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 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.

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 hemicryptophytes 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.

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, 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 6^th to 5^th 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 Saponario 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 apenninicolaePirone et al. 2001 and Fumano procumbentis-Stipetum apenninicolaeTaffetani et al. 2004 for the former; Lino tommasinii-Stipetum apenninicolae onobrychietosum albaeBallelli et al. 2006 and Stipo apenninicolae-Seslerietum juncifoliaeCatorci 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). 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, Saponaria 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 Globu­lario 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 long-term 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 (Saponaria bellidifolia and Rhamnus saxatilis) are exclusive species. At the 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 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 Saponario 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).

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.

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

Saponario 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; Saponaria 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 +.

Saponario 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 +.

Globulario meridionalis-Stipetum capillatae Tammaro 1984; Lino tommasinii-Stipetum apenninicolae Pirone, Corbetta, Ciaschetti, Frattaroli et Burri 2001; Fumano procumbentis-Stipetum apenninicolae Taffetani, Zitti et Giannangeli 2004; Lino tommasinii-Stipetum apenninicolae onobrychietosum albae Ballelli, Gatti, Raponi et Catorci 2006; Stipo apenninicolae-Seslerietum juncifoliae Catorci, Gatti et Ballelli 2007.