: Quercus robur comm.; C: Carpinus betulus comm.; U: Ulmus minor comm.; P: Populus spp. and Salix alba comm.).

Vegetation types A F Q C U P
Alnus glutinosa (L.) Gaertn. 47.1 14.7 . . . .
Lycopus europaeus L. 38.8 . . . . .
Solanum dulcamara L. 38.7 . . . . .
Equisetum arvense L. 36.8 . . . . .
Thelypteris palustris Schott 35.7 . . . . .
Hydrocotyle vulgaris L. 35.6 . . . . .
Carex remota L. 33.5 . . . . .
Angelica sylvestris L. subsp. sylvestris 29.5 . . . . .
Urtica dioica L. subsp. dioica 28.2 . . . . .
Sambucus nigra L. 27.2 . . . . .
Carex pendula Huds. 26.1 . . . . .
Mentha aquatica L. subsp. aquatica 24.4 27.7 . . . .
Potentilla reptans L. 22.9 . . . . .
Fraxinus angustifolia Vahl subsp. oxycarpa (Willd.) Franco & Rocha Afonso . 54.5 . . . .
Carex elata All. subsp. elata . 46.3 . . . .
Limniris pseudacorus (L.) Fuss . 43.1 . . . .
Ficus carica L. . 31.7 . . . .
Galium palustre L. s.l. . 30.7 . . . .
Samolus valerandi L. . 28 . . . .
Lysimachia nummularia L. . 28 . . . .
Carex riparia Curtis . 27.9 . . . .
Juncus acutus L. subsp. acutus . 24.1 . . . .
Leucojum aestivum L. subsp. aestivum . 24.1 . . . .
Periploca graeca L. . 24 . . . .
Agrostis stolonifera L. . 23.4 . . . .
Juncus articulatus L. subsp. articulatus . 22.3 . . . .
Juncus inflexus L. subsp. inflexus . 18.8 . . . .
Carex otrubae Podp. . 14.4 . . . .
Quercus robur L. subsp. robur . . 54.9 . . .
Cornus mas L. . . 42.9 . . .
Lonicera caprifolium L. . . 40.4 . . .
Acer campestre L. . . 40.3 . . .
Luzula forsteri (Sm.) DC. . . 39.2 . . .
Ajuga reptans L. . . 36.1 . . .
Ligustrum vulgare L. . . 36.1 . . .
Chamaeiris foetidissima (L.) Medik. . . 35 . . .
Quercus ilex L. subsp. ilex . . 31.4 . . .
Crataegus monogyna Jacq. . . 29.6 . . .
Moehringia trinervia (L.) Clairv. . . 29.4 . . .
Carex sylvatica Huds. . . 29.3 . . .
Ruscus aculeatus L. . . 28.6 . . .
Cardamine pratensis L. . . 28.4 . . .
Arctium lappa L. . . 28.1 . . .
Cyclamen repandum Sm. subsp. repandum . . 28.1 . . .
Geum urbanum L. . . 27.5 . . .
Fraxinus ornus L. subsp. ornus . . 27 . . .
Hedera helix L. subsp. helix . . 26.8 . . .
Viola reichenbachiana Jord. ex Boreau . . 24.8 . . .
Myosotis sylvatica Hoffm. subsp. sylvatica . . 24.5 . . .
Holcus lanatus L. subsp. lanatus . . 23.9 . . .
Veronica montana L. . . 23.6 . . .
Brachypodium sylvaticum (Huds.) P.Beauv. subsp. sylvaticum . . 22.4 . . .
Ulmus minor Mill. subsp. minor . . 20.9 . . .
Clematis vitalba L. . . 19.9 . . .
Quercus cerris L. . . 19.7 . . .
Stellaria media (L.) Vill. subsp. media . . 19.4 . . .
Laurus nobilis L. . . 16.4 . . .
Euonymus europaeus L. . . 16 . . .
Pteridium aquilinum (L.) Kuhn subsp. aquilinum . . 14.2 . . .
Robinia pseudoacacia L. . . 7.2 . . .
Ilex aquifolium L. . . . 99.2 . .
Polygonatum multiflorum (L.) All. . . . 99.2 . .
Anemonoides nemorosa (L.) Holub . . . 97 . .
Corylus avellana L. . . . 94 . .
Lonicera etrusca Santi . . . 92.2 . .
Malus sylvestris (L.) Mill. . . . 91.7 . .
Carpinus betulus L. . . 4.9 87 . .
Asparagus tenuifolius Lam. . . . 78.1 . .
Festuca heterophylla Lam. . . . 77.1 . .
Mespilus germanica L. . . . 76.2 . .
Rubus hirtus Waldst. & Kit. group . . . 75.3 . .
Sorbus torminalis (L.) Crantz . . . 75.2 . .
Crataegus laevigata (Poir.) DC. . . . 73.5 . .
Dioscorea communis (L.) Caddick & Wilkin . . . 61.9 . .
Rhamnus alaternus L. subsp. alaternus . . . . 82.2 .
Smilax aspera L. . . . . 69.5 .
Rubia peregrina L. . . . . 68 .
Rosa sempervirens L. . . . . 66.3 .
Phillyrea angustifolia L. . . . . 59.2 .
Salix alba L. . . . . . 58.6
Populus nigra L. . . . . . 49.3
Rubus caesius L. . . . . . 44.5
Populus alba L. . . . . . 44.2
Xanthium italicum Moretti . . . . . 43.9
Humulus lupulus L. . . . . . 43
Salix cinerea L. . . . . . 42.2
Galega officinalis L. . . . . . 35.8
Pulicaria dysenterica (L.) Bernh. . . . . . 35.3
Cornus sanguinea L. . . . . . 26.4
Phragmites australis (Cav.) Trin. ex Steud. subsp. australis . . . . . 22.7

HYDROCOTYLO VULGARIS-ALNETUM GLUTINOSAE Gellini, Pedrotti et Venanzoni, 1986. Holotypus: Table 1, rel. 26 in Gellini et al. 1986 (corresponding to rel. n. 13 in Table 2 this paper) – Interdune swampy woods dominated by Alnus glutinosa (Table 2)

Table 2.

Hydrocotylo vulgaris-Alnetum glutinosae Pedrotti, Gellini & Venanzoni, 1986. Species reported by the original authors as diagnostic are written in italics.

*: typus relevé *
Rel. n. 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30
Phi Hydrocotylo vulgaris-Alnetum glutinosae
33.5 Carex remota L. + . . 3 . . . . + . + . 1 + + + + + . + 1 2 1 1 3 3 1 2 . 1
38.8 Lycopus europaeus L. . + + . . . 1 + + . . 1 + . + . . . . . + . + + + + + r . r
Periploca graeca L. 2 1 + . . . r + 1 + 1 1 + . . . + 1 . + + . + . . . . . . .
28.2 Urtica dioica L. subsp. dioica . . . 1 . . . . + + . . + + + . . + . + . + + + 1 2 1 2 .
26.1 Carex pendula Huds. . . . 3 1 + . 2 . . . . + . 1 . . + + . . 1 + + 1 . r . + .
22.9 Potentilla reptans L. . . . + . . . . + + . . + 1 . + + + + . 1 + + 1 . . . . .
35.6 Hydrocotyle vulgaris L. . . . . . . . . + + + . 1 . . + 1 1 1 1 . . + + . . . . . .
27.2 Sambucus nigra L. . . . . 1 1 . . . . . . . . 1 . . + . . . 1 . . . 1 r + + .
38.7 Solanum dulcamara L. . + . . . . 1 . . . . 1 + . . . . + . . . . . + . . . 1 . .
36.8 Equisetum arvense L. . . . 1 1 . . . . . . . . . . . . . . . . . . . . + 2 2 + +
29.5 Angelica sylvestris L. subsp. sylvestris . . . . . . . . . . . . . . . . . . . . . . . . . 3 2 1 r .
Alnion glutinosae, Alnetalia, Alnetea
47.1 Alnus glutinosa (L.) Gaertn. 5 5 5 5 5 5 5 5 5 5 5 5 5 5 5 4 4 4 3 3 5 5 5 4 5 5 5 5 5 5
Galium palustre L. s.l. + r . . . . r . + . + r + + . + . + + . + . + + + . . . . .
Ranunculus repens L. . . . 1 . . . . . . . . + + . . . + . + . . + . + 2 1 3 + +
35.7 Thelypteris palustris Schott . . . . . . . . + + . . 2 . . 1 1 1 . 5 1 . 3 3 . . . . . .
Samolus valerandi L. . . . . . . . . + + . . + . . + + . . + + . + + + . . . . .
Fraxinus angustifolia Vahl subsp. oxycarpa (M.Bieb. ex Willd.) Franco & Rocha Afonso . . . . . . . . . . + + + + . . . + . . 2 . 1 1 1 . . . . .
Ulmus minor Mill. subsp. minor 1 . . . . . . 2 . . . . + + . . . . . . + + + + . . . . 1 .
Frangula alnus Mill. subsp. alnus . 2 . . . . . + . . . 1 . . . . . . 1 . . . . . . r . + . +
Populus alba L. . . + . . . . . . . . . + . . + . 1 . . . . . . + . . . . 3
Scutellaria galericulata L. . . . . . . . . . . . . . . . . . 1 . . . . + + + . . . . .
Humulus lupulus L. . . . . 1 . . . . . . . . . 1 . . . . . . . . . . . + + . .
Rubus caesius L. . . . . . . 1 . . . . . . . . . . . . . . 1 . . . . . . . .
Osmunda regalis L. . . . . . . . . . . . . . . . . . . 3 . . . . . . . . . . .
Salix cinerea L. . . . . . . . . . . . . . . . . . . . . . . . . . . . r . .
Magnocaricion elatae, Magnocaricetalia elatae, Phragmiti-Magnocaricetea
24.4 Mentha aquatica L. subsp. aquatica + + . . . . . . 1 + + + 1 + . + + 2 . 1 1 . 1 2 1 . . . . .
Lythrum salicaria L. . . . . . . . . . . . + + . . + . + . . . . + . + . . . . +
Carex elata All. subsp. elata . . . . . . . . + . . . + . . . . . + . + . + . + . . . . .
Lysimachia vulgaris L. . . . . . . . . . . . . . . . . . . + . . . + . . . . . . +
Alisma plantago-aquatica L. . . . . . . . . + . . r . . . . . . . . . . . . . . . . . .
Limniris pseudacorus (L.) Fuss . . . + . + . . . . . . . . . . . . . . . . + . + . . . . .
Other species
Rubus ulmifolius Schott 1 + r + 3 5 3 1 + + 2 1 + 1 . + + 1 1 . + . + + 1 + 1 + . 3
Hedera helix L. subsp. helix + r . 2 4 3 . . + . + + . + 3 + . + + . + + + . . + . . . +
Juncus effusus L. subsp. effusus . . . . . . . . + . + . 1 + . + . + 1 . + . + + . . . . . +
Agrostis stolonifera L. . . . . . . . . + . . . + 1 . . . + + . . . + . . 1 r 3 + .
Crataegus monogyna Jacq. . . . . . . . . + . . . . + + . . + . . . . + + . + . . . .
Ficus carica L. . . . . 2 1 . + . . . . . + . 1 . + . . . . + . . . . . . .
Prunus spinosa L. subsp. spinosa . . . . . . . . . . . + . . + . . . . . . + . . . . 2 . + 1
Euonymus europaeus L. . . . . . . . . . . . . . . + . . . . . . . . . . + + + + +
Carex sylvatica Huds. . . . . . . . . + . . . + + . . . + . . . . + . + . . . . .
Cornus sanguinea L. s.l. . . . . . 1 . . . . . . . . . . . . . . . . . . + 1 1 + . 1
Arum italicum Mill. subsp. italicum . . . + . . . . . . . . . + . . . + . . . . . . . r . . r .
Brachypodium sylvaticum (Huds.) P.Beauv. subsp. sylvaticum . . . . . . . . . . . + . 1 . . . + . . . . . . . . . . + +
Quercus robur L. subsp. robur . . . . . . . . . . . . . . + . . . + . . 1 . . . r . . . +
Clematis vitalba L. . + . 1 . . . . + . . . . . + + . . . . . . . . . . . . . .
Rubia peregrina L. . r . . . . . . . . . + . . + . . . . . . + . . . . . . . .
Equisetum telmateia Ehrh. . . . . . . . . . . . . . . 1 1 + . . . . . . . 1 . . . . .
Poa trivialis L. . . . 1 . . . . . . . . + + . . . . . . . . . . + . . . . .
Hypericum androsaemum L. . . . . . . . . + . . . . + . . . + . . . . . . + . . . . .
Salix alba L. . . . . 1 . . + . . . . . . . . . . . . . 1 . . . . . . 2 .
Vitis vinifera L. s.l. . + . . . . . . . . . r . . . . . + . . . . . . + . . . . .
Laurus nobilis L. . . . + 1 . . . . . . . . . + . . . . . . 1 . . . . . . . .
Convolvulus sepium L. . . . 2 . . . . . . . . . . . . . + . . . . + . . . . . .
Bidens frondosa L. . . . . . . . . . . . r . . . . . . . . . . . . . . . . 1 r
Persicaria hydropiper (L.) Delarbre . . . . . . . . . + . . . . . . . . . 1 . + . . . . . . . .
Ruscus aculeatus L. . . . . . . . . . . . . . + + . . . . . . 1 . . . . . . . .
Smilax aspera L. . . r . . . + + . . . . . . . . . . . . . . . . . . . . .
Acer campestre L. . . . . . . . . . . . . . . . . . . . . . 1 . . . + . . . +
Carex canescens L. . . . . . . . . . . . . + . . . . + . . . . . . + . . . . .
Erigeron canadensis L. . . . . . . . . . . . . . + . . . + . . . . . . + . . . . .
Euphorbia peplus L. . . . . . . . . . . . . . . . + + . . + . . . . . . . . . .
Acer negundo L. . . . 1 . . . . . . . . . . . . . . . . . + . . . + . . . .
Populus nigra L. . . . . 1 . . . . . . 1 . . . . . + . . . . . . . . . . . .
Carex canescens L. . . . . . . . . x . . . . . . . . . . . . . x . x . . . . .
Pinus pinaster Aiton subsp. pinaster . . + . . . . 1 . . . . . . . . . . . . . . . . . . . . . .
Eupatorium cannabinum L. subsp. cannabinum . . . . . . . . . . . . . . . . . . . . . . . . . r . + . .
Lonicera caprifolium L. . . . . . . . . . . . . . . . . . + . . . + . . . . . . . .
Oxalis corniculata L. . . . . . . . . . . . . . + . . . + . . . . . . . . . . .
Persicaria dubia (Stein.) Fourr. . . . . . . . . . . . . . . . . . . . . . . . . . . . + 3 .
Parthenocissus quinquefolia (L.) Planch. . 2 . . . . . . . . . r . . . . . . . . . . . . . . . . . .
Carex riparia Curtis . . . 1 . . . . . . . . . . . . . . . . . . . . . . . . . .
Rubus hirtus Waldst. & Kit. group . . . . . . . . . . . . . . 4 . . . . . . . . . . . . . . .

Diagnostic species : Alnus glutinosa, Hydrocotyle vulgaris, Periploca graeca, Thelypteris palustris, Lycopus europaeus, Solanum dulcamara, Equisetum arvense, Carex remota.

Dominant trees : Alnus glutinosa, Fraxinus angustifolia subsp. oxycarpa

Ecology and Chorology : Alnus glutinosa tolerates the submersion better than the other tree species considered (Mondino and Bernetti 1998) and it is present only in freshwater. It forms a hygrophilous vegetation type, characterized from a chorological viewpoint by a low percentage of Mediterranean and European-Mediterranean chorotypes and a high presence of species with broad distribution (Cosmopolitan-Circumboreal and Eurosiberian chorotypes, see Fig. 2). According to the EIV analysis (Fig. 1) these woods are located in the first quadrant, positively correlated with L (Light), U (Soil Moisture, C (Continentality), and N (Nutrients), while they are negatively correlated with T (Temperature).

Figure 2. 

Chorological spectra of the vegetation types. A = Hydrocotylo-Alnetum glutinosa; F = Limnirido-Fraxinetum oxycarpae; P = Dioscoreo-Populetum nigrae; U = Periploco-Ulmetum minoris; C = Asparago-Carpinetum betuli; Q = Fraxino-Quercetum roboris.

Distribution in Tuscany (Fig. 3): These woods were found in the north-western Tuscany, especially in the plains of Serchio and Arno rivers: Selva di San Rossore (Gellini et al.1986), Macchia lucchese (Arrigoni 1990), Cerbaie (Arrigoni 1997), Tenuta di San Rossore (Tomei et al. 2004), Tenuta di Migliarino (Sani et al. 2011), Lago di Massaciuccoli (Lastrucci et al. 2017; Viciani et al. 2017), Lago di Porta (Lastrucci et al. 2016).

Figure 3. 

Distribution map of the floodplain woods of Tuscany studied. Hydrocotylo-Alnetum glutinosae (square); Limnirido-Fraxinetum oxycarpae (black circle); Dioscoreo-Populetum nigrae (white circle); Periploco-Ulmetum minoris (diamond); Asparago-Carpinetum betuli (asterisk); Fraxino-Quercetum roboris (triangle).

Syntaxonomy : the communities dominated by Alnus glutinosa can be referred to the association described by Gellini et al. (1986) for the Selva di San Rossore (PI), and previously reported by Arrigoni (1998) for Tuscany and Sburlino et al. (2011) for northern and central Italy.

LIMNIRIDO PSEUDACORI-FRAXINETUM OXYCARPAE ass. nova. Holotypus relevé 23 in Table 3 this paper – Swampy woods dominated by Fraxinus angustifolia subsp. oxycarpa (Table 3)

Table 3.

Limnirido pseudacori-Fraxinetum oxycarpae ass. nova. Rels 1–32: typical aspects; rels 33–47: transition aspects to Hydrocotylo-Alnetum.

*: typus relevé *
N. rel. 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47
Phi Limnrido pseudocori-Fraxinetum oxycarpae ass nova
54.5 Fraxinus angustifolia Vahl subsp. oxycarpa (M.Bieb. ex Willd.) Franco & Rocha Afonso 5 5 5 5 5 5 5 5 4 4 4 3 5 5 5 4 5 5 5 5 5 5 5 5 5 5 5 4 4 5 4 5 3 3 3 5 5 5 4 4 4 4 4 3 3 2 2
24 Periploca graeca L. 2 3 2 + + + + . . . . 1 1 + . . + + + 1 + + 2 . . . . . . 2 . + + 1 1 1 + + + 2 + + . . . + 1
43.1 Limniris pseudacorus (L.) Fuss . . . . . + + + . . r . . . . + + + . + + + 1 + + + . + + + + + + + + + + . . . 1 . + + . + 1
46.3 Carex elata All. subsp. elata . . . . . . . + + . . . . + + 2 + + . 4 . . 1 1 . . . . 2 + 1 1 1 + . 1 3 . . + 1 . r + . 2 +
30.7 Galium palustre L. s.l. + . + + . . + + 1 + . . . + . . . + + . + + + 2 + + + . + . . . . . + . + + + . . . r . + r 2
31.7 Ficus carica L. . . . . . . . . . . . . . . . . . . . . + 1 . . + + . . . + . + . . 1 + + r . . 1 . 1 + 2 + 2
28 Samolus valerandi L. . . . . . . + + + + . + . + . . . + + . + . . . + + . + + . + . . . . . . . . + . . . . . . .
28 Lysimachia nummularia L. . . . + . . + 1 + + . . . . . . . . + . . . . + + . . . . . 3 . . . . . . . . . . . . . . . .
24.1 Juncus acutus L. subsp. acutus . . . . . . . . . . . 1 + + . . . . . . . . . . . . . . . . + . . . . . . . . . . . . . . . .
24.1 Leucojum aestivum L. subsp. aestivum . . . . . + . . . . . . . . . . . . . . . . . + . . . + 2 . . . . . . . . . . . . . . . . . .
Frangulo-Fraxinion, Alnetalia, Alnetea
Alnus glutinosa (L.) Gaertn. . . . . . . . . + . . . . . . . + + + 1 + 1 1 + + + + 1 2 . . 1 4 3 3 3 1 2 3 2 1 3 1 3 3 3 4
Frangula alnus Mill. subsp. alnus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1 . 2 . . . + 1 1 . . 1 1 1 1 . 2 1
Thelypteris palustris Schott . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1 . . . 2 2 . . . 1 . . + . 2 1
Hydrocotyle vulgaris L. . . . . . . . + . . . . . . . . . . + . . . . . . + . 2 . . . . + . . . . . . . . . . . + . .
Lycopus europaeus L. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + + r . . . . . . . + .
Salix cinerea L. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1 . . . . . . . . . . .
Scutellaria galericulata L. . . . . . . . . . . . . . . . . . . . . . . . . + . . . . . . . . . . . . . . . . . . . . . .
Solanum dulcamara L. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . . . . . . . . . .
Magnocaricion elatae, Magnocaricetalia elatae, Phragmiti-Magnocaricetea
Mentha aquatica L. subsp. aquatica . . . . . . . . 1 . . + . + . . . 2 2 1 1 + + . 1 + . + 1 . 2 . 1 + 2 . . 2 3 + + . . . . + 1
Lythrum salicaria L. 1 + + . . + + + + . . . . . . . + . . . . + . . + + + . . 1 . . . . . . . r . . . . . . . . .
Lysimachia vulgaris L. . . . . . . . . + . . . . . . . . . + . . . + + . . . . . . . r . . . + + 1 . . r r 1 . . + .
27.9 Carex riparia Curtis . . . . . . . . + . . . . . . 3 5 . + . . + 1 . + + . . 3 . . . . + . . + . . . . . . . . . .
Alisma plantago-aquatica L. . . . . . . . . . . . . . . + . . . . . . . . . . . . . . . . r . . . . . . . . . . + . . r .
Populion albae, Populetalia, Alno-Populetea
Ulmus minor Mill. subsp. minor 3 + 1 + + 1 + 1 + 1 2 1 . . . . . . 2 + + 1 + 1 + + + . . 1 . + + 4 2 . 2 . . 4 . . 1 + 2 . 2
Carex remota L. 1 + . + . . + 4 1 . . + + . . . . . . . 1 . . 2 3 1 1 + 1 . . . . . + . . r + + . . r . 1 . .
Populus alba L. . . . . . 2 . . . . . . . + . . . . . . . . . . . . . . . 1 . 2 + . 1 + 2 + + . . . 2 + . 3 .
Carex pendula Huds. . . . . . . 4 . + . . . . . . . . . . . . . . + + 1 . . 2 . . . . . + . . r . . . . 2 . 1 . .
Brachypodium sylvaticum (Huds.) P.Beauv. subsp. sylvaticum . . . . . . 1 . + . . . . + . . . . . . . + . . . . + + . + . . . . + . . . . . . . . . + . .
Rumex sanguineus L. . . . . . + + + . . 3 . . . . . . . . . . . . 1 + + + . . . . . . . . . . . . . . . . . . . .
Populus nigra L. . . . . . . . . . . . . . . . . . . 1 . . . . . . . . 2 . . . . . . . . . . . . 2 . 1 . . 2 .
Populus canescens (Aiton) Sm. . . . . . . . . . . . . . . . . . . . . . . 2 . . . . . . . . . . . + . . . . 2 . . . . . . .
Other species
Rubus ulmifolius Schott + . + . . . + + 1 . . + + + . . . . . . . 2 . 1 + + + + . + + . + . 2 + . + 3 + . r + 1 + + 2
Agrostis stolonifera L. + 2 1 . . + . . 1 3 . 1 . 1 . . . + 2 1 + . . . + 2 + + . 1 . . 1 . . . . . + . . . . . . . .
Chamaeiris foetidissima (L.) Medik. . . . . + . . + . + . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Prunus spinosa L. subsp. spinosa . 1 1 + + . . . . . . 3 . . . . . . . . . . . . . + . . . 1 . 1 . + . 3 . + . . 1 + 2 2 . 2 +
Hedera helix L. subsp. helix . . . . + . + + . . . . . . . . . . + . . + . + + . + . 1 + . . . . . . . . + . r . . r + . .
Potentilla reptans L. . . + + . . . . 1 . . + . + . . . . . . . . . . + + + + . 1 + . + . . . . . . . . . . . + . .
Crataegus monogyna Jacq. . . + . + + . . . + . 1 . + . . . . . . . . . . . . + + . . . 1 1 . . + . . . . + . . . + . .
Carex otrubae Podp. . + + + + + . + . + 1 . . . . . . . . . . . . . . . . 1 + . . . . . . . . . . . . . . . . . .
Carex distans L. . . . . . . . . . . . . . r . . . . 1 + + + . . . . . + . . 1 . . . . . . . . . . . . + + . .
Juncus effusus L. subsp. effusus . . . . . . + . 1 . . . . . . . . . . + . . . + . + + + . . . . . . . . . . + . . . . . . . .
Ranunculus repens L. . 1 . . . . . . + . . . . . . . . . . . . . . + 1 + + . . . . . . . . . . . . . . . . . + . .
Prunella vulgaris L. subsp. vulgaris . . . . . . . . . . . . . + . . . + + . . . . + . . . + . . + . . . . . . . . . . . + . . . .
Juncus inflexus L. subsp. inflexus . . . . . 1 . . . . . . . . . + . . + . . . . . . . . . . . + . + . . . . . . . . . . . r . .
Erigeron canadensis L. . . . . . . . . + . . + . + . . . . . . . . . + . + + . . . . . . . . . . . . . . . . . . . .
Poa trivialis L. . . . . . . . . + . . . . . . . . . . . . . . + + + + . . . . . . . . . . . . . . . . . + . .
Rhamnus cathartica L. . . . . . . . . . . . . . . . . . . . . + . . . . + . . . . . . . . . + . 1 . . + . + . . . .
Cornus sanguinea L. . . . . . . . . + . . . . . . . . . 1 . . . . . . + . . . 1 . . . + . . . . . . . . . . . . .
Urtica dioica L. subsp. dioica . . . . . . . + . . . . . . . . . . . . . . . 1 1 . 2 . . . . . . . . . . . . . . . . . + . .
Oenanthe lachenalii C.C.Gmel. . . . + . + . . . . . . . . + . . . . . . . . . . . . + . . + . . . . . . . . . . . . . . . .
Plantago major L. . . . . . + . . + . . . . + . . . . . . . . . . . + . . . . + . . . . . . . . . . . . . . . .
22.3 Juncus articulatus L. subsp. articulatus . . . . . . . . . . . . . . . . . . . + + . . . . . . . . . + . + . + . . . . . . . . . . . .
Laurus nobilis L. . . . . . + . . . . . . . . . . . . . . . . . . . . . . . . . 1 . . . . . . . . 1 . . . . . +
Quercus robur L. subsp. robur 1 . . . . . . . . . . . . . . . . . . . . . + 1 . + . . . . . . . . . . . . . . . . . . . . .
Oenanthe pimpinelloides L. 1 . . . r . . . . . . . . . . . . + . . . . . . . . . . . + . . . . . . . . . . . . . . . . .
Rubus caesius L. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . r . . . + r . . . + . . . . . .
Euonymus europaeus L. . . + . r . . . . . . . . . . . . . . . . . . + . . . . . . . . . . . . . . . . . . . . . . .
Ruscus aculeatus L. . . . . + . . . . . . . . . . . . . . . . . . . . . . . . + . . . . . . . . . . . . . . + . .
Viola reichenbachiana Jord. ex Boreau . . . . . . . . . + . . . . . . . . . . . . . + . . . . . . . . . . . . . . . . . . . . + . .
Carex sylvatica Huds. . . . . . . . . + . . . . . . . . . . . . . . . . + + . . . . . . . . . . . . . . . . . . . .
Euphorbia peplus L. . . . . . . . . . + . . . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + . .
Vitis vinifera L. subsp. sylvestris (C.C. Gmel.) Hegi . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + + . . . . . . . . + . . .
Smilax aspera L. . . . . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + + . . . . . . .
Quercus ilex L. subsp. ilex . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1 . . . . + . . . . . . . r . . . .
Ranunculus sardous Crantz . . . . . . . . . . . + . . . . . . . . . . . . . . . + + . . . . . . . . . . . . . . . . . .
Rumex conglomeratus Murray + + + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Ligustrum vulgare L. . . . . r . . . . . . . . . . . . . . . . . . . . . . . . + . . . . . . . . . . . . + . . . .
Symphyotrichum squamatum (Spreng.) G.L.Nesom + + + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Schoenus nigricans L. . 2 . . . . . . . . . + . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Phragmites australis (Cav.) Trin. ex Steud. subsp. australis . . . . . . . . . . . . . . . . + . . . . . . . . . . + . + . . . . . . . . . . . . . . . . .
Persicaria lapathifolia (L.) Delarbre subsp. lapathifolia . . . . . . . . + . . . . . . . . . . . . . . + + . . . . . . . . . . . . . . . . . . . . . .
Jacobaea erucifolia (L.) G.Gaertn., B.Mey. & Scherb. subsp. erucifolia + + + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Clematis vitalba L. . . . . . . . . . . . . . . . . . . . . . . . . + . + . . + . . . . . . . . . . . . . . . . .
Carex vulpina L. . . . . . . . + . . . + . + . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Carex viridula Michx. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . + r . . . . 2 . . . .
Cardamine pratensis L. . . . . . . . . . . . . . . . . . . . . . . . + + . + . . . . . . . . . . . . . . . . . . . .
Convolvulus sepium L. . . . . . . . . . . . . . . . + 2 . . . . . . . . . . . . + . . . . . . . . . . . . . . . . .

Diagnostic species : Fraxinus angustifolia subsp. oxycarpa, Carex elata, Ficus carica, Galium palustre, Limniris pseudacorus, Samolus valerandi, Leucojum aestivum, Lysimachia nummularia, Carex riparia, Periploca graeca.

Dominant trees : Fraxinus angustifolia subsp. oxycarpa, Alnus glutinosa.

Ecology and Chorology : Fraxinus angustifolia subsp. oxycarpa can tolerate short periods of moderate water deficit and low concentrations of salt in soil waters. In general terms, this species is more Mediterranean, relatively more thermophilous than Alnus glutinosa. From a chorological viewpoint, these woods are firstly characterized by the European-Mediterranean and Eurasian chorotypes (Fig. 2). According to the EIV analysis (Fig. 1), these woods are positively related to Soil Moisture (U) and Light (L).

Distribution in Tuscany (Fig. 3): These woods occupy the interdune areas along the Tyrrhenian coasts, such as San Rossore (Gellini et al. 1986), Bosco dell’Ulivo (Coaro 1987), Castagneto Carducci (Foggi et al. 2000), Macchia Lucchese (Arrigoni 1990), Tenuta del Tombolo di San Rossore (Tomei et al. 2004), Tenuta di Migliarino (Sani et al. 2011), Tenuta di Coltano (Bertacchi and Lombardi 2016) and Padule di Bolgheri.

Syntaxonomy : Gellini et al. (1986) referred this type of woods to the Carici remotae-Fraxinetum oxycarpae Pedrotti corr. Pedrotti 1993, a riparian forest association. The woods dominated by Fraxinus angustifolia subsp. oxycarpa here presented, instead, are clearly characterized by a higher presence of swamp species with respect to riverine Carici remotae-Fraxinetum oxycarpae coenoses. For example, species like Stachys sylvatica, Symphytum tuberosum, Ranunculus lanuginosus are not present (Gellini et al. 1986), while several Carex species were frequently found.

Gellini et al. (1986) suggested the establishment of a new subassociation, Carici-Fraxinetum oxycarpae alnetosum glutinosae, but the name is invalid on the basis of Art. 5 of the ICPN (Theurillat et al. 2021). Furthermore, the two relevés attributed to this subassociation were included in the Hydrocotylo-Alnetum table, therefore, we decided to not validate the subassociation. However, it must be noted that the relevés 33–47 of Table 3, where Alnus glutinosa has high cover values, show a transition to the swampy woods of Hydrocotylo-Alnetum.

Fraxinus angustifolia subsp. oxycarpa, in Tuscany, has broad ecological needs; according to Mondino and Bernetti (1998) it can be found in swamp areas with Alnus glutinosa and Quercus robur, riparian habitats with Ulmus minor and, finally, in hilly meso-hygrophilous conditions with Quercus cerris (Scoppola and Filesi 1995; Foggi et al. 2000; Terzi et al. 2020). Marshy woods are often characterized by a vegetation mosaic of forest stands with Juncetalia maritimi, Magnocaricion elatae and Phragmition. A similar situation is reported for the Bosco della Mesola site, in North-Eastern Italian Adriatic coast (Piccoli et al. 1983; Piccoli and Gerdol 1984; Gerdol et al. 2018). In in a similar landscape context, for the Tuscan north-western Thyrrenian coast, Tomei et al. (2004) proposed the new association Carici elatae-Fraxinetum oxycarpae, not validly published according to the Art. 5 of the ICPN (Theurillat et al. 2021). Moreover, it was based on only two relevés that, in our opinion, are not representative of a wood, being the cover value of the tree layer very low. For these reasons, we decided not to validate this name and we here propose to attribute these coenoses to a new association named Limnirido pseudacori-Fraxinetum oxycarpae, which is characterized by the presence of a high contingent of Phragmiti-Magnocariceta species compared to other similar associations, i.e. Fraxino-Quercetum roboris observed for the low interdune swamp areas of Migliarino-San Rossore by Gellini et al. (1986), Carici remotae-Fraxinetum oxycarpae reported by Pedrotti (1970, 1993), Gellini et al. (1986), Mondino and Bernetti (1998), Arrigoni (1998), Mercadal and Vilar (2013) and, finally, Alno-Fraxinetum oxycarpae (Arrigoni 1998).

The floristic differences between the other associations dominated by Fraxinus angustifolia subsp. oxycarpa described for Italy (Tables 1 and 2 in Poldini and Sburlino 2018) and the similar coenoses detected in the Tuscan floodplain woods are displayed in a synoptic table (Table 4) and in the dedicated NMDS ordination (Fig. 4). Table 4 shows that columns on the left side (columns 1–12) can be assigned to the swampy alliance Frangulo alni-Fraxinion, while those at the right side (columns 13–20) can be attributed to the riparian alliance Carici remotae-Fraxinion (Poldini and Sburlino 2018). According to this scheme, we refer the association Limnirido-Fraxinetum oxycarpae to the alliance Frangulo-Fraxinion. This interpretation was also confirmed by the cluster analysis (Suppl. material 1: Figure S1) through which the riparian woods are clustered together and separated from the swampy woods. Even if these coenoses also host a high rate of Alno-Populetea and Populetalia species (Table 3), the attribution of this new association to the Alnetea and Alnetalia is also justified for its ecological position, as it is typically located in the interdune swamps. Also, the Fraxinus angustifolia subsp. oxycarpa communities recently indicated for Corsica (Gauberville et al. 2018) as Carici remotae-Fraxinetum can be probably attributed to Limnirido-Fraxinetum, considering their floristic composition and ecology.

In the end, we note that in Tuscany the distinction between the alliances Alnion glutinosae and Frangulo-Fraxinion is not so clear, as many diagnostic species are in common, and a comprehensive work devoted to clarify this problem is needed.

Figure 4. 

NMDS ordination scatterplot for the synthetic tables of the Italian woods dominated by Fraxinus angustifolia subsp. oxycarpa. Crem-Fx: Carici remotae-Fraxinetum; Cla-Fx: Cladio marisci-Fraxinetum; Sal-Fx: Salici apenninae-Fraxinetum; Rub-Fx: Rubo caesi-Fraxinetum; Lys-Fx: Lysimachio-Fraxinetum; Leu-Fx: Leucojo verni-Fraxinetum; Val-Fx: Valeriano-Fraxinetum; Lim-Fx: Limnirido pseudacori-Fraxinetum. Data from Tables 12 by Poldini and Sburlino (2018) with the exception of Lim-Fx, data from this paper.

Table 4.

Synoptic table of Fraxynus angustifolia subsp. oxycarpa communities, including Tuscan relevés and Italian relevés by Poldini and Sburlino (2018). Column legend: 1: Tuscany, Macchia Lucchese (Arrigoni, 1990); 2: Tuscany, Tenuta di San Rossore (Tomei et al., 2004); 3: Tuscany, Selva di San Rossore (Gellini et al., 1986); 4: Tuscany, Tenuta di Migliarino (Sani et al., 2011); 5: Tuscany, Bosco dell’Ulivo (Coaro, 1987); 6: Tuscany, Tenuta del Tombolo di Pisa (AAVV, 2005); 7–20: from tables 1 and 2 by Poldini and Sburlino (2018); 7: Emilia-Romagna; 8: Emilia-Romagna; 9: Latium; 10: Croatia; 11: Croatia; 12: Friuli-Venezia Giulia; 13: Abruzzo; 14: Abruzzo; 15: Apulia; 16: Abruzzo; 17: Friuli-Venezia Giulia and Veneto; 18: Campania; 19: Emilia-Romagna; 20: Marche. Association abbreviation legend: Lim-Fx: Limnirido pseudacori-Fraxinetum; Cla-Fx: Cladio marisci-Fraxinetum; Leu-Fx: Leucojo verni-Fraxinetum; Val-Fx: Valeriano dioicae-Fraxinetum; Crem-Fx: Carici remotae-Fraxinetum; Lys-Fx: Lysimachio nummulariae-Fraxinetum; Rub-Fx: Rubo caesi-Fraxinetum; Sal-Fx: Salici apenninae-Fraxinetum.

Column n. 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20
Association abbreviation Lim-Fx1 Lim-Fx2 Lim-Fx3 Lim-Fx4 Lim-Fx5 Lim-Fx6 Cla-Fx1 Cla-Fx2 Cla-Fx3 Leu-Fx1 Leu-Fx2 Val-Fx Crem-Fx1 Crem-Fx2 Crem-Fx3 Crem-Fx4 Lys-Fx Rub-Fx Rub-Ulm Sal-Fx
Frangulo-Fraxinion, Alnetalia, Alnetea, Magnocaricion elatae species n. 14 8 9 10 4 8 16 8 12 19 13 14 0 3 2 1 5 1 2 0
Carici-Fraxinion, Populetalia, Alno-Populetea species n. 5 6 7 8 6 5 4 2 2 3 4 2 11 18 16 5 14 11 6 7
Frangulo-Fraxinion, Alnetalia, Alnetea
Alnus glutinosa (L.) Gaertn. 100 22 86 82 10 11 22 . . 17 100 50 . . . . . . . .
Dryopteris carthusiana (Vill.) H.P.Fuchs . . . . . . . . . 13 60 . . . . . . . . .
Frangula alnus Mill. subsp. alnus 90 . . . 10 22 100 100 100 17 80 100 . . . . . . 40 .
Galium palustre L. s.l. 30 56 100 53 30 33 11 86 83 100 100 83 . . . 56 17 . . .
Hydrocotyle vulgaris L. . 11 14 12 . . . 29 . . . . . . . . . . . .
Lycopus europaeus L. 50 . . . . . 67 14 33 96 80 67 . 47 50 . . 29 10 .
Cladium mariscus (L.) Pohl . . . . . . 56 100 50 . . 33 . . . . . . . .
Salix cinerea L. 10 . . . . . 44 . 50 . . 17 . . . . . . . .
Thelypteris palustris Schott 50 . . . . 11 22 71 . . . 50 . . . . . . . .
Magnocaricion elatae
Mentha aquatica L. subsp. aquatica 30 11 43 82 . 11 89 57 83 91 40 33 . . . . . . . .
Limniris pseudacorus (L.) Fuss 60 11 43 71 . 22 89 . 50 100 100 67 . 29 17 . 67 . . .
Carex elata All. subsp. elata 60 11 43 41 . 11 78 . 17 26 . 100 . . . . . . . .
Lythrum salicaria L. 20 17 57 18 . 56 56 . 100 100 . 67 . . . . . . . .
Lysimachia vulgaris L. 70 . 29 12 . . 89 . 67 78 100 50 . . . . . . . .
Carex riparia Curtis 10 . 43 35 . . 56 . . 43 . . . . . . . . . .
Alisma plantago-aquatica L. 20 . . . . . 22 . . 30 40 . . . . . . . . .
Rumex triangulivalvis (Danser) Rech.f. . . . . 20 . 11 . . 4 . . . . . . . . . .
Stachys palustris L. . . . . . . 78 . . 91 100 . . . . . . . . .
Alisma lanceolatum With. 10 . . . . . . . . 100 . . . . . . . . . .
Glyceria fluitans (L.) R.Br. . . . . . . . . . 39 20 . . . . . . . . .
Poa palustris L. subsp. palustris . 11 . . . . . . . . 20 . . . . . . . . .
Carex acutiformis Ehrh. . . . . . . . 86 . 4 . . . . . . 67 . . .
Eupatorium cannabinum L. subsp. cannabinum . . . 6 . . . . 33 . 40 67 . 6 . . 17 . . .
Cyperus longus L. . . . . . . . . 33 . . . . . . . . . . .
Oenanthe aquatica (L.) Poir. . . . . . . . . . 13 . . . . . . . . . .
Sparganium erectum L. . . . . . . . . . 22 . . . . . . . . . .
Veronica beccabunga L. . . . . . . . . . . 17 . . . . . . . .
Phalaris arundinacea L. subsp. arundinacea . . . . . . . . . . . . . . . . 50 . . .
Carici-Fraxinion, Populetalia, Alno-Populetea . . . . . . . . . . . . . . . . . . . .
Carex pendula Huds. 20 33 57 12 . . . . . . . . 80 94 100 86 67 57 100 33
Brachypodium sylvaticum (Huds.) P.Beauv. subsp. sylvaticum 10 56 43 18 10 44 . . . . . . 60 59 100 . 67 86 60 .
Carex remota L. 20 61 100 35 30 33 . . . 43 80 . 100 47 . . 83 . . .
Vitis vinifera L. 10 . . 6 10 . 22 29 67 . . . . 29 33 . 14 20 .
Convolvulus sepium L. . . . 12 . 11 44 29 83 . 40 17 . . . . 33 14 . .
Populus nigra L. 20 . 14 12 . . 33 . . . . 50 . 12 100 . 67 . . 67
Rumex sanguineus L. . 22 86 6 . . . . . 39 80 . 100 88 100 . 67 . . .
Geum urbanum L. . . . . . 22 . . . . 60 . 20 18 67 57 . 57 . .
Ranunculus lanuginosus L. . . . . 30 11 . . . . . . 100 88 83 86 . 86 . .
Dioscorea communis (L.) Caddick & Wilkin . . . . . . . . . . . . . 71 100 43 17 43 . 22
Arum italicum Mill. subsp. italicum . . . . 60 . . . . . . . 100 82 71 17 71 . .
Equisetum telmateia Ehrh. . 17 . . . . . . . . . . 20 18 50 . 33 . . 55
Salix alba L. . . . . . . 67 . . 39 . . . . 50 . 50 . 60 89
Carex divulsa Stokes . 6 14 . . . . . . . . . 40 12 67 . . . . .
Bryonia dioica Jacq. . . . . . . . . . . . . 20 47 17 . 17 . . .
Clematis viticella L. . . . . . . . . . . . . . 35 . . 83 43 90 .
Symphytum tuberosum L. subsp. angustifolium (A.Kern.) Nyman . . . . . . . . . . . . 20 18 100 . . 57 . .
Sambucus nigra L. . . 14 . . . . . . . . . . . 17 . 83 29 . .
Malus sylvestris (L.) Mill. . . . 12 40 . . . . . . . . . 17 . . . 22 .
Humulus lupulus L. . . . . . . . . . . . . . 12 67 . 17 . . .
Salix purpurea L. . . . . . . . . . . . . . 53 . . . . . 44
Chaerophyllum temulum L. . . . . . . . . . . . . . 47 . . . . . .
Salix triandra L. subsp. triandra . . . . . . . . . . . . . . . . . . . 11
Other species
Fraxinus angustifolia Vahl subsp. oxycarpa (M.Bieb. ex Willd.) Franco & Rocha Afonso 100 100 100 100 100 100 100 100 100 100 100 100 100 100 100 100 100 100 100 100
Ulmus minor Mill. subsp. minor 40 83 100 47 50 100 44 86 . 22 80 50 100 94 100 86 67 86 100 67
Rubus ulmifolius Schott 70 89 100 41 70 89 22 71 100 . . . 80 100 83 78 17 43 . 67
Quercus robur L. subsp. robur . 56 57 6 70 56 56 71 67 48 80 . 80 18 . . 67 . 100 11
Populus alba L. 60 34 14 36 30 44 78 100 . . . . 100 82 . 14 . 14 100 78
Cornus sanguinea L. . . 43 12 . 56 11 43 . . . 67 60 65 67 89 83 43 80 100
Prunella vulgaris L. subsp. vulgaris 20 17 29 24 . . . . . 17 60 17 80 12 83 56 . 14 . 44
Ligustrum vulgare L. 20 . . . 60 56 . 43 . . . 17 . 76 83 22 17 57 50 11
Ruscus aculeatus L. 20 39 . . 80 56 . 29 . . . 50 80 53 83 89 . 100 . 11
Potentilla reptans L. . 28 71 12 10 22 . 43 33 30 . . 20 35 . . . . . 11
Ranunculus repens L. . 11 100 . 20 11 . . . 100 60 67 47 50 . 33 . . 11
Agrostis stolonifera L. . 39 86 41 . 44 11 57 50 13 20 . . . . . . 43 . .
Hedera helix L. subsp. helix 40 67 57 29 80 67 11 57 67 . . 50 80 88 50 100 67 100 . 44
Smilax aspera L. 20 . . 6 80 22 . . 50 . . . 40 47 17 22 29 .
Crataegus monogyna Jacq. 40 56 43 12 70 56 67 71 . . 100 . . 100 17 71 . 89
Carex otrubae Podp. . 17 . 18 30 22 . . 67 . . . . . 50 11 17 14 .
Clematis vitalba L. 10 . 29 . 10 44 . . . . . . 20 18 50 . . . 30 22
Euonymus europaeus L. . . 29 6 50 56 . . . . . . . 76 83 56 17 71 . .
Quercus ilex L. subsp. ilex 30 6 . 6 30 44 . 43 33 . . . . 6 . . . 14 . .
Acer campestre L. 10 28 29 . 10 11 . . 17 . 20 17 20 59 33 33 100 14 40 55
Carex flacca Schreb. subsp. flacca 10 6 . . . 11 . 43 67 . . . . . . 22 29 . 44
Carex sylvatica Huds. . 22 71 . . 22 . . . . . . 20 18 . 100 17 14 . .
Leucojum aestivum L. subsp. aestivum . . 14 18 . . 11 . . 100 60 100 . 24 . . 67 . . .
Lysimachia nummularia L. . 22 43 12 20 . 44 . . 100 100 . . . . . 83 . . .
Poa trivialis L. . 11 86 . . . . . . 83 80 . 100 35 83 . 50 . . .
Viola reichenbachiana Jord. ex Boreau . 44 43 . 50 22 . . . . . . 20 12 . . 17 14 . .
Prunus spinosa L. subsp. spinosa 80 6 29 . 70 56 89 86 . . . . 60 12 83 44 50 57 100 100
Rhamnus cathartica L. 40 . 14 6 . 11 78 14 . . . 50 . . . . . . . .
Asparagus acutifolius L. . . . . 20 11 . . . . . . . 24 17 22 . 86 . 11
Crataegus laevigata (Poir.) DC. . . . . . . . . . . 60 80 71 . 67 17 14 30 .
Lonicera caprifolium L. . . . . 70 67 . . . . . . 20 59 . . 17 29 . 33
Rosa sempervirens L. . . . . 30 22 . . . . . . . 59 83 44 . 100 . 22
Rubia peregrina L. 10 . . . 70 56 . . . . . . . 71 33 11 . 86 . .
Urtica dioica L. subsp. dioica . 11 57 . . . 11 . . 57 60 . . 24 . . 67 . . .
Rubus caesius L. 50 . . . . . 11 29 . 26 60 33 . . 17 100 14 100 89
Carex distans L. 20 11 . 35 10 . . 57 17 . . . . . . . . . . .
Ajuga reptans L. . . 14 . 20 . . . . . . . . 12 . 33 17 14 . .
Ficaria verna Huds. . . . . . . . . . 9 20 . 100 71 100 . . 29 . .
Chamaeiris foetidissima (L.) Medik. . 44 14 . 80 33 . . . . . . . . . 100 . 57 . .
Samolus valerandi L. . 28 43 41 10 . . 57 50 . . . . . . . . . . .
Solanum dulcamara L. 10 . . . . . 89 . . 74 100 17 . 47 83 . . . . .
Carex vulpina L. . 17 . . . . . . . 57 80 20 47 . . . . . .
Ficus carica L. 70 6 29 29 . 22 . . . . . . . . . . . . . .
Fraxinus ornus L. subsp. ornus . . . . 10 . . . . . . . . . . 43 . 43 10 44
Juncus effusus L. subsp. effusus . 17 86 18 . . . . . 22 60 . . . . . . . . .
Laurus nobilis L. 30 11 . 6 . 33 . . . . . . . . . . . 86 . .
Periploca graeca L. 60 28 . 71 70 67 . . . . . . . . . . . . . .
Phragmites australis (Cav.) Trin. ex Steud. subsp. australis . . . 12 . 11 11 . 17 . . 50 . . . . . . . .
Pyrus communis L. . . . . 10 11 56 43 . . 40 . . . . . . . . .
Stellaria media (L.) Vill. subsp. media . . 14 . . . . . . . . . . 6 17 14 . 14 . .
Cyclamen repandum Sm. subsp. repandum . . 14 . 10 . . . . . . . . 24 17 . . . . .
Holcus lanatus L. subsp. lanatus . 28 14 . . . . 14 17 . . . . . . . . . . .
Ranunculus sardous Crantz . 6 . 18 10 . . . . . . . . . . 44 . . . .
Stachys sylvatica L. . . . . . . . . . . . . 40 53 100 11 . . . .
Thysselinum palustre (L.) Hoffm. . . . . . . 11 . . 9 40 17 . . . . . . . .
Viola alba Besser subsp. alba . . . . . . . . . . . . . . 33 29 . 29 . 22
Bellevalia romana (L.) Sweet . . . . . . . . . . . . . 18 33 29 . 29 . .
Oenanthe pimpinelloides L. . . . 6 10 22 . . . . . . 100 41 83 . . . . .
Arum maculatum L. . . . . . . . . . . . . . 47 100 . . 14 . .
Bidens tripartita L. s.l. . . . . . . 33 . . 17 20 . . . . . . . . .
Caltha palustris L. . . . . . . . . . 22 40 33 . . . . . . . .
Cardamine pratensis L. . 6 43 . 20 . . . . . . . . . . . . . . .
Carpinus betulus L. . 6 . . 20 11 . . . . . . . . . . . . . .
Galium aparine L. . . . . . . . . . . . . . . 67 29 . 29 . .
Genista tinctoria L. . . . . . . . . . 48 40 . . . . . . . . 67
Hypericum tetrapterum Fr. . . 14 . . . . . 17 . 60 . . . . . . . . .
Juncus articulatus L. subsp. articulatus . . 24 . . . . 17 . . 17 . . . . . . . .
Luzula forsteri (Sm.) DC. . 6 14 . 10 . . . . . . . . . . . . . . .
Phalaris arundinacea L. subsp. arundinacea . . . 6 . . . . . 30 . 17 . . . . . . . .
Plantago major L. . 6 29 12 . . . . . . . . . . . . . . . .
Pteridium aquilinum (L.) Kuhn subsp. aquilinum . 11 . 6 . 11 . . . . . . . . . . . . . .
Scutellaria galericulata L. . . 14 . . . . . . 57 20 . . . . . . . . .
Sison amomum L. . . . . . . . . . . . . 20 29 50 . . . . .
Viburnum opulus L. . . . . . . 33 . . . . 67 . . . . . . . .
Althaea officinalis L. . . . . . . 11 . 17 . . . . . . . . . . .
Amorpha fruticosa L. . . . 12 . . . . . . . . . . . . . . 90 .
Aristolochia clematitis L. . . . . . . . . . 13 20 . . . . . . . . .
Carex ornithopoda Willd. . . . . . . . . . 52 80 . . . . . . . . .
Carex vesicaria L. . . . . . . . . . 83 40 . . . . . . . . .
Cornus mas L. . . . . 50 11 . . . . . . . . . . . .
Deschampsia cespitosa (L.) P.Beauv. . . . . . . . . . 22 20 . . . . . . . . .
Dittrichia viscosa (L.) Greuter subsp. angustifolia (Bég.) Greuter . . . 6 20 . . . . . . . . . . . . . . .
Erigeron canadensis L. . 11 57 . . . . . . . . . . . . . . . . .
Euphorbia palustris L. . . . . . . 33 . . 91 . . . . . . . . . .
Glechoma hederacea L. . . . . . . . . . 4 80 . . . . . . . . .
Gratiola officinalis L. . . . 12 . . . . . 57 . . . . . . . . . .
Hypericum androsaemum L. . 11 29 . . . . . . . . . . . . . . . . .
Juncus acutus L. subsp. acutus . 17 . 6 . . . . . . . . . . . . . . . .
Juncus inflexus L. subsp. inflexus . 11 . 24 . . . . . . . . . . . . . . . .
Juniperus communis L. . . . . 10 . . . . . 40 . . . . . . . . .
Melica uniflora Retz. . . . . 40 11 . . . . . . . . . . . . . .
Myosotis sylvatica Hoffm. subsp. sylvatica . 6 29 . . . . . . . . . . . . . . . . .
Oenanthe fistulosa L. . . . . . . . . . 91 20 . . . . . . . . .
Oenanthe lachenalii C.C.Gmel. . . . 18 60 . . . . . . . . . . . . . . .
Oxalis corniculata L. . 6 14 . . . . . . . . . . . . . . . . .
Persicaria hydropiper (L.) Delarbre . . . . . . . . . 78 80 . . . . . . . . .
Pinus pinea L. 20 . . . 10 . . . . . . . . . . . . . . .
Primula vulgaris Huds. subsp. rubra (Sm.) Arcang. . . . . . . . . . . . . . 6 . . . . . 11
Quercus cerris L. . . . . . . . . . . . . 20 12 . . . . . .
Quercus pubescens Willd. subsp. pubescens . . . . . . . . . . . . . . 17 . . . . 55
Rorippa amphibia (L.) Besser . . . . . . 22 . . 87 . . . . . . . . . .
Rumex conglomeratus Murray . 6 . . . 44 . . . . . . . . . . . . . .
Schoenus nigricans L. . 11 . . . 11 . . . . . . . . . . . . . .
Sium latifolium L. . . . . . . 33 . . 83 . . . . . . . . . .
Symphytum officinale L. . . . . . . 11 . . . . 17 . . . . . . . .
Teucrium scordium L. subsp. scordioides (Schreb.) Arcang. . . . . . . . . . 87 20 . . . . . . . . .
Tussilago farfara L. . . . . . . . . . . . . . . 17 . . . . 22
Valeriana dioica L. . . . . . . . . . . 80 100 . . . . . . . .
Veronica montana L. . 17 14 . . . . . . . . . . . . . . . . .
Veronica scutellata L. . . . . . . . . . 61 40 . . . . . . . . .
Veronica serpyllifolia L. . 6 29 . . . . . . . . . . . . . . . . .
Aegopodium podagraria L. . . . . . . . . . . 40 . . . . . . . . .
Althaea cannabina L. . . . 6 . . . . . . . . . . . . . . . .
Anthoxanthum odoratum L. . 6 . . . . . . . . . . . . . . . . . .
Aristolochia rotunda L. . . 14 . . . . . . . . . . . . . . . . .
Asplenium onopteris L. . . . . 10 . . . . . . . . . . . . . . .
Athyrium filix-femina (L.) Roth . . . . . . . . . . 60 . . . . . . . . .
Berula erecta (Huds.) Coville . . . . . . . . . . . 17 . . . . . . . .
Bidens frondosa L. 10 . . . . . . . . . . . . . . . . . . .
Blackstonia perfoliata (L.) Huds. . 6 . . . . . . . . . . . . . . . . . .
Brachypodium rupestre (Host) Roem. & Schult. . . . . . . . . . . . . . . . . . . . 55
Cardamine pratensis L. . . . . . . . . . 100 . . . . . . . . . .
Carex canescens L. . . 14 . . . . . . . . . . . . . . . . .
Carex caryophyllea Latourr. . 11 . . . . . . . . . . . . . . . . . .
Carex flava L. . . . . . . . . . . . 50 . . . . . . . .
Carex punctata Gaudin . . 14 . . . . . . . . . . . . . . . . .
Carex strigosa Huds. . . . . . 22 . . . . . . . . . . . . . .
Carex viridula Michx. 30 . . . . . . . . . . . . . . . . . . .
Centaurium pulchellum (Sw.) Druce subsp. pulchellum . 6 . . . . . . . . . . . . . . . . . .
Centaurium tenuiflorum (Hoffmanns. & Link) Fritsch . 6 . . . . . . . . . . . . . . . . . .
Cerastium semidecandrum L. . 11 . . . . . . . . . . . . . . . . . .
Euphorbia peplis L. . . 14 . . . . . . . . . . . . . . . . .
Circaea lutetiana L. subsp. lutetiana . . . . . 11 . . . . . . . . . . . . . .
Clematis flammula L. . . . . 10 . . . . . . . . . . . . . . .
Crepis bellidifolia Loisel. . 6 . . . . . . . . . . . . . . . . . .
Silene baccifera (L.) Durande . . . . . . . . . . . . . . . . 17 . . .
Cyclamen hederifolium Aiton . . . . . . . . . . . . . . . . . 43 . .
Elymus repens (L.) Gould subsp. repens . . . . . 11 . . . . . . . . . . . . . .
Equisetum arvense L. . . 14 . . . . . . . . . . . . . . . . .
Euphorbia peplis L. . 22 . . . . . . . . . . . . . . . . . .
Fallopia convolvulus (L.) Á.Löve . . . . . 11 . . . . . . . . . . . . . .
Fragaria vesca L. subsp. vesca . . . . 10 . . . . . . . . . . . . . . .
Galium debile Desv. . . 14 . . . . . . . . . . . . . . . . .
Galium rotundifolium L. subsp. rotundifolium . 6 . . . . . . . . . . . . . . . . . .
Geranium purpureum Vill. . . 14 . . . . . . . . . . . . . . . . .
Geranium robertianum L. . 11 . . . . . . . . . . . . . . . . . .
Geranium sylvaticum L. . 6 . . . . . . . . . . . . . . . . . .
Hypochaeris glabra L. . . . . 10 . . . . . . . . . . . . . . .
Isolepis setacea (L.) R.Br. . . . 6 . . . . . . . . . . . . . . . .
Jacobaea erucifolia (L.) G.Gaertn., B.Mey. & Scherb. . . . . . 33 . . . . . . . . . . . . . .
Juncus conglomeratus L. . 6 . . . . . . . . . . . . . . . . . .
Lonicera japonica Thunb. 10 . . . . . . . . . . . . . . . . . . .
Lonicera etrusca Santi . . . . . . . . . . . . . . 67 . . . . .
Lotus corniculatus L. subsp. alpinus (DC.) Rothm. . . . . 10 . . . . . . . . . . . . . . .
Lysimachia arvensis (L.) U.Manns & Anderb. . . 14 . . . . . . . . . . . . . . . . .
Mentha x verticillata L. . . . . . . . . . 83 . . . . . . . . . .
Moehringia trinervia (L.) Clairv. . . 29 . . . . . . . . . . . . . . . . .
Myosotis laxa Lehm. subsp. cespitosa (Schultz) Hyl. ex Nordh. . 11 . . . . . . . . . . . . . . . . .
Myosotis scorpioides L. subsp. scorpioides . . . . . . . . . 96 . . . . . . . . . .
Myrtus communis L. . . . . 30 . . . . . . . . . . . . . . .
Phelipanche purpurea (Jacq.) Soják . 6 . . . . . . . . . . . . . . . . . .
Parietaria officinalis L. . . . . . . . . . . . . . . . . 50 . . .
Persicaria lapathifolia (L.) Delarbre subsp.
lapathifolia
. . 57 . . . . . . . . . . . . . . . . .
Phillyrea angustifolia L. . . . . 10 . . . . . . . . . . . . . . .
Phytolacca americana L. . . 14 . . . . . . . . . . . . . . . . .
Pinus pinaster Aiton subsp. pinaster . . . . 10 . . . . . . . . . . . . . . .
Platanthera chlorantha (Custer) Rchb. . . . . 30 . . . . . . . . . . . . . . .
Platanus sp. 10 . . . . . . . . . . . . . . . . . . .
Poa annua L. . . 14 . . . . . . . . . . . . . . . . .
Poa balbisii Parl. . 6 . . . . . . . . . . . . . . . . . .
Potentilla sterilis (L.) Garcke 20 . . . . . . . . . . . . . . . . . . .
Pulicaria dysenterica (L.) Bernh. . . 14 . . . . . . . . . . . . . . . . .
Pyracantha coccinea M.Roem. . . . . . 11 . . . . . . . . . . . . . .
Ranunculus flammula L. . . 14 . . . . . . . . . . . . . . . . .
Ranunculus sp. . . 14 . . . . . . . . . . . . . . . . .
Rorippa sylvestris (L.) Besser subsp. sylvestris . . . . . . . . . 43 . . . . . . . . . .
Rosa canina L. . . . . . . . . . . . . . . . . . . 44
Rubus sp. . . . 12 . . . . . . . . . . . . . . . .
Salix apennina A.K.Skvortsov . . . . . . . . . . . . . . . . . . . 100
Lolium arundinaceum (Schreb.) Darbysh. subsp. arundinaceum . 6 . . . . . . . . . . . . . . . . . .
Schenkia spicata (L.) G.Mans. . 6 . . . . . . . . . . . . . . . . . .
Scirpoides holoschoenus (L.) Soják . . . . 10 . . . . . . . . . . . . . . .
Scrophularia nodosa L. . . 14 . . . . . . . . . . . . . . . . .
Lychnis flos-cuculi L. subsp. flos-cuculi . . 29 . . . . . . . . . . . . . . . . .
Betonica officinalis L. . . . . 10 . . . . . . . . . . . . . . .
Succisa pratensis Moench . . . . . . . . . . 60 . . . . . . . . .
Symphyotrichum squamatum (Spreng.) G.L.Nesom . . . . . 33 . . . . . . . . . . . . . .
Taxodium distichum (L.) Rich. . . . 18 . . . . . . . . . . . . . . . .
Thalictrum flavum L. . . . . . . . . . 43 . . . . . . . . . .
Thalictrum lucidum L. . . . . . . . . . . . 83 . . . . . . . .
Trifolium fragiferum L. subsp. fragiferum . . . . 10 . . . . . . . . . . . . . . .
Triglochin barrelieri Loisel. . . . . 20 . . . . . . . . . . . . . . .
Galatella pannonica (Jacq.) Galasso, Bartolucci & Ardenghi . . . . 10 . . . . . . . . . . . . . . .
Veronica anagallis-aquatica L. subsp.
anagallis-aquatica
. . . . . . . . . 57 . . . . . . . . . .
Veronica chamaedrys L. subsp. chamaedrys . . . . 20 . . . . . . . . . . . . . . .
Veronica officinalis L. . . 14 . . . . . . . . . . . . . . . .

FRAXINO OXYCARPAE-QUERCETUM ROBORIS Gellini, Pedrotti et Venanzoni, 1986. Holotypus: Table 3, rel. 14 in Gellini, Pedrotti et Venanzoni, 1986 (corresponding to rel. n. 40 in Suppl. material 2: Table S1) – Mesohygrophylous forests dominated by Quercus robur and Fraxinus angustifolia subsp. oxycarpa (Suppl. material 2: Table S1)

Diagnostic species : Quercus robur, Cornu mas, Acer campestre, Luzula forsteri, Moheringia trinervia, Chamaeiris foetidissima, Lonicera caprifolium, Ruscus aculeatus, Viola reichembachiana, Ajuga reptans, Ligustrum vulgare, Veronica montana.

Dominant trees : Quercus robur, Fraxinus angustifolia subsp. oxycarpa, Acer campestre, Carpinus betulus, Ulmus minor, Quercus ilex (only in the lauretosum nobilis subassociation).

Ecology and Chorology : Meso-hygrophilous, Mediterranean woods developing on fertile and deep soils. The chorotypes spectrum is very similar to that of the Fraxinus angustifolia subsp. oxycarpa woods, apart for a slight increase in European species (Figure 2). These communities were found in the more elevated areas of the swamps, in contact with the climatic woods, which act as suppliers of many euryoecious species, tolerating seasonal variations of hygromorphic conditions. According to the EIV analysis (Fig. 1), these woods are positively related to Temperature (T), and negatively to Soil Moisture (U), Continentality (C) and Light (L).

Distribution in Tuscany (Fig. 3) : These communities were found in San Rossore (Gellini et al. 1986; Tomei et al. 2004), Bosco dell’Ulivo (Coaro 1987), Macchia Lucchese (Arrigoni 1990), Cerbaie (Arrigoni1997), Bosco dei Renacci (Viciani and Gabellini 2012), Bosco di Chiusi (Tomei and Cenni 1986), Coltano (Bertacchi and Lombardi 2016), and according to Gellini et al. (1986), also in Migliarino and Viareggio.

Syntaxonomy : English oak forests in Tuscany were originally described by Gellini et al. (1986) for the Selva di San Rossore and referred to the association Fraxino oxycarpae-Quercetum roboris Gellini, Pedrotti, Venanzoni, 1986. The presence of the association was then confirmed by Mondino and Bernetti (1998) and Arrigoni (1998) for the same areas. The association can be divided into four subassociations (Suppl. material 2: Table S1):

1. subass. typicum Rels 1–45 in Suppl. material 2: Table S1; it represents the typical aspects of the association (holotypus – the same of the association – Table 3, rel. 14 in Gellini et al. 1986).

2. carpinetosum betuli Gellini, Pedrotti, Venanzoni, 1986 ex Gennai et al. subass. nova hoc loco (lectotypus: rel. 19 Table 3, in Gellini et al. 1986, corresponding to rel. n. 50 in Suppl. material 2: Table S1; lectotypus hoc loco) Rels 46–52 in Suppl. material 2: Table S1; it is differentiated by the dominance of Carpinus betulus and by the presence of Pteridium aquilinum. This subassociation was reported but not validly typified by Gellini et al. (1986) (Art. 5 of the ICPN – Theurillat et al. 2021), so here we validate this name. These communities can be interpreted as a transition to zonal woody vegetation.

3. lauretosum nobilis subass. nova, Rels 54–57 in Suppl. material 2: Table S1; holotypus relevé 55 in Suppl. material 2: Table S1. This subassociation occurs in more thermophilous and relatively xeric areas and it is differentiated by high cover values of Quercus ilex and Laurus nobilis. Similar communities were previously described by Bertacchi and Lombardi (2016) sub Fraxino-Quercetum roboris facies with Laurus nobilis.

4. sambucetosum nigrae subass. nova, Rels 58–75 in Suppl. material 2: Table S1; holotypus relevé 63 in Suppl. material 2: Table S1. This subassociation is defined by a group of relevés that underlines the transition with the woods of the Crataego-Quercion alliance (Querco-Fagetea). This evidence is highlighted by several subnitrophilous species such as Sambucus nigra, Chaerophyllum temulum, Geum urbanum, Arctium lappa, Galium aparine and less water-demanding species like Fraxinus ornus and Quercus cerris. In this subassociation we found a decrease in the number of species belonging to the class Alnetea and an increase for the species of the class Alno-Populetea.

ASPARAGO TENUIFOLII-CARPINETUM BETULI Arrigoni 1997. Holotypus: Table 2, rel. 90 in Arrigoni (1997) – Mesophylous floodplain forests dominated by Carpinus betulus (Table 5)

Table 5

. Asparago tenuifolii-Carpinetum betulii Arrigoni 1997. Species reported by the original authors as diagnostic are written in italics.

Rel. n° 1 2 3
Phi Asparago tenuifolii-Carpinetum betuli
87 Carpinus betulus L. 4 4 4
97 Anemonoides nemorosa (L.) Holub 2 2 +
94 Corylus avellana L. + 1 1
99.2 Ilex aquifolium L. + + r
91.7 Malus sylvestris (L.) Mill. 1 1 +
92.2 Lonicera etrusca Santi + + 1
99.2 Polygonatum multiflorum (L.) All. + r 1
78.1 Asparagus tenuifolius Lam. . + +
73.5 Crataegus laevigata (Poir.) DC. + r .
61.9 Dioscorea communis (L.) Caddick & Wilkin . r +
77.1 Festuca heterophylla Lam. + r .
76.2 Mespilus germanica L. + r .
75.3 Rubus hirtus Waldst. & Kit. group + + .
75.2 Sorbus torminalis (L.) Crantz r r .
Asarum europaeum L. r . .
Crataego-Quercion, Quercetalia pubescenti-petraeae, Querco-Fagetea
Physospermum cornubiense (L.) DC. + r r
Quercus robur L. subsp. robur 4 4 4
Hedera helix L. subsp. helix 2 2 3
Euonymus europaeus L. . r +
Viola reichenbachiana Jord. ex Boreau r r .
Luzula pilosa (L.) Willd. + + .
Prunus avium L. subsp. avium . . r
Crataegus monogyna Jacq. . . 1
Acer campestre L. . + .
Pteridium aquilinum (L.) Kuhn subsp. aquilinum . . r
Other species
Ruscus aculeatus L. 1 1 1
Arisarum proboscideum (L.) Savi + . r
Alnus glutinosa (L.) Gaertn. . 2 1
Lathraea clandestina L. + r .
Arum italicum Mill. subsp. italicum . . r
Pyrus communis L. . . 2
Frangula alnus Mill. subsp. alnus + . .
Myosotis sicula Guss. . r .
Cornus sanguinea L. . . +
Dactylis glomerata L. s.l. . r .
Quercus petraea (Matt.) Liebl. subsp. petraea 1 . .
Castanea sativa Mill. r . .
Ligustrum lucidum W.T.Aiton . . 1
Euphorbia dulcis L. . . r
Holcus mollis L. subsp. mollis . r .
Loncomelos pyramidale (L.) Raf. . . r
Neottia nidus-avis (L.) Rich. . . r
Viola riviniana Rchb. . . r

Diagnostic species : Carpinus betulus, Anemonoides nemorosa, Asparagus tenuifolius, Corylus avellana, Crataegus laevigata, Polygonatum multiflorum, Dioscorea communis, Ilex aquifolium, Lonicera etrusca, Rubus hirtus, Sorbus torminalis, Asarum europaeum.

Dominant trees : Carpinus betulus, Quercus robur.

Ecology and Chorology : Mesophilous woods can be found in sites where submersion is very occasional. This is a community with a more evident C-European character, highlighted by the lowest percentage of Mediterranean species and the highest percentages of European and European-Mediterranean ones (Fig. 2). According to the EIV analysis (Fig. 1), the woods dominated by Carpinus betulus show ecological requirements similar to those of Quercus robur communities, except for a slightly preference for lower temperature.

Distribution in Tuscany (Fig. 3) : Cerbaie (Arrigoni 1997, 1998).

Syntaxonomy : The association Asparago tenuifolii-Carpinetum betulii was described by Arrigoni (1997) for the Cerbaie hills, in north-western Tuscany. Arrigoni (1997; Table 2) reported two different associations: Polygonato multiflori-Quercetum roboris Sartori 1984 (Sartori 1984; Brullo and Spampinato 1999) and the new association Asparago tenuifolii-Carpinetum betulii (Arrigoni 1997). According to our analysis, and also to some important floristic differences (i.e. the lack of Convallaria majalis, diagnostic for Polygonato-Quercetum roboris, see Brullo and Spampinato 1999), the totality of the relevés in question can be assigned to the association Asparago tenuifolii-Carpinetum betulii and the association Polygonato-Quercetum roboris is to be excluded from Tuscany. These forests were found in the small plain sites of the lower parts of the Cerbaie hills. This association was assigned to the Crataego-Quercion and could be considered the wettest association of this alliance.

PERIPLOCO GRAECAE-ULMETUM MINORIS Vagge et Biondi, 1999. Holotypus: table 20, rel. 3 in Vagge and Biondi (1999) – Elm interdune groves (Table 6)

Table 6.

Periploco graecae-Ulmetum minoris Vagge et Biondi, 1999. Species reported by the original authors as diagnostic are written in italics.

Rel. n° 1 2 3 4 5 6 7
Phi Periploco graecae-Ulmetum minoris
Ulmus minor Mill. subsp. minor 4 5 5 4 5 4 5
Periploca graeca L. . 3 + . . 3 2
82.2 Rhamnus alaternus L. subsp. alaternus r . + r 1 . +
69.5 Smilax aspera L. r 1 + + . 2 1
68 Rubia peregrina L. + 1 + 2 1 1
66.3 Rosa sempervirens L. . . 1 + 1 . +
59.2 Phillyrea angustifolia L. . . . 2 1 . 1
Lauro-Ulmion, Rubio-Ulmetalia, Alno-Populetea
Rubus ulmifolius Schott 3 3 . + 1 3 2
Hedera helix L. subsp. helix r 3 . . 4 2 2
Brachypodium sylvaticum (Huds.) P.Beauv. subsp. sylvaticum + . . + 3 . +
Fraxinus angustifolia Vahl subsp. oxycarpa (M.Bieb. ex Willd.) Franco & Rocha Afonso 3 . . . . . +
Alnus glutinosa (L.) Gaertn. . 1 . . . .
Euonymus europaeus L. . . . . 1 . .
Laurus nobilis L. . . . . 1 . .
Dioscorea communis (L.) Caddick & Wilkin . . . + . . .
Chamaeiris foetidissima (L.) Medik. . . . r . . .
Quercetea ilicis . . . . . . .
Asparagus acutifolius L. . . . . + . 1
Ruscus aculeatus L. . . . . + . +
Pistacia lentiscus L. . . . 3 + . .
Viola alba Besser subsp. dehnhardtii (Ten.) W. Becker r . . . . . .
Myrtus communis L. . . . + . . .
Quercus ilex L. subsp. ilex . . . . 1 . .
Other species
Prunus spinosa L. subsp. spinosa r . 1 . + . +
Prunella vulgaris L. subsp. vulgaris + . . r . . .
Carex otrubae Podp. + . . + . . .
Robinia pseudoacacia L. . 2 . . . 2 .
Phragmites australis (Cav.) Trin. ex Steud. subsp. australis . + 1 . . . .
Pinus pinea L. . . . . . . 5
Lonicera etrusca Santi . . . . . . +
Crataegus monogyna Jacq. . . . . + . .
Ligustrum vulgare L. . . . . + . .
Potentilla reptans L. r . . . . . .
Rumex sanguineus L. r . . . . . .
Poa trivialis L. 2 . . . . . .
Carex distans L. r . . . . . .
Carex divulsa Stokes 2 . . . . . .
Oenanthe lachenalii C.C.Gmel. r . . . . . .
Plantago major L. r . . . . . .
Pyrus communis L. . . . . + . .
Vitis vinifera L. subsp. sylvestris (C.C. Gmel.) Hegi . + . . . . .
Mentha suaveolens Ehrh. r . . . . . .
Jacobaea erratica (Bertol.) Fourr. r . . . . . .
Phillyrea latifolia L. r . . . . . .
Rosa sp. r . . . . . .
Trifolium pratense L. subsp. pratense r . . . . . .
Catapodium rigidum (L.) C.E.Hubb. s.l. r . . . . . .
Limonium narbonense Mill. . . + . . . .
Sison amomum L. + . . . . . .
Tamarix sp. . . 3 . . . .
Taraxacum sp. r . . . . . .
Quercus pubescens Willd. . . . . . . +

Diagnostic species : Ulmus minor, Rhamnus alaternus, Smilax aspera, Rosa sempervirens, Rubia peregrina, Phillyrea angustifolia, Periploca graeca, Asparagus acutifolius.

Dominant tree : Ulmus minor.

Ecology and Chorology : Recently established woods, located on wet clay soils rich in nutrients. This is the most typical Mediterranean vegetation type among those investigated: here we found the highest percentage of Mediterranean species (Figure 2). According to the EIV analysis (Fig. 1), the groves with Ulmus minor show a negative relation to Soil Moisture (U).

Distribution in Tuscany (Fig. 3) : This vegetation type was found in the coastal systems of Tuscany. Tenuta di Migliarino (Vagge and Biondi 1999; Sani et al. 2011), Tombolo di Tirrenia (AA.VV. 2005), Tombolo di Cecina (Vagge and Biondi 1999).

Syntaxonomy : Ulmus minor groves of Tuscany were referred to the association Periploco graecae-Ulmetum minoris Vagge and Biondi 1999), described for the coastal areas of Tuscany from Migliarino to Cecina (Vagge and Biondi 1999).

DIOSCOREO COMMUNIS-POPULETUM NIGRAE Poldini et Vidali in Poldini, Sburlino et Vidali, 2017. Holotypus: rel. 1 of Table I of Poldini et Vidali in Poldini, Sburlino et Vidali, 2017 – Riparian wood with poplars and/or willows (Table 7)

Table 7.

Dioscoreo communis-Populion nigrae Poldini et Vidali in Poldini, Sburlino et Vidali, 2017. Rels 1–7: Dioscoreo communis-Populetum nigrae Poldini et Vidali in Poldini, Sburlino et Vidali, 2017 populetoum albae (Biondi, Vagge, Baldoni et Taffetani, 1999) Poldini, Vidali et Castello, 2020; rels 8–18: Dioscoreo communis-Populetum nigrae Poldini et Vidali in Poldini, Sburlino et Vidali, 2017 typicum.

Phi Rel. n. 1 2 3 4 5 6 7 8 9 11 12 10 13 14 15 16 17 18
Dioscoreo communis-Populetum nigrae
58.6 Salix alba L. . . . + . . 2 1 . . . . 5 4 3 4 4 5
49.3 Populus nigra L. 2 . . . . . 1 3 4 3 4 5 . . . 1 . .
Hedera helix L. s.l. 4 . 2 . 5 . . + . . . . . . . . . .
Brachypodium sylvaticum (Huds.) P.Beauv. subsp. sylvaticum . . . . + . . . . . + . . + . . . .
Ulmus minor Mill. subsp. minor . + 1 1 . . . . . . . . . . . . . .
Clematis vitalba L. . . + + + . . . . . . . . . . . . .
Robinia pseudoacacia L. . . . . . . . 1 . . . . . . . . . .
Corylus avellana L. . . . . 2 . . . . . . . . . . . . .
populetosum albae . . . . . . . . . . . . . . . . . .
44.2 Populus alba L. 4 3 4 4 5 4 2 2 + . . . . . . . + .
Populus canescens (Aiton) Sm. . . . . . . 2 . . . . . . . . . . .
Dioscoreo communis-Populion nigrae, Populetalia, Alno-Populetea
36.4 Cornus sanguinea L. . . 1 . . . . r . 3 3 3 3 1 + . . .
Rubus ulmifolius Schott . . 2 + 1 2 . . + 1 3 . . . . . . .
Lythrum salicaria L. . . . + . . + + + . + 2 . . . + . .
44.5 Rubus caesius L. . . . . . . + . . 2 . 4 2 + 5 + . .
Mentha aquatica L. subsp. aquatica . r . . . 1 . . + . + + . . . 3 . .
43 Humulus lupulus L. 1 . . . 1 . + . . . . . . 1 + + . .
42 Salix cinerea L. 1 . . . 1 . . . . . 1 . . + . 2 . .
Agrostis stolonifera L. + . . + . . . + . . . + 2 . . . . .
Alnus glutinosa (L.) Gaertn. 2 2 . 1 . . . . 3 . . . . . . . . .
Lycopus europaeus L. . . . + . 1 . . . . + . . . . 1 . .
Periploca graeca L. . + . + . . . . + . . . . . . . . .
Carex pendula Huds. + . . 1 + . . . . . . . . . . . r .
Fraxinus angustifolia Vahl subsp. oxycarpa (Willd.) Franco & Rocha Afonso . 3 2 1 . . . . . . . . . . . . . .
Galium palustre L. s.l. . . . + . 1 . . . . . + . . . . . .
Solanum dulcamara L. . . . + . . . . . . . . . . . + . .
Vitis vinifera L. . . + + . . . . . . . . . . . . . .
Carex otrubae Podp. . . . . . + . . . . . 1 . . . . . .
Carex sp. . . . . . . . . + . . + . . . . . .
Frangula alnus Mill. subsp. alnus . 1 . . . . . . + . . . . . . . . .
Sambucus nigra L. 1 . . . . . . . . . . . . . . . . .
Pastinaca sativa L. s.l. . . . . . . . . . . . . . . . + . .
Salix purpurea L. subsp. purpurea . . . . . . . + . . . . . . . . . .
Ludwigia palustris (L.) Elliott . . . . . + . . . . . . . . . . . .
Lythrum hyssopifolia L. . . . . . + . . . . . . . . . . . .
Carex remota L. . . . 1 . . . . . . . . . . . . . .
Carex sylvatica Huds. subsp. sylvatica . . . + . . . . . . . . . . . . . .
Other species
Crataegus monogyna Jacq. . . . . . . . . . 3 1 2 3 + + . . .
22.7 Phragmites australis (Cav.) Trin. ex Steud. subsp. australis . . . . . 2 . + . + + . . . . . 2 .
Urtica dioica L. subsp. dioica + . . . . . + . . . . . . 3 1 + . .
Limniris pseudacorus (L.) Fuss . r . + . . . r . . 1 . . . . + . .
43.9 Xanthium orientale L. subsp. italicum (Moretti) Greuter . . . . . . + . . . . r + . . . . 2
Lysimachia vulgaris L. . . . + . . . . + . . . + . . 1 . .
Laurus nobilis L. 1 . + . 1 . r . . . . . . . . . . .
35.3 Pulicaria dysenterica (L.) Bernh. . . . . . + . . . + . + . . . . . .
45.8 Galega officinalis L. . . . . . . . . . . . 1 3 . . + . .
Cirsium arvense (L.) Scop. . . . . . . . . . . . 1 2 + . . . .
Dipsacus fullonum L. . . . . . . . . . . . 1 + . . + . .
Galium aparine L. . . . . . . . . . . . 1 . 3 1 . . .
Hypericum perforatum L. . . . . . . . . . 1 . 1 2 . . . . .
Aristolochia clematitis L. . . . . . . + . . . . . . 2 1 . . .
Prunus spinosa L. subsp. spinosa . . + + + . . . . . . . . . . . . .
Acer campestre L. . . . . . . . . . . . . . 1 2 . . .
Acer negundo L. . . . . 1 . 1 . . . . . . . . . . .
Agrimonia eupatoria L. s.l. . . . . . . . . . . . + 2 . . . . .
Althaea officinalis L. . . . . . . . . . . . 1 . . + . . .
Euonymus europaeus L. 1 . 1 . . . . . . . . . . . . . . .
Amorpha fruticosa L. 1 . . . . . 1 . . . . . . . . . . .
Artemisia vulgaris L. . . . . . . 1 . . . . . . . . + . .
Cruciata laevipes Opiz . . . . . . . . . . . . . 1 + . . .
Daucus carota L. s.l. . . . . . . . . . . . + + . . . . .
Equisetum telmateia Ehrh. 1 . . . . . . . . r . . . . . . . .
Erigeron canadensis L. . . . + . . . . . . . . . . . + . .
Jacobaea aquatica (Hill) G.Gaertn., B.Mey. & Scherb. . . . . . . . . . . . . + . . + . .
Juncus inflexus L. subsp. inflexus . . . + . . + . . . . . . . . . . .
Ligustrum ovalifolium Hassk. 2 . . . + . . . . . . . . . . . . .
Phalaris arundinacea L. subsp. arundinacea . . . . . 3 2 . . . . . . . . . . .
Plantago major L. . . . + . . . . . . . + . . . . . .
Potentilla reptans L. . . . . . . . . . r . + . . . . . .
Quercus robur L. subsp. robur 1 . 2 . . . . . . . . . . . . . . .
Ruscus aculeatus L. . . 2 . . . . . . . . + . . . . . .
Sambucus ebulus L. . . . . . . . . . . . . . 2 2 . . .
Teucrium scordium L. s.l. . . . . . . . . . . + + . . . . . .
Torilis japonica (Houtt.) DC. . . . . . . . . . . . . . + + . . .
Typha latifolia L. . . . . . . . 1 . . . . . . . . + .
Verbena officinalis L. . . . . . . . . . . . r + . . . . .
Carex riparia Curtis . . . . . . . . . . . + . . . . . .
Artemisia verlotiorum Lamotte . . . . . . 2 . . . . . . . . . . .
Arundo donax L. . . . . . . . 2 . . . . . . . . . .
Brachypodium rupestre (Host) Roem. & Schult. . . . . . . . . . . . . 2 . . . . .
Bromus arvensis L. . . . . . . . . . . . . 2 . . . . .
Convolvulus sepium L. . . . . . 2 . . . . . . . . . . . .
Convolvulus arvensis L. . . . . . . . . . . . 2 . . . . . .
Ficus carica L. . . 2 . . . . . . . . . . . . . . .
Ligustrum vulgare L. . . 2 . . . . . . . . . . . . . . .
Paspalum distichum L. . . . . . . . . . . . . . . . . . 2
Thalictrum lucidum L. . . . . . . . . . . . . 2 . . . . .
Atriplex prostrata Boucher ex DC. . . . . . . . . . . . 1 . . . . . .
Lolium perenne L. . . . . . . . . . . . 1 . . . . . .
Lonicera caprifolium L. . . 1 . . . . . . . . . . . . . . .
Lotus corniculatus L. s.l. . . . . . . . . . . . 1 . . . . . .
Mentha pulegium L. subsp. pulegium . . . . . . . . . . . 1 . . . . . .
Mentha suaveolens Ehrh. . . . 1 . . . . . . . . . . . . . .
Quercus cerris L. . . . . . . . . . . . 1 . . . . . .
Quercus ilex L. subsp. ilex . . 1 . . . . . . . . . . . . . . .
Rubia peregrina L. . . 1 . . . . . . . . . . . . . . .
Rubus canescens DC. 1 . . . . . . . . . . . . . . . . .
Thalictrum flavum L. . . . . . . . . . . . 1 . . . . . .

Diagnostic species : Populus nigra, Salix alba, Rubus caesius, Xanthium orientale subsp. italicum, Salix cinerea, Humulus lupulus (phi >40), Populus alba, Ulmus minor, Galega officinalis, Pulicaria disenterica (phi > 30).

Dominant trees : Populus sp.pl., Salix alba.

Ecology and Chorology : This vegetation encompasses the woods that are located along riparian areas, placed in sites where the influence of rivers is marginal, like abandoned meanders or alluvial sectors in sedimentation. In the wet season, these communities are subject to even prolonged periods of submersion. This is a heterogeneous vegetation type, in which patches dominated by trees like poplars were found mixed with tall willow trees and shrubs. This vegetation is characterized by chorotypes with wide distribution (Eurosibiric, Cosmopolitan/Circumboreal) (Fig. 2). Unfortunately, from the conservation point of view, these vegetation types are rich in alien species. According to the EIV analysis (Fig. 1), the woods dominated by Populus sp.pl., and Salix alba are positively related to N (Nutrient).

Distribution in Tuscany (Fig. 3) : Macchia Lucchese (Arrigoni 1990), Selva di San Rossore (Gellini et al. 1986), Lago di Montepulciano and Lago di Chiusi (Lastrucci et al. 2014), Lago di Santa Luce (Bertacchi et al. 2005), Lago di San Floriano, Piana di Firenze and Valdarno superiore (Viciani and Raffaelli 2003). Many other sites with this vegetation type are probably present in other parts of Tuscany in addition to those reported here.

Syntaxonomy : In the past, similar coenoses have been attributed by many Italian authors (e.g. Pedrotti and Gafta 1996; Venanzoni and Gigante 2000; Pirone et al. 2003) to Populetum albae (Br.-Bl. 1931) Tchou or to Salici-Populetum nigrae (Tüxen, 1931) Meyer-Drees, 1936, mainly in relation to the dominant species. Very recently, some authors (Poldini et al., 2017; 2020) faced this issue and found that many Salici-Populetum nigrae communities sensu Auct. Ital. can be attributed to a recent established association named Dioscoreo communis-Populetum nigrae, which encompasses typical aspects dominated by Salix alba and Populus nigra, together with aspects dominated by Populus alba. Dioscoreo communis-Populetum nigrae is attributed to a new alliance, named Dioscoreo communis-Populion nigrae. Our relevés belonging to group 6 (Table 7) are not easy to classify, due to their heterogeneous floristic composition (common in these coenoses) but can be provisionally attributed to this syntaxon. The relevés 1–7, dominated by Populus alba, can be attributed to Dioscoreo communis-Populetum nigrae populetosum albae, while the relevés 8–18, dominated by Salix alba and Populus nigra, to the subass. typicum. These communities are floristically well characterized only for the tree layer, and from the successional and ecological points of view.

The spatial distribution scheme of these associations in the mosaic of the floodplain vegetation of Tuscany is shown in Figs 1 and 3 of Gellini et al. (1986). We add to these the groves dominated by Ulmus minor that can be found in Mediterranean coastal areas of central-southern Tuscany. Furthermore, we added the poplar and willow vegetation that, with the only exception of coastal areas, was also found in the areas around the large lakes of Chiusi and Montepulciano (Lastrucci et al. 2014).

Nature conservation remarks

Our results clarified the phytosociological classification of lowland woody communities in Tuscany, that is crucial to attribute them to the appropriate habitat type of conservation concern, listed in the Directive 92/43/EEC. The coenoses here described represent, in general, threatened plant communities that deserve strict conservation measures. Accordingly, based on Biondi et al. (2009, 2012) and the European Commission (2013), the floodplain woods of Tuscany can be referred to Natura 2000 habitats of Community interest as follows.

Alnus glutinosa and Fraxinus angustifolia subsp. oxycarpa dominated communities (Hydrocotylo vulgaris-Alnetum glutinosae; Limnirido pseudacori-Fraxinetum oxycarpae): habitat 91E0* – Alluvial forests with Alnus glutinosa and Fraxinus excelsior (Alno-Padion, Alnion incanae, Salicion albae).

Quercus robur and Ulmus minor dominated communities (Fraxino-Quercetum roboris; Periploco graecae-Ulmetum minoris): habitat 91F0 – Riparian mixed forests of Quercus robur, Ulmus laevis and Ulmus minor, Fraxinus excelsior or Fraxinus angustifolia, along the great rivers (Ulmenion minoris); when there is a codominance of Carpinus betulus, the habitat 9160 – Sub-Atlantic and medio-European oak or oak-hornbeam forests of the Carpinion betuli – can be used.

Carpinus betulus dominated communities (Asparago tenuifolii-Carpinetum betuli): habitat 9160 – Sub-Atlantic and medio-European oak or oak-hornbeam forests of the Carpinion betuli.

Populus spp. and Salix spp. dominated communities (Dioscoreo communis-Populetum nigrae): habitat 92A0 – Salix alba and Populus alba galleries.

Syntaxonomic scheme

ALNETEA GLUTINOSAE Br.-Bl. et Tüxen ex Westhoff, Dijk et Passchier, 1946

ALNETALIA GLUTINOSAE Tx., 1937

Alnion glutinosae Malcuit, 1929

Hydrocotylo vulgaris-Alnetum glutinosae Gellini, Pedrotti et Venanzoni, 1986

Frangulo alni-Fraxinion oxycarpae Poldini, Sburlino et Venanzoni in Biondi, Allegrezza, Casavecchia, Galdenzi, Gasparri, Pesaresi, Poldini, Sburlino, Vagge et Venanzoni, 2015

Limnirido pseudacori-Fraxinetum oxycarpae Gennai, Gabellini, Viciani, Venanzoni, Dell’Olmo, Giunti, Lucchesi, Monacci, Mugnai et Foggi ass. nova

ALNO GLUTINOSAE-POPULETEA ALBAE P. Fukarek et Fabijanić, 1968

POPULETALIA ALBAE Br.-Bl. ex Tchou, 1948

Carici remotae-Fraxinion oxycarpae Pedrotti ex Pedrotti, Biondi, Allegrezza et Casavecchia in Biondi, Allegrezza, Casavecchia, Galdenzi, Gasparri, Pesaresi, Vagge et Blasi, 2014

Fraxino oxycarpae-Quercetum roboris Gellini, Pedrotti et Venanzoni, 1986

typicum

carpinetosum betuli Gellini, Pedrotti et Venanzoni, 1986 ex Gennai, Gabellini, Viciani, Venanzoni, Dell’Olmo, Giunti, Lucchesi, Monacci, Mugnai et Foggi subass. nova

lauretosum nobilis Gennai, Gabellini, Viciani, Venanzoni, Dell’Olmo, Giunti, Lucchesi, Monacci, Mugnai et Foggi subass. nova

sambucetosum nigrae Gennai, Gabellini, Viciani, Venanzoni, Dell’Olmo, Giunti, Lucchesi, Monacci, Mugnai et Foggi subass. nova

Dioscoreo communis-Populion nigrae Poldini et Vidali in Poldini, Sburlino et Vidali, 2017

Dioscoreo communis-Populetum nigrae Poldini et Vidali in Poldini, Sburlino et Vidali, 2017 typicum

Dioscoreo communis-Populetum nigrae Poldini et Vidali in Poldini, Sburlino et Vidali, 2017 populetoum albae (Biondi, Vagge, Baldoni et Taffetani 1999) Poldini, Vidali et Castello, 2020

RUBIO PEREGRINAE–ULMETALIA MINORIS Biondi, Casavecchia, Gasparri et Pesaresi in Allegrezza, Casavecchia, Galdenzi, Gasparri, Pesaresi, Poldini, Sburlino, Vagge et Venanzoni, 2015

Lauro nobilis-Ulmion minoris Biondi, Casavecchia, Gasparri et Pesaresi in Allegrezza, Casavecchia, Galdenzi, Gasparri, Pesaresi, Poldini, Sburlino, Vagge et Venanzoni 2015

Periploco graecae-Ulmetum minoris Vagge et Biondi, 1999

QUERCO ROBORIS-FAGETEA SYLVATICAE Br.-Bl. et Vlieger in Vlieger, 1937

QUERCETALIA PUBESCENTIS Klika, 1933

Crataego laevigatae-Quercion cerridis Arrigoni 1997

Asparago tenuifolii-Carpinetum betuli Arrigoni 1997

Acknowledgements

Thanks are due to Simona Bacci for English language checking, and to three anonymous reviewers for their suggestions and comments that allow us to greatly improve the manuscript.

Funding

The research was funded by Regione Toscana “The floodplain woods of Tuscany: Distribution, Ecology, Characterization” (resp. prof. F. Lucchesi, Department of Architecture – DIDA, University of Florence).

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Appendixes

Appendix I – Literature concerning the floodplain vegetation of Tuscany from which we selected the relevés

AA.VV., 2005 – Piano di Gestione Forestale della “Tenuta di Tombolo” di proprietà del Comune di Pisa. Periodo di validità 2005 – 2014. D.R.E.AM. Italia. Poppi (AR). Relazione tecnica, 193 pagine.

Arrigoni P.V., 1990 – Flora e vegetazione della Macchia lucchese di Viareggio (Toscana). Webbia, 44(1): 1–62.

Arrigoni P.V., 1997 – Documenti per la carta della vegetazione delle Cerbaie (Toscana settentrionale). Parlatorea 2: 39–71.

Arrigoni P.V., Nardi E., Raffaelli M., 1985 – La vegetazione del Parco Naturale della Maremma (Toscana). Con carta in scala 1:25.000. Università degli Studi di Firenze. Dip. Biol. Veg. 39 pagine.

Bertacchi A., Bocci G., Lombardi T., Tomei P.E., 2005 – Le fitocenosi riparie della Toscana nord-occidentale. Il fiume Fine. Edizioni ETS. Pisa. 59 pagine.

Bertacchi A., Lombardi T., 2016 – I boschi di Coltano: aspetti storici, fisionomici e vegetazionali di un paesaggio forestale relitto nella pianura di Pisa (Toscana). Atti Soc. tosc. Sci. nat., Mem., Ser. B 122(2015): 111–122.

Coaro E., 1987 – Flora e vegetazione del Bosco dell’Ulivo (Parco di Migliarino, S. Rossore e Massaciuccoli). Quad. Mus. Stor. Nat. Livorno 8, Suppl. 1: 1–45.

Ferretti G., Lastrucci L., 2010. Lago di San Floriano, provincia di Grosseto. Rilievi inediti.

Foggi B., Selvi F., Viciani D., Bettini D., Gabellini A., 2000 – La vegetazione forestale del bacino del Fiume Cecina (Toscana centro-occidentale). Parlatorea 4: 39–73.

Gabellini A. & al., 2018 – Boschi planiziali di Toscana. Rilievi inediti.

Gellini R., Pedrotti F., Venanzoni R., 1986 – Le associazioni forestali ripariali e palustri della Selva di San Rossore (Pisa). Doc. Phytosoc., n.s. 10(2): 27–41.

Gennai M., Lastrucci L., 2013 – Corso del Fiume Arno nella città di Firenze. Rilievi inediti.

Lastrucci L., Bonari G., Angiolini C., Casini F., Giallonardo T., Gigante D., Landi M., Landucci F., Venanzoni R., Viciani D., 2014 – Vegetation of Lakes Chiusi and Montepulciano (Siena, central Italy): updated knowledge and new discoveries. Plant Sociology 51(2): 29–55.

Lastrucci L., Dell'Olmo L., Foggi B., Massi L., Nuccio C., Vicenti C., Viciani D., 2017 – Contribution to the knowledge of the vegetation of the Lake Massaciuccoli (northern Tuscany, Italy). Plant Sociology 54(1): 67–87.

Lastrucci L., Valentini E., Dell'Olmo L., Vietina B., Foggi B., 2016 – Hygrophilous vegetation and habitats of conservation interest in the area of the Lake Porta (Tuscany, Central Italy). Atti Soc. tosc. Sci. nat., Mem., Ser. B 122(2015): 131–146.

Sani A., Monacci F., Trimarchi S., Tomei P.E., 2011 – La vegetazione della Tenuta di Migliarino. Inter Nos 2 (2010): 5–72.

Tomei P.E., Bertacchi A., Sani A., Consiglio M., 2004 – La vegetazione della Tenuta di San Rossore. Note esplicative della Carta della Vegetazione di San Rossore 1.10.000. Ente Parco Regionale Migliarino San Rossore Massaciuccoli. 67pagine.

Tomei P.E., Cenni M., 1986 – Il Bosco di Chiusi e la Paduletta di Ramone (PT): note floristiche e vegetazionali. Quad. Mus. Stor. Nat. Livorno 7: 55–79.

Vagge I., Biondi E., 1999 – La vegetazione delle coste sabbiose del Tirreno settentrionale italiano. Fitosociologia 36(2): 61–95.

Viciani D., Gabellini A., 2012 – La vegetazione relitta a dominanza di farnia (Quercus robur L.) del Bosco dei Renacci e di altri popolamenti simili presenti nella porzione basale della conca intermontana del Valdarno Superiore tra Incisa e San Giovanni Valdarno (Toscana). Atti Soc. tosc. Sci. nat., Mem., Ser. B 118(2011): 57–64.

Appendix II – Correspondence between the relevé numbers in our tables and original bibliographic references

Table 2 – Rels. 4, 5, 6 = rels. 16, 10, 12 in Lastrucci L. & al. (2016); Rels. 1, 11, 21 = rels. 46, 45, 50 from Tab. 12 in Sani A. & al. (2011); Rels. 2, 3, 7, 8, 12 = rels. 144, 32, 54, 109, 142 from Tab. 14 in Arrigoni P.V. (1990); Rels. 9, 13, 18, 23, 24 = rels. 27, 26, 25, 24, 28 from Tab. 1 in Gellini & al. (1986); Rels. 14, 25 = rels. 8, 7 from Tab. 3 in Gellini & al. (1986); Rels. 10, 16, 17, 20 = rels. 21, 57, 59, 63 from Tab. 6 in Tomei & al. (2004); Rels. 15, 22 unpublished in Gabellini A. & al. (2018); Rels. 26, 27, 28, 29, 30 = rels. 73, 74, 75, 76, 84 from Tab. 3 in Arrigoni P.V. (1997).

Table 3 – Rels. 45 = rels. 38 from Tab. 5 in Tomei & al. (2004); Rels. 33, 34, 35, 39, 40 = rels. 32, 80, 83, 29, 67 from Tab. 11 in Sani A. & al. (2011); Rels. 6, 16, 17, 18, 19, 20, 21, 22, 23, 28, 29, 31 = rels. 18, 9, 42, 49, 63, 65, 68, 85, 95, 33, 34, 47 from Tab. 10 in Sani A. & al. (2011); Rels. 1, 2, 3, 30, 47 = unpublished rels. 10, 11,12, 35 by Gabellini A. in Piano di Gestione Forestale della “Tenuta di Tombolo”; Rels. 32, 36, 37, 38, 41, 42, 43, 44, 46 = rels. 80, 91, 103, 124, 79, 97, 123, 81, 94 from Tab. 15 in Arrigoni P.V. (1990); Rels. 11, 15 = rels. X023, X026 from Tab. 16 in Foggi & al. (2000); Rels. 4, 5 = rels. 3, 36 from Tab. 8 in Coaro E. (1987); Rels. 7, 8, 10, 12, 13, 14 = rels. 3, 84, 53, 55, 20, 99 from Tab. 3. in Tomei & al. (2004); Rels. 9, 24, 25, 26, 27 = rels. 4, 2, 6, 5, 3 from Tab. 3. in Gellini & al. (1986).

Suppl. material 2: Table S1 – Rels. 34, 36, 37, 38, 39, 40, 41, 42, 43, 47, 49, 50 = rels. 1, 16, 11, 15, 12, 14, 10, 22, 9, 18, 20, 19 from Tab. 3 in Gellini & al. (1986); Rels. 7, 51, 88, 39, 56, 6, 2, 11, 17, 1, 9, 34, 33, 14, 43, 4 = rels. 6, 49, 43, 14, 17, 2, 11, 7, 1, 33, 39, 56, 51, 88, 34 from Tab. 1 in Tomei & al. (2004); Rels. 33, 44 = rels. 14, 42 from Tab. 3 in Tomei & al. (2004); Rels. 9, 25, 26, 32, 47 = rels. 18, 12, 20, 10, 30 from Tab. 4 in Coaro E. (1987); Rels. 39 = rels. 42 from Tab. 5 in Coaro E. (1987); Rels. 35, 36, 37, 38, 39 = rels. 11, 13, 21, 33, 2 from Tab. 8 in Coaro E. (1987); Rels. 6, 51, 52= rels. 8, 44, 36 from Tab. 8 in Sani A. & al. (2011); Rels. 54, 55, 57 = rels. 39, 43, 40 from Tab. 9 in Sani A. & al. (2011); Rels. 18, 19, 22, 23, 56 = rels. 9, 13, 7, 8, 6 in patch n. 255, patch n. 257, patch n. 262, patch n. 262, patch n. 262 in AA.VV. (2005); Rels. 69, 75, 61, 62 = rels. unpublished in Gabellini A. & al. (2018); Rels. 7, 13, 14 = rels. 1, 4, 10 from Tab. 1 in Bertacchi A. & Lombardi T. (2016); Rels. 15, 31 = rels. 17, 4 from Tab. 2 in Tomei P.E. & Cenni M. (1986); Rels. 17 = rels. 53 from Tab. 15 in Arrigoni P.V. (1990); Rels. 58, 59, 60, 63, 64, 65, 66, 67, 68, 70, 71 = rels. G007, V003, V004, G008, V002, G001, G009, G010, G006, V001, V005 from Tab. 1 in Gabellini A. & Viciani D. (2012); Rels. 16 = rels. 63 from Tab. 9 in Arrigoni P.V. (1997); Rels. 30 = rels. 94 from Tab. 2 in Arrigoni P.V. (1997).

Table 6 – Rels. 2, 6 = rel. 220, 210 Tab. 20 from Arrigoni P.V. & al. (1985). Rel. 4, 5, 7, 8, 9 = rels. 2, 4, 5, 1, 3 Tab. 20 from Vagge I., Biondi E. (1999).

Table 7 – Rels. 1, 5 unpublished from Gabellini A. & al. (2018). Rels. 10–16 = rels. 7–11, 13, 14 Tab. 10 from Lastrucci L. & al. (2014). Rels. 8, 17 = rels. 9, 4 Tab. 3 from Bertacchi A. & al. (2005). Rels. 7, 18 = rels. 37, 35 unpublished from Gennai M. & Lastrucci L. (2013). Rels. 6 = rels. 8 unpublished from Ferretti G. & Lastrucci L. (2010). Rels. 2, 9 = rels. 59, 126 Tab. 15 from Arrigoni P.V. (1990). Rel. 3 = rel. 5, from AA. VV (2005). Rel. 4 = rel. 30 Tab. 4 from Gellini & al. (1986).

Appendix III – Sporadic species

Table 2Aegopodium podagraria L. (22), Anemonoides nemorosa (L.) Holub (22), Aristolochia rotunda L. s.l. (14, 25), Athyrium filix-femina (L.) Roth (15), Aulacomnium palustre (Hedw.) Schwägr. (22), Baldellia ranunculoides (L.) Parl. (13), Bidens tripartitus L. s.l. (22), Calypogeia sphagnicola (Arn. & Perss.) Warnst. & Loeske (22), Carex distans L. (7), Carex divulsa Stokes (10), Carex flacca Schreb. s.l. (20), Carex punctata Gaudin (9), Carex sp. (12), Chamaesyce peplis (L.) Prokh. (18, 23), Circaea lutetiana L. subsp. lutetiana (15, 22), Corylus avellana L. (15), Dioscorea communis (L.) Caddick & Wilkin (15), Emerus major Mill. s.l. (4), Euonymus japonicus Thunb. (5), Fallopia convolvulus (L.) Á. Löve (29), Galium aparine L. (29, 4), Galium debile Desv. (13, 18), Galium mollugo L. subsp. erectum Syme (15, 29), Geum rivale L. (35), Geum urbanum L. (26), Gratiola fromficinalis L. (19, 13), Holcus lanatus L. (19), Iris foetidissima L. (16), Juncus anceps Laharpe (12), Juncus articulatus L. (13), Juniperus oxycedrus L. subsp. macrocarpa (Sibth. & Sm.) Neilr. (3), Ligustrum ovalifolium Hassk. (15), Ligustrum sinense Lour. (5, 6), Ligustrum vulgare L. (6, 17), Lysimachia nummularia L. (22), Malus sylvestris (L.) Mill. (21), Moehringia trinervia (L.) Clairv. (18), Myosotis laxa Lehm. (27), Myosotis scorpioides L. subsp. scorpioides (30), Oenanthe pimpinelloides L. (15), Persicaria lapathifolia (L.) Delarbre subsp. lapathifolia (13, 23), Platanus hispanica Mill. ex Münchh. (4), Poa annua L. (18), Poa sylvicola Guss. (15), Populus canescens (Aiton) Sm. (4), Potentilla indica (Andrews) Th. Wolf (4), Potentilla sterilis (L.) Garcke (26), Prunella laciniata (L.) L. (30), Prunella vulgaris L. subsp. vulgaris (9, 14), Prunus laurocerasus L. (15), Prunus sp. (5), Pteridium aquilinum (L.) Kuhn subsp. aquilinum (30), Pyracantha coccinea M. Roem. (2), Pyrus communis L. (26), Ranunculus flammula L. (13), Ranunculus lanuginosus L. (15, 4), Ranunculus ophioglossifolius Vill. (9), Ranunculus sardous Crantz subsp. sardous (16, 17), Rhamnus cathartica L. (2), Rumex conglomeratus Murray (15, 22), Rumex sanguineus L. (13, 23), Silene latifolia Poir. subsp. alba (Mill.) Greuter & Burdet (29), Solanum nigrum L. (12, 22), Stellaria aquatica (L.) Scop. (4), Stellaria media (L.) Vill. subsp. media (14), Symphytum tuberosum L. subsp. angustifolium (A. Kern.) Nyman (29), Thyselium palustre (L.) Raf. (19), Veronica montana L. (17), Veronica persica Poir. (14), Veronica serpyllifolia L. subsp. serpyllifolia (14, 25), Vinca minor L. (15), Viola reichenbachiana Jord. ex Boreau (22,14), Vitis sp. (4).

Table 3Ajuga reptans L. (5), Alisma lanceolatum With. (38), Althaea cannabina L. (6), Amorpha fruticosa L. (28, 31), Aristolochia rotunda L. (26), Asparagus acutifolius L. (30), Bidens frondosa L. (42), Bidens tripartitus L. s.l. (47), Blackstonia perfoliate (L.) Huds. s.l. (12), Carex canescens L. (24), Carex caryophyllea Latourr. (10, 14), Carex divulsa Stokes (27, 33), Carex flacca Schreb. s.l. (10, 30), Carex strigosa Huds. (30, 47), Centaurium pulchellum (Sw.) Druce subsp. pulchellum (12), Centaurium tenuiflorum (Hfromfmanns. & Link) Fritsch subsp. tenuiflorum (14), Cerastium semidecandrum L. (45), Crepis bellidifolia Loisel. (14), Dittrichia viscosa (L.) Greuter s.l. (33; 57), Elymus repens (L.) Gould subsp. repens (2), Equisetum arvense L. subsp. arvense (25), Equisetum telmateja Ehrh. (10, 12), Eupatorium cannabinum L. subsp. cannabinum (6), Falloppia convolvulus (L.) A. Love (3, 47), Fraxinus ornus L. subsp. ornus (34), Galium debile Desv. (9), Galium palustre L. subsp. elongatum (C.Presl) Lange (11), Geum urbanum L. (47), Gratiola fromficinalis L. (20, 39), Holcus lanatus L. (12, 45), Hypericum androsaemum L. (26), Hypericum tetrapterum Fr. (24), Isolepis setacea (L.)R.Br. (28), Juncus conglomeratus L. (12), Lemna minor L. (15), Lonicera caprifolium L. (30), Lonicera caprifolium L. (30), Lotus corniculatus L. s.l. (4), Luzula forsteri (Sm.) DC. (45), Lysimachia arvensis (L.) U. Manns & Anderb. s.l. (26), Lysimachia foemina (Mill.) U. Manns & Anderb. s.l. (11), Lythrum hyssopifolia L. (11), Malus sylvestris (L.) Mill. (18, 23), Melica uniflora Retz. (30), Moehiringia trinervia (L.) Clairv. (24), Myosotis laxa Lehm. (45), Myosotis sylvatica Hfromfm. subsp. sylvatica (45), Phytolacca americana L. (24), Phalaris arundinacea L. subsp. arundinacea (20), Phytolacca americana L. (24), Pinus pinea L. (36), Poa annua L. (27), Poa palustris L. (45), Poa sterilis L. (38, 43), Potentilla sterilis (L.) Garcke (38, 43), Pulicaria dysenterica (L.) Berhn. (25), Pulicaria odora (L.) Rchb. (15), Ranunculus bulbosus L. subsp. aleae (Willk.) Rouy & Foucaud (11), Ranunculus flammula L. (24), Ranunculus ophioglossifolius Vill. (15), Ranunculus velutinus Ten. (4, 3), Ranunculus sp. (24), Rosa sempervirens L. (30), Rubia peregrina L. s.l. (30), Rubus sp. (19, 21), Rumex hydrolapathum Huds. (33), Sambucus nigra L. (25), Schenkia spicata (L.) G. Mans. (12), Scirpoides holoschoenus (L.) Soják (4), Scrophularia nodosa L. (24), Silene flos-cuculi (L.) Clairv. (24), Solanum dulcamara L. (37), Thalictrum lucidum L. (15), Trifolium fragiferum L. subsp. fragiferum (5), Triglochin bulbosum L. subsp. barrelieri (Loisel.) Rouy (4), Tripolium pannonicum (Jacq.) Dobrocz. subsp. pannonicum (4), Veronica anagallis-aquatica L. subsp. anagallis-aquatica (1524), Veronica montana L. (), Veronica serpyllifolia L. subsp. serpyllifolia (24, 27).

Suppl. material 2: Table S1 – Ailanthus altissima (Mill.) Swingle (65), Alliaria petiolata (M. Bieb.) Cavara & Grande (58), Amorpha fruticosa L. (31), Anemoides nemorosa (L.) Holub (16, 30), Anthoxanthum odoratum L. s.l. (62, 45), Arisarum proboscideum (L.) Savi (4), Asparagus tenuifolius Lam. (30), Asperula laevigata L. (61), Bellevalia romana (L.) Sweet (15), Buglossoides purpurocaerulea (L.) I.M. Johnst. (69), Calluna vulgaris (L.) Hull (15), Carex flacca Schreb. s.l. (6), Carex flacca Schreb. subsp. flacca (39, 40), Carex leporina L. (37), Carex pallescens L. (41), Carex punctata Gaudin (36), Carex riparia Curtis (43), Castanea sativa Mill. (62), Cerastium brachypetalum Desp. ex Pers. s.l. (74), Chamaesyce peplis (L.) Prokh. (36), Clematis flammula L. (3), Clinopodium nepeta (L.) Kuntze s.l. (11), Clinopodium vulgare L. s.l. (71), Clinopodium vulgare L. subsp. vulgare (72), Cruciata glabra (L.) Ehrend. s.l. (6), Cyclamen hederifolium Aiton subsp. hederifolium (42), Cytisus scoparius (L.) Link subsp. scoparius (70), Dactylis glomerata L. s.l. (70, 16), Danthonia decumbens (L.) DC. subsp. decumbens (61), Dittrichia viscosa (L.) Greuter s.l. (8), Dryopteris filix-mas (L.) Schott (41), Equisetum palustre L. (47), Erica arborea L. (15), Erigeron canadensis L. (37), Euphorbia cyparissias L. (72), Fallopia convolvulus (L.) Á. Löve (56), Ficus carica L. (56), Fragaria vesca L. subsp. vesca (2, 26), Gagea lutea (L.) Ker Gawl. (68), Galega officinalis L. (70), Galium mollugo L. subsp. erectum Syme (69), Geranium molle L. (26), Geranium purpureum Vill. (41, 36), Geranium robertianum L. (21, 29), Geranium sylvaticum L. (10), Gladiolus palustris Gaudin (15), Glechoma hederacea L. (32), Hottonia palustris L. (43), Hypericum montanum L. (75, 39), Hypochaeris glabra L. (8), Ilex aquifolium L. (30), Iris graminea L. (68), Iris pseudacorus L. (13), Juglans regia L. (68), Juncus conglomeratus L. (43), Juncus inflexus L. (42), Juniperus communis L. (31), Lamium maculatum L. (58), Lapsana communis L. subsp. communis (62), Lathyrus sp. (43), Lysimachia arvensis (L.) U. Manns & Anderb. s.l. (45), Lysimachia vulgaris (L.) U. Manns & Anderb. s.l. (37, 43), Melissa officinalis L. s.l. (42), Molinia caerulea (L.) Moench s.l. (15), Muscari comosum (L.) Mill. (67, 68), Myosotis scorpioides L. subsp. scorpioides (43), Narcissus tazetta L. s.l. (15), Oenanthe pimpinelloides L. (62, 15), Ornithogalum umbellatum L. (67), Orobanche purpurea Jacq. (35), Parietaria officinalis L. (61), Phillyrea angustifolia L. (46), Poa pratensis L. (45, 48), Polygonatum multiflorum (L.) All. (30), Polygonatum odoratum (Mill.) Druce (42), Polystichum setiferum (Forssk.) T. Moore ex Woyn. (62), Potentilla erecta (L.) Raeusch. (42), Prunus domestica L. s.l. (75), Pyracantha coccinea M. Roem. (19), Quercus petraea (Matt.) Liebl. subsp. petraea (62), Quercus pubescens Willd. subsp. pubescens (69), Ranunculus lanuginosus L. (68), Ranunculus polyanthemos L. (16), Ranunculus sardous Crantz subsp. sardous (45), Rumex acetosella L. s.l. (15), Rumex hydrolapathum Huds. (8, 66), Salix alba L. (61), Samolus valerandi L. (1), Schedonorus arundinaceus (Schreb.) Dumort. s.l. (12), Scilla bifolia L. (68), Scirpoides holoschoenus (L.) Soják (6), Sedum cepaea L. (60, 64), Serratula tinctoria L. s.l. (15), Silene dioica (L.) Clairv. (61), Silene dioica (L.) Clairv. (61), Solanum nigrum L. (61), Sorbus domestica L. (62), Stachys officinalis (L.) Trevis. (42, 3), Symphyotrichum squamatum (Spreng.) G.L. Nesom (26), Ulex europaeus L. subsp. europaeus (62), Verbena officinalis L. (16), Veronica chamaedrys L. subsp. chamaedrys (5, 58), Veronica hederifolia L. subsp. hederifolia (67), Viburnum tinus L. subsp. tinus (69, 71), Vincetoxicum hirundinaria Medik. subsp. hirundinaria (15), Viola alba Besser subsp. dehnhardtii (Ten.) W. Becker (50, 66), Viola odorata L. (15, 75).

Table 7Alisma plantago-aquatica L. (17), Aquilegia vulgaris auct. Fl. Ital. (10), Arctium nemorosum Lej. (16), Bidens frondosus L. (18), Carex canescens L. (4), Chaerophyllum temulum L. (1), Chara sp. (11), Cirsium vulgare (Savi) Ten. (16), Cuscuta campestris Yunck. (18), Datura stramonium L. subsp. stramonium (7), Epilobium tetragonum L. s.l. (10), Eupatorium cannabinum L. subsp. cannabinum (16), Fragaria vesca L. subsp. vesca (10), Galium rotundifolium L. subsp. rotundifolium (10), Hypericum montanum L. (5), Juncus effusus L. subsp. effusus (7), Juncus maritimus Lam. (4), Lactuca serriola L. (13), Ligustrum lucidum W.T. Aiton (5), Lilium bulbiferum L. subsp. croceum (Chaix) Jan (10), Liquidambar styraciflua L. (6), Melissa officinalis L. s.l. (6), Mentha arvensis L. (11), Persicaria hydropiper (L.) Delarbre (16), Persicaria lapathifolia (L.) Delarbre subsp. lapathifolia (18), Phytolacca americana L. (1), Pinus pinaster Aiton subsp. pinaster (9), Prunus cerasifera Ehrh. (5), Ranunculus repens L. (4), Ranunculus serpens Schrank subsp. nemorosus (DC.) G. López (10), Rosa canina L. (13), Samolus valerandi L. (4), Saponaria officinalis L. (17), Scutellaria galericulata L. (4), Smilax aspera L. (3), Stellaria media (L.) Vill. subsp. media (18), Symphyotrichum squamatum (Spreng.) G.L. Nesom (10), Tordylium maximum L. (10), Veronica anagallis-aquatica L. subsp. anagallis-aquatica (17).