The Stipa dasyvaginata subsp. apenninicola mountain grasslands of the Velino Massif (central Italy)

The Stipa dasyvaginata subsp. apenninicola mountain grasslands of the Velino Massif (central Italy)* Claudia Zitarelli1, Andrea De Toma1, Flavio Marzialetti2,3, Maurizio Cutini1,3 1 Department of Science, Roma Tre University, Rome, Italy 2 Department of Agricultural Sciences, University of Sassari, Sassari, Italy 3 National Biodiversity Future Center (NBFC), Palermo, Italy Corresponding author: Claudia Zitarelli ([email protected]) * Topical Collection: “Vegetation classification: from classic to numeric approaches”. Edited by Bianca Ott Andrade, Claudia Angiolini, Lorenzo Lazzaro. Academic editor: Federico Fernández-González♦ Received 30 July 2025♦ Accepted 31 October 2025♦ Published 14 November 2025 Abstract The central Italian Stipa mountain grasslands are a fascinating vegetation type of notable biogeographic interest, having remained along the Apennine chain since the last Pleistocene glacial phases. The present study focuses on the phytosociological description of grasslands dominated by Stipa dasyvaginata subsp. apenninicola, an endemic Apennine species. We sampled twenty-five plots in the Velino Massif and surrounding areas facing the northwestern side of the Fucino Basin and performed a cluster analysis combining our dataset with all published relevés of Stipa dasyvaginata subsp. apenninicola dominated communities in the Italian peninsula. The floristic and environmental differences between our dataset and the other communities led us to describe the new association Sileno bellidifoliae-Stipetum apenninicolae (class Helianthemo cani-Seslerietea nitidae), showing a xerophilous ecology. We further analyzed the variance in environmental data within our dataset, as well as its biological and chorological spectra. The peculiar floristic and ecological characteristics of these steppic grasslands highlight the need of preserving this type of vegetation, which may represent a valuable example contributing to the understanding of the formation of the contemporary vegetation landscape. Keywords Central Apennines, dry grasslands, phytosociology, syntaxonomy Introduction The Eurasian Steppe is one of the largest continuous terrestrial natural habitats, spanning a wide area of the Palearctic region from central and eastern Europe to northern China (Willner et al. 2021). This habitat is mainly characterized by a dry continental climate with low annual precipitation and pronounced temperature seasonality (Török et al. 2020). It is also marked by high biodiversity and zonal vegetation typical of temperate grasslands and forest-steppes (Pokorný et al. 2015; Wesche et al. 2016). During the glacial phases of the Pleistocene, steppes were widespread across most of central and northern Eurasia but retreated with the beginning of the Holocene, whose more humid climate led to the expansion of forests (Hejcman et al. 2013; Dengler et al. 2014; Feurdean et al. 2018; Willner et al. 2021). However, under pressure from herbivores and agricultural societies during the mid-Holocene climatic optimum (Pokorný et al. 2015), many species retreated to refugial areas, where they found favorable environmental conditions and suitable habitats, leading to the development of central European extrazonal steppes, as in the dry valleys of the Alps, the Pannonian Basin, the Vegetation Ecology and Diversity 62 (2025) e167266|DOI 10.3897/ved.167266 Copyright Claudia Zitarelli et al. This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited. Italian Society of Vegetation Science (SISV) Topical Collection Claudia Zitarelli et al.: The S. dasyvaginata subsp. apenninicola mountain grasslands of the Velino Massif2 Rhine Valley, northern and central Bohemia, Thuringia and the lower Oder Valley, southern Moravia, and Lower Austria (Kajtoch et al. 2016; Kirschner et al. 2020; Chytrý et al. 2022; Divíšek et al. 2022). Several species with ancient continuous distribution throughout Eurasia are currently considered either rare or with a highly fragmented distributional range. This is the case for several species of the genus Stipa, occurring in many dry valleys of the Alps and the Apennines (Becker 2010; Hensen et al. 2010). In the Italian peninsula, most of the species of the genus Stipa are found on calcareous substrates, preferably in semi-rocky habitats, where competition with other species is reduced (Pignatti 2017–2019). In the Apennines, three endemic species can be found: S. aquilana (Conti et al. 2007) and S. crassiculmis subsp. picentina (Spada et al. 2010), restricted to isolated patches, and S. dasyvaginata subsp. apenninicola, which is common in mountainous areas. This latter species shows a fragmented distribution along central and southern Apennines (Martinovský and Moraldo 1980; Moraldo 1983, 1986), reaching up to 2000 m a.s.l. from Marche Region to Calabria (Pignatti 2017–2019). In central Apennines, as well as in the Alps, grasslands dominated by Stipa capillata, a relict steppic species with a Eurasian distribution, can also be found (Tammaro 1984). A few studies on communities dominated by Stipa dasyvaginata subsp. apenninicola have been conducted in the dry valleys of the central Apennines (Pirone et al. 2001; Taffetani et al. 2004; Ballelli et al. 2006; Catorci et al. 2007), but none of them regarded the Fucino Basin. The intermontane basins of central Italy provide remarkably suitable areas for this vegetation, with an overall species composition which resembles that of the central European steppes and rich in glacial relicts, endemic and rare species (Petriccione 1993; Di Pietro and Catonica 1999; Pezzetta 2011; Cancellieri et al. 2024). In the Fucino Basin and surrounding mountains, the peculiar environmental and land-use conditions of mountain sites hindered the development of forest vegetation, thus enabling the persistence of fragmented populations of steppic species in glacial refugia, as well documented by the recent findings of Festuca valesiaca, Astragalus exscapus subsp. exscapus, Goniolimon tataricum subsp. italicum, Alyssum desertorum, Spiraea hypericifolia subsp. hypericifolia, and Adonis fucensis (Bartolucci and Conti 2016; Cancellieri et al. 2017; Filibeck et al. 2020; Conti and Bartolucci 2023; Conti et al. 2023). Here, the presence of Stipa dasyvaginata subsp. apenninicola has already been sporadically recorded (Cancellieri et al. 2020), but the grasslands dominated by this species have never been fully investigated. Given the biogeographical and ecological importance of these communities along with the lack of studies in the area, the main aim of the present investigation was the phytosociological analysis, as well as the coenological and chorological characterization, of the Stipa dasyvaginata subsp. apenninicola dominated grasslands in the Velino Massif. Materials and methods Study area The study area is located on the Velino Massif and surrounding areas facing the northwestern side of the Fucino Basin in Abruzzo, which is the largest tectonic intermontane basin in central Italy. It hosted an ancient lake before being completely drained at the end of 19th century (Petriccione 1993; Cavinato et al. 2002). The sampling occurred in seven limestone mountain areas, namely the slopes between Mount Rozza (2064 m a.s.l.) and Mount Sevice (2355 m a.s.l.), Mount Velino (2487 m a.s.l.), Mount Cafornia (2424 m a.s.l.), the access of Valle Majelama, the upper side of Forme, Mount Tino (1923 m a.s.l.), and the slopes near Collarmele (Fig. 1a). The area belongs to the sub-mediterranean bioclimate, with upper orotemperate thermotype and upper subhumid ombrotype (Pesaresi et al. 2017; Cutini et al. 2021). Furthermore, the basin is affected by the rain shadow effect caused by the northern and southwestern mountain massifs of the Sirente-Velino and Simbruini Mountains, which drive moist air upslope and thus cause a dry continental climate with low annual precipitation, winter frosts, and summer drought (Petriccione 1993; Pirone and Cutini 2002), as shown in the ombrothermic diagram below (Fig. 1b). Dataset and data analysis Twenty-five phytosociological relevés were carried out in summers 2007 and 2012 (7 relevés) and in summer 2023 (18 relevés). The preferential sampling was performed according to the Braun-Blanquet (1932) method, along south-facing rocky outcrops on calcareous soils ranging from 930 m to 1820 m a.s.l. Relevés ranged in size from 30 to 100 m2 and were located where Stipa dasyvaginata subsp. apenninicola dominated communities were found. During the sampling, we recorded the geographic coordinates of each plot and also collected environmental data on elevation, aspect, slope, rockiness, stoniness, and total vegetation cover. Taxonomic nomenclature follows Bartolucci et al. (2024), while life forms and chorotypes follow Pignatti (2017–2019). The syntaxonomical nomenclature follows Mucina et al. (2016) and Terzi et al. (2021). For the statistical analyses, cover data were converted into van der Maarel (2007) ordinal scale and aspect was converted into a southing index (with the formula S = -cos(aspect)+1). The cluster analysis of the vegetation data from our dataset, along with all existing published data of Stipa dasyvaginata subsp. apenninicola dominated communities, was performed using the Bray-Curtis index as similarity measure (Bray and Curtis 1957) and complete linkage as the agglomerative criterion to reduce chaining effects (package “vegan”, Oksanen et al. 2025). Subsequently, all the relevés were organized into a synoptic table, to summarize the floristic and syntaxonomical differences among these communities (see Table 1, which Vegetation Ecology and Diversity 62 (2025) ee167266 3 shows percentage values with cover ranges indicated as superscripts). Furthermore, to provide a more detailed characterization of our dataset, the variance of environmental data among the resulting subclusters was tested using the non-parametric Kruskall-Wallis test. In addition, normal and weighted biological and chorological spectra were developed by grouping the plants’ life forms and chorotypes into main categories (package “ggplot2”, Wickham 2016; see Suppl. material 1), to better investigate the ecology of the sampled community. All the statistical analyses were performed using R 4.5.1 (R Core Team 2025). Results The cluster analysis (Fig. 2) identified four main vegetation groups. Cluster A represents our dataset and can be further divided into two subclusters (A1 and A2). Cluster B–C represents respectively Stipa dasyvaginata subsp. apenninicola dominated grasslands from Norcia Basin (Ballelli et al. 2006; subcluster B) and from Umbria-Marche and Marche Ridge (Catorci et al. 2007; subcluster C). Cluster D represents Stipa dasyvaginata subsp. apenninicola dominated grasslands from Cingoli Ridge (Taffetani et al. 2004). Lastly, cluster E represents Stipa dasyvaginata subsp. apenninicola dominated grasslands from Tirino Valley (Pirone et al. 2001). The groups identified through multivariate analysis differ both in species composition and in environmental characteristics, with elevation emerging as the primary differentiating factor (see Table 1). Cluster A, in particular, reaches much higher elevations compared to the other groups and is clearly well separated from the other clusters. Regarding the two subclusters A1 and A2, as shown in Suppl. material 2, they differ both in elevational range and in the presence of high-elevation species. Moreover, subcluster A1 is the only group with a small number of relevés showing the presence of Stipa capillata (see Suppl. material 3). The results from ANOVA (Fig. 3) show a statistically significant difference in elevation (p-value: <0.001; Fig. 3a) and total cover (p-value: 0.014; Fig. 3f) between the two subclusters within our dataset. By contrast, the other environmental variables did not show any statistically significant difference (Fig. 3b, 3c, 3d, 3e). In conclusion, the biological and chorological spectra enabled a better understanding of the sampled community (Fig. 4). The Stipa dasyvaginata subsp. apenninicola mountain grasslands found in the Velino Massif and surrounding areas are mainly composed of hemicrypFigure 1. a) Map of the study area showing the mountain areas where field sampling occurred: sampling plots are represented by blue dots. b) Ombrothermic diagram sensu Walter and Breckle (2002) from Ovindoli station (L’Aquila, Italy): mean annual temperature and precipitation values are shown above the diagram, while mean maximum and minimum temperatures are shown on the left; frost months (in blue) and pre-frost months (in light blue) are shown below the diagram; the orange line represents the mean monthly temperature, while the blue lines represent, respectively, precipitation equal to three times the temperature (upper line) and precipitation equal to twice the temperature (lower line); the horizontal black line indicates the reference level above which the shaded blue area represents months with precipitation exceeding 100 mm, with one division corresponding to 200 mm (scale 1:10). Claudia Zitarelli et al.: The S. dasyvaginata subsp. apenninicola mountain grasslands of the Velino Massif4 tophytes and to a lesser extent chamaephytes. Other life forms are very rare, especially phanerophytes (including only few shrubs). The chorotypes highlight the peculiarity of this vegetation, characterized by the predominance of endemic species, species with a Eurasian distribution and several Mediterranean and Pontic elements. Discussion In this study, we investigated the Velino Massif and surrounding areas facing the northwestern side of the Fucino Basin, which host a fragment of the steppic grasslands typically occurring in refugia along the Apennines. These sites are characterized by local mesoclimatic and edaphic conditions that, over time, have favored the persistence of long-lasting grasslands, allowing the identification of primary grasslands (Cancellieri et al. 2017; Filibeck et al. 2020), as also observed in some Stipa dasyvaginata subsp. apenninicola grassland sites. In fact, pollen analysis conducted in the Fucino Basin, and correlated with existing profiles from other sites in southern Europe, confirm the dominance of herbaceous vegetation characterized by xerophytic taxa (Artemisia, Amaranthaceae, Ephedra, Figure 2. Dendrogram resulting from the cluster analysis. Letters refer to: Our dataset (A), Ballelli et al. 2006 (B), Catorci et al. 2007 (C), Taffetani et al. 2004 (D), and Pirone et al. 2001 (E). Figure 3. Boxplots of environmental variables with group comparisons between subclusters A1 and A2 based on ANOVA (different letters indicate significant differences between groups). Vegetation Ecology and Diversity 62 (2025) ee167266 5 Hippophae), as indicators of pronounced cold and arid conditions during the transitional period between the Pleistocene and the Holocene (MIS 12-MIS11c transition, i.e. around 430-425 ka; Vera-Polo et al. 2024). These high-resolution pollen data (Giaccio et al. 2019; Vera-Polo et al. 2024; Roberts et al. 2025) have enabled the identification of several abrupt landscape transformation events, characterized by phases dominated by steppic herbaceous vegetation, alternating with climatically humid phases in which tree vegetation prevailed (Ulmus, Acer, Tilia and subsequently Carpinus, Ostrya, Corylus, Celtis, Alnus) during the transition between the Penultimate and Last Glaciations (138-129 ka; Roberts et al. 2025). Although generalizations are difficult, a multifunctional use of ecosystems has been also repeatedly documented across different areas of the Mediterranean Basin, from Spain to Greece, varying according to the local characteristics of the biotopes (Mercuri et al. 2019). Farming communities progressively expanded northwards along the Italian Peninsula, promoting the maintenance of forest-free areas within which grassland species were preserved. The coexistence of herding and hunting activities was common during the Middle Neolithic (dating from the 6th to 5th millennia B.P.) and was later supplemented by mixed agriculture (Malone 2003) in the foothill landscapes of the Apennines (Abruzzo and Molise). Evidence emphasizes processes in which human activities, such as local deforestation, fire use and cultivation, modified an already open landscape (Sadori et al. 2004). In a comparable period (around 2800 BP, Copper Age), several evidence from the Abruzzo Apennines (Abruzzo, Molise, and Lazio National Park) indicate the first significant human impacts on forests, together with intensive livestock grazing reaching altitudes of up to 2000 m (Brown et al. 2013; Filibeck et al. 2020). From this perspective, faunal remains from the southernmost sector of the Fucino Basin (Grotta La Punta, near Ortucchio) have revealed the presence of red deer and sheep/goat remains (Malone 2003), indirectly confirming human presence and resource use during the Neolithic. Therefore, both paleoclimatic factors and human activity played a role in shaping the relict herbaceous vegetation described by Vera-Polo et al. (2024) and Roberts et al. (2025). This vegetation occurs on calcareous, calcareous-marly, and calcarenitic substrates, and occasionally on detrital deposits, similarly to other syntaxa that describe xeric grasslands dominated by Stipa dasyvaginata subsp. apenninicola in other central Apennines areas (Pirone et al. 2001; Taffetani et al. 2004; Ballelli et al. 2006; Catorci et al. 2007). These sites are invariably characterized by steep slopes and pronounced rocky outcrops, morphotypes that have allowed the conservation of relict species and communities (Kajtoch et al. 2016; Cancellieri et al. 2017; Filibeck et al. 2020; Divíšek et al. 2022). A similar consideration can be applied to the vegetation dominated by Stipa austroitalica (Fanelli et al. 2001; Terzi and D’Amico 2016) and by Achnatherum calamagrostis, which is common in highly dynamic habitats of the southern Apennines such as scree slopes and sites with shallow soil (Corbetta and Pirone 1981; Brullo et al. 1998). Analogously to the Stipa dasyvaginata subsp. apenninicola fragments found in central Apennines, this fragmented vegetation in southern Italy can be interpreted as a remnant of a much drier climatic period that occurred during the Quaternary (Spada et al. 2011). The proposal of the new syntaxon Sileno bellidifoliae-Stipetum apenninicolae (Fig. 5) is here justified by the absence of an association that adequately captures its floristic and ecological distinctiveness, as demonstrated by the clear separation of its cluster. Indeed, currently described associations refer either to sub-Mediterranean hill communities or to transitional submontane communities (Lino tommasinii-Stipetum apenninicolae Pirone et al. 2001 and Fumano procumbentis-Stipetum apenninicolae Taffetani et al. 2004 for the former; Lino tommasinii-Stipetum apenninicolae onobrychietosum albae Ballelli et al. 2006 and Stipo apenninicolae-Seslerietum juncifoliae Catorci et al. 2007 for the latter), but none of them exhibit a clear montane character with continental bioclimatic traits (Di Lena et al. 2014; Crespi et al. 2018; Cancellieri et al. 2024). Figure 4. Normal and weighted biological and chorological spectra (see Suppl. material 1). a) Biological spectrum showing the following life forms: Chamaephytes (C), Geophytes (G), Hemicryptophytes (H), Phanerophytes (P), and Therophytes (T). b) Chorological spectrum showing the following distributional ranges: Atlantic (A), Circumboreal (C), Endemic (End), Eurasiatic (Euras), Eurimediterranean (Eurim), Mediterranean-Montane (MM), South-European Orophytes (O), Pontic (P), Stenomediterranean (Stenom), and Subcosmopolitan (Subc). Claudia Zitarelli et al.: The S. dasyvaginata subsp. apenninicola mountain grasslands of the Velino Massif6 Moreover, the syntaxon we propose is more closely associated with shallow and poorly developed soils and occurs at much higher elevations (Table 1). The new association includes Stipa dasyvaginata subsp. apenninicola, Silene bellidifolia, and Rhamnus saxatilis as diagnostic species. Moreover, we found a transitional form with hilly grasslands on sites with a gentle slope, which we have interpreted as the new subassociation stipetosum capillatae with Stipa capillata as diagnostic species. This subassociation highlights the transition to other steppic grasslands dominated by Stipa capillata, which to date have been interpreted within the framework of Globu lario meridionalis-Stipetum capillatae. This latter association was described in the inner areas of the Lazio-Abruzzi Apennines (in Marsica, Capestrano, Sulmona, L’Aquila and Barisciano Basins and in Salto valley; Tammaro 1984) and later recorded in the Norcia Basin as well, where a transitional community to Stipa dasyvaginata subsp. apenninicola dominated grasslands (subass. stipetosum apenninicolae) was also identified (Ballelli et al. 2006). Regarding the new syntaxon here proposed, the longterm persistence of this relict vegetation is generally associated with specific ecological constraints and the peculiar continental mesoclimate of the area (Cancellieri et al. 2024). This vegetation, in particular, is predominantly found on steep slopes with shallow soils, where two factors act synergistically: first, the slow progression of vegetation dynamics, driven by limited soil depth and the presence of rocky outcrops (Petriccione 1993; Török et al. 2020); and second, reduced grazing pressure from sheep and horses, which preferentially utilize flatter areas with higher water availability (Adler 2001; Napoleone et al. 2021). In these areas, several shrub associations have been documented, highlighting phases of colonization of grasslands that indicate the abandonment or reduction of pastoral activities (Cutini et al. 2002; Pirone and Cutini 2002). As shown by the biological and chorological spectra, these grasslands are mainly characterized by hemicryptophytes and chamaephytes, with a high percentage of endemic species (many of which are endemic to the Apennines, such as Stipa dasyvaginata subsp. apenninicola, Sesleria nitida, and Centaurea ceratophylla subsp. ceratophylla). The presence of several species with Mediterranean and Eurasiatic distributions highlights the typical characteristics of dry grasslands in the central Apennines (Biondi et al. 2005; Di Pietro et al. 2015, 2017). The synoptic table (Table 1; see Suppl. material 4) allowed us to better explain the distinctiveness of our group, in which the diagnostic species (Silene bellidifolia and Rhamnus saxatilis) are exclusive species. At the Figure 5. Sileno bellidifoliae-Stipetum apenninicolae ass. nova: a–c–d) southern slopes of Mount Velino; b) southern slopes of Mount Cafornia. Photo credits: Claudia Zitarelli, Maurizio Cutini. Vegetation Ecology and Diversity 62 (2025) ee167266 7 Table 1. Synoptic table. Cluster A: relevés from Fucino Basin, Abruzzo (our dataset); Cluster B: relevés from Norcia Basin, Umbria (Ballelli et al. 2006); Cluster C: relevés from Umbria-Marche and Marche Ridge, Marche (Catorci et al. 2007). Cluster D: relevés from Cingoli Ridge, Marche (Taffetani et al. 2004); and Cluster E: relevés from Tirino Valley, Abruzzo (Pirone et al. 2001). The diagnostic species identified in these associations align with those chosen by the cited authors. Values represent percentage frequencies, with superscripts showing cover ranges. Cluster A B C D E Elevation range (m a.s.l.) 930–1820 1200–1250 1100–1350 580–695 320–880 Mean vegetation cover (%) 54 92 88 84 88 No. Plot 25 311 8 8 Sileno bellidifoliae-STIPETUM APENNINICOLAE ass. nova + stipetosum capillatae subass. nova* Stipa dasyvaginata subsp. apenninicola 100 3-4 100 364 +-1 75 1-3 100 1-5 Silene bellidifolia 24 +-2 . . . . Rhamnus saxatilis 32 +-1 . . . . Stipa capillata*12 4...13 + Lino tommasinii-STIPETUM APENNINICOLAE Pirone, Corbetta, Ciaschetti, Frattaroli et Burri 2001 + onobrychietosum albae Ballelli, Gatti, Raponi et Catorci 2006** Convolvulus elegantissimus ....63 +-3 Linaria purpurea .100 +. . 50 +-2 Linum tommasinii 28 +-1 67 +. . 63 +-1 Aethionema saxatile 76 +-1 100 +.38 +50 +-2 Thymus striatus subsp. striatus** 96 +-2 100 +36 +-1 . . Onobrychis alba subsp. alba** 8 +-1 100 +27 +. . Fumano procumbentis-STIPETUM APENNINICOLAE Taffetani, Zitti et Giannangeli 2004 Artemisia alba 28 +-2 .27 +100 +-2 13 + Thymus longicaulis subsp. longicaulis 4 +33 + 27 +88 +-1 . Globularia bisnagarica .33 + .88 +-1 25 +-1 Fumana procumbens 32 +-1 . . 75 +-3 25 +-1 Stipo apenninicolae-SESLERIETUM JUNCIFOLIAE seslerietosum juncifolii Catorci, Gatti et Ballelli 2007 + onobrychietosum albae Catorci, Gatti et Ballelli 2007*** Anthyllis vulneraria subsp. polyphylla . . 91 +-1 . . Sesleria juncifolia 44 +-3 .100 +-4 . . Thliphthisa purpurea subsp. purpurea 28 +-1 .82 +.13 1 Centaurea ambigua subsp. ambigua . . 64 +-1 . . Crepis lacera subsp. lacera 36 +-2 100 +73 +-1 . . Takhtajaniantha austriaca*** . . 45 +. . Helianthemum nummularium subsp. obscurum*** .67 +36 1-2 100 +-1 . Cytiso-Bromopsion erectae Bonin 1978 mut. Terzi, Di Pietro et Theurillat 2021 / Cytiso spinescentis-Saturejion montanae Pirone et Tammaro 1997 Bromopsis erecta 84 +-3 100 3100 +-2 100 1-3 63 1-4 Eryngium amethystinum 20 +-1 100 +91 +-1 .25 +-1 Phleum hirsutum subsp. ambiguum 12 +-2 100 +-1 82 +13 165 1-3 Silene otites subsp. otites 16 +-1 33 +82 +.13 1 Erysimum pseudorhaeticum 44 +-1 33 + 100 +-1 . . Centaurea rupestris 28 +-1 100 +-1 91 +-1 . . Leontodon crispus 72 +-1 33 + .38 + 38 +-1 Globularia cordifolia subsp. bellidifolia 100 +-3 .18 +-1 .13 1 Galium lucidum subsp. lucidum 68 +-3 ...25 +-1 Cytisus spinescens 92 +-2 . . . . Muscari neglectum 4 +100 +55 +.13 + Festuco laevigatae-Seslerietalia nitidae Ubaldi 2003 nom. corr. Terzi, Di Pietro et Theurillat 2021 Anthyllis vulneraria subsp. rubriflora 92 +-1 67 +-1 .63 +-1 . Allium sphaerocephalon subsp. sphaerocephalon 40 +-1 100 +64 +-1 .13 + Helichrysum italicum subsp. italicum 12 +-1 . . 63 +-3 75 +-2 Teucrium chamaedrys subsp. chamaedrys 36 +-1 100 +73 +75 +-1 . Teucrium capitatum subsp. capitatum 4 167 +.75 +-1 25 1 Teucrium montanum 52 +-1 67 +55 +-1 . . Argyrolobium zanonii subsp. zanonii ...100 +-1 13 1 Claudia Zitarelli et al.: The S. dasyvaginata subsp. apenninicola mountain grasslands of the Velino Massif8 Cluster A B C D E Elevation range (m a.s.l.) 930–1820 1200–1250 1100–1350 580–695 320–880 Mean vegetation cover (%) 54 92 88 84 88 No. Plot 25 311 8 8 Convolvulus cantabrica 4 +.9 +100 +-1 25 1 Coronilla minima subsp. minima 36 +-1 100 +. . 25 +-1 Pentanema montanum 12 +100 +27 +-2 .13 + Scabiosa columbaria subsp. columbaria 12 +. . . 38 +-2 Stachys recta subsp. recta 40 +-1 100 +.75 +25 +-1 Odontites luteus subsp. luteus .100 +. . 38 +-1 Thesium humifusum 20 1-2 100 +18 +. . Melica ciliata 48 +-2 . . . 13 + Plantago subulata .100 +64 +-2 . . Helianthemo cani-Seslerietea nitidae Terzi, Di Pietro et Theurillat 2021 Koeleria splendens 96 +-2 100 1100 +-1 13 +25 +-2 Dianthus virgineus 52 +-2 67 +100 +-1 .13 + Carex humilis 88 +-3 .82 1-2 . . Sesleria nitida 36 +-2 . . . . Helianthemum oelandicum subsp. incanum 52 +-2 .100 +-2 .13 + Helianthemum apenninum subsp. apenninum 72 +-2 100 164 +-1 .13 + Satureja montana subsp. montana 60 +-1 ...50 +-2 Festuca spp. (F. inops, F. marginata subsp. marginata)68 +-2 100 1-2 36 1-2 38 +-2 50 +-1 Linum tenuifolium 24 +-1 33 +.25 +38 +-1 Anthericum liliago 36 +-2 . . . . Centaurea triumfettii 28 +-2 .36 +. . Carex liparocarpos subsp. liparocarpos 8 +100 +-2 ... Ononis pusilla subsp. pusilla .67 +.13 2. Valeriana tuberosa 4 +100 +27 +. . Cerastium arvense subsp. suffruticosum . . 64 +-1 .13 + Other species Petrosedum rupestre 64 +-1 67 +100 +-1 .50 +-1 Poterium sanguisorba 48 +100 1-2 .63 +75 +-2 Dactylis glomerata subsp. glomerata 16 +-2 ...75 +-3 Petrorhagia saxifraga subsp. saxifraga 16 +67 +36 +88 +-1 50 +-1 Galium corrudifolium .100 +-1 64 +-1 100 +-2 13 + Medicago prostrata subsp. prostrata .100 +-1 . . 13 + Seseli tommasinii .100 2-3 . . 38 + Arenaria serpyllifolia subsp. serpyllifolia 48 +33 +. . . Anthyllis montana subsp. jacquinii 20 +-1 67 +64 +. . Eryngium campestre ...38 +63 +-2 Cephalaria leucantha 28 +-1 100 1.38 150 +-2 Crupina vulgaris 12 +33 +9 +50 +-1 50 +-2 Bupleurum baldense .33 +.63 +13 + Plantago lanceolata .100 +.50 +38 +-1 Hypericum perforatum subsp. perforatum 24 +-1 100 +. . 75 +-1 Reichardia picroides ...75 +38 +-1 Ornithogalum comosum .67 +9 +63 +. same time, a floristic uniformity at higher ranks becomes evident, due to the ecological constraints of the habitat. Therefore, given the presence of different diagnostic taxa of the class (Aethionema saxatile, Artemisia alba, Centaurea ambigua subsp. ambigua, Crepis lacera subsp. lacera, Globularia cordifolia subsp. bellidifolia, Helianthemum nummularium subsp. obscurum, Leontodon crispus, Linum tommasinii, and Thymus striatus subsp. striatus) and the ecological characterization of these grasslands, we interpret all these communities as belonging to the Helianthemo cani-Seslerietea nitidae class described by Terzi et al. (2021). This class encompasses sub-Mediterranean and sub-montane dry grasslands on calcareous substrates, spanning from the Balkan Peninsula to Spain, with the Apennines as the core. The peculiar biogeographical and floristic features of these communities underline the importance of preserving this vegetation, which displays a relictual and Vegetation Ecology and Diversity 62 (2025) ee167266 9 fragmented distribution along the Apennine chain. These considerations offer an opportunity to include these grasslands under the protection of Habitat 6210 of the Habitats Directive 92/43/EEC (Semi-natural dry grasslands and scrubland facies on calcareous substrates; Biondi et al. 2009), reported in Italy in the Alps and throughout the Apennines. It is worth mentioning that, as indicated by our subassociation Sileno bellidifoliae-Stipetum apenninicolae stipetosum capillatae, at hilly and lower elevations these grasslands come into contact with steppe grasslands dominated by Stipa capillata, which are protected under the priority Habitat 6240 (Sub-Pannonic steppic grasslands; Biondi et al. 2009). This habitat is reported in Italy only in inner alpine valleys and, more recently, on the southeastern side of the Fucino Basin, within a Natura 2000 area managed by the Abruzzo, Lazio and Molise National Park (Filibeck et al. 2022). Conclusions The present work provides an overview of the steppic grasslands of the Velino Massif and surrounding areas, investigating the ecology and syntaxonomy of a relict vegetation dominated by the endemic species Stipa dasyvaginata subsp. apenninicola. The new association here described characterizes continental mountain communities, often localized in conservative primary sites. Thus, despite the continuous landscape transformations caused by millennial climatic and environmental evolutions during the Pliocene-Holocene (Vera-Polo et al. 2024), the variable pressure along time by livestock grazing (Brown et al. 2013) as well as the different anthropogenic impacts over time (Malone 2003; Mercuri et al. 2019), the mountain biotopes facing the Fucino Basin still preserve species and communities that are unique along the Italian peninsula. Syntaxonomic scheme Helianthemo cani-Seslerietea nitidae Terzi, Di Pietro et Theurillat 2021 Festuco laevigatae-Seslerietalia nitidae Ubaldi 2003 corr. Terzi, Di Pietro et Theurillat 2021 Cytiso-Bromopsion erectae Bonin 1978 mut. Terzi, Di Pietro et Theurillat 2021 Sileno bellidifoliae-Stipetum apenninicolae Cutini, Zitarelli, De Toma et Marzialetti ass. nova Holotypus: Relevé n. 3 [Monte Velino, 42°8'46"N, 13°21'21"E, 04/07/2007, 1540 m a.s.l., 100 sq m, aspect 0.93, slope 30°, rockiness 20%, stoniness 40%, vegetation cover 50%], Stipa dasyvaginata subsp. apenninicola 4; Silene bellidifolia +; Rhamnus saxatilis +; Bromopsis erecta +; Cytisus spinescens 2; Galium lucidum subsp. lucidum +; Sesleria juncifolia 2; Aethionema saxatile +; Leontodon crispus +; Brachypodium genuense +; Pimpinella tragium 1; Seseli montanum subsp. montanum +; Erysimum pseudorhaeticum 1; Cerastium tomentosum +; Crepis lacera subsp. lacera +; Centaurea rupestris 1; Thliphthisa purpurea subsp. purpurea 1; Cynanchica aristata subsp. aristata +; Anthyllis vulneraria subsp. rubriflora +; Stachys recta subsp. recta 1; Polygala major +; Artemisia alba 1; Teucrium chamaedrys subsp. chamaedrys +; Teucrium montanum 1; Pentanema montanum +; Globularia cordifolia subsp. bellidifolia 2; Carex humilis 1; Koeleria splendens +; Thymus striatus subsp. striatus 1; Dianthus virgineus 1; Sesleria nitida 2; Helianthemum oelandicum subsp. incanum 2; Satureja montana subsp. montana 1; Anthericum liliago 1; Centaurea triumfettii +; Plantago argentea subsp. argentea 1; Petrosedum rupestre +; Arenaria serpyllifolia subsp. serpyllifolia +; Sempervivum tectorum +; Juniperus oxycedrus +; Sedum dasyphyllum subsp. dasyphyllum +; Euphorbia cyparissias +; Euphorbia myrsinites subsp. myrsinites +; Amelanchier ovalis subsp. ovalis +; Siler montanum subsp. stabianum +; Daphne oleoides subsp. oleoides +. Sileno bellidifoliae-Stipetum apenninicolae stipetosum capillatae Cutini, Zitarelli, De Toma et Marzialetti subass. nova Holotypus: Relevé n. 24 [Collarmele, 42°4'39"N, 13°38'39"E, 24/07/23, 930 m a.s.l., 64 sq m, aspect 1.98, slope 60°, rockiness 30%, stoniness 40%, vegetation cover 60%], Stipa dasyvaginata subsp. apenninicola 3; Rhamnus saxatilis +; Stipa capillata 3; Bromopsis erecta 3; Cytisus spinescens 1; Festuca inops 2; Stachys italica 1; Galium lucidum subsp. lucidum 3; Pimpinella tragium +; Silene otites subsp. otites +; Eryngium amethystinum +; Melica ciliata 2; Anthyllis vulneraria subsp. rubriflora 1; Teucrium chamaedrys subsp. chamaedrys 1; Teucrium montanum +; Scabiosa columbaria subsp. columbaria +; Globularia cordifolia subsp. bellidifolia 2; Carex humilis 2; Koeleria splendens 2; Thymus striatus subsp. striatus 2; Dianthus virgineus 1; Helianthemum apenninum subsp. apenninum +; Satureja montana subsp. montana 1; Petrosedum rupestre 1; Dactylis glomerata subsp. glomerata 2; Arenaria serpyllifolia subsp. serpyllifolia +; Cephalaria leucantha 1; Poterium sanguisorba +; Petrorhagia saxifraga subsp. saxifraga +; Allium flavum subsp. flavum +; Centaurea ceratophylla subsp. ceratophylla 1; Cuscuta epithymum subsp. epithymum +; Silene saxifraga +.