The MNP, located in the central Apennines, Italy (Fig. 1), was established in 1995 by National Law 1991, n. 394, to preserve, protect and enhance the high value of the inherent natural, historical, and cultural resources of the area. The Park consists mainly of carbonate mountains, separated by valleys and karst high plateaus, with a broad altitudinal range (130–2,793 m a.s.l.). The Majella massif has more than 60 peaks, with half of them rising above 2,000 m, and includes the second highest peak in the Apennines, Mount Amaro (2,793 m). The landscape is dominated by NW-SE-oriented limestone ridges reaching above the beech tree-line, i.e., Morrone, Rotella, Pizzalto, and Porrarra, and by the Majella massif. These mountains are composed almost exclusively of deep layers of limestone where all the geological eras from the Triassic onwards are represented. A unique periglacial plateau at ca. 2000 m lies above the Majella massif and includes more than 15 smooth summits (Whitehead 1951; Stanisci et al. 2011). The Apennines were repeatedly glaciated during the Pleistocene (Giraudi 2005), while simultaneously the Adriatic Sea retreated resulting in the formation of a broad periglacial plain. During the last glacial maximum, the upper part of the massif was covered by a thick ice layer 30 km² wide and more than 200 m thick (Jaurand 1994), with glacial tongues spreading over all the adjacent valleys down to 1330/1400 m of altitude. The Würm glaciation gave origin to macroscopic forms of erosion, circles and moraine deposits; the current periglacial landscape is exposed to frost wedging and long-term snow persistence (7–8 months a year; Stanisci et al. 2005). From a bioclimatic point of view, the study area is included in the alpine biogeographical region (Cervellini et al. 2020) and the climate corresponds to the subalpine-alpine humid type as far as the lower summit is concerned, whereas the other summits belong to the alpine humid type (Blasi 2005). Sub-Mediterranean conditions prevail up to ≈ 1000 m a.s.l. (van Gils et al. 2012). The landscape below 1000 m is a patchwork of villages, farmlands, oak forest fragments (Quercus pubescens Willd. subsp. pubescens), and shrubs. At mid-elevation (ca. 1000–1750 m), the territory is dominated by contiguous monospecific beech (Fagus sylvatica L. subsp. sylvatica) forests (van Gils et al. 2010). Secondary montane grasslands, currently unutilized, prevail on south-eastern to south-western slopes. Summer pastoralism above the beech tree-line has been gradually abandoned over the past century.

Figure 1. 

Geographical position of the Majella National Park.

The Park’ territories are part of the Natura 2000 network. The boundaries coincide with a Special Protection Area (SPA) for the conservation of wild birds (established by the Birds Directive 79/409/EEC). Furthermore, within the Park, there are four Special Areas of Conservation (SAC), established by Habitat Directive 92/43/EEC.

The MNP is extremely rich in plant species, indeed its flora stands out for the high number of specific and subspecific taxa, for a total of 2,286 (Conti et al. 2019), including 15 exclusive endemics, such as Pinguicula fiorii Tammaro & Pace, Soldanella minima Hoppe subsp. samnitica Cristof. & Pignatti, Aquilegia magellensis F.Conti & Soldano, Centaurea tenoreana Willk., Crepis magellensis F.Conti & Uzunov. Italian endemic taxa are 201 (8.8%), grouped in 104 genera and 35 families (Conti et al. 2019).

Moreover, the MNP is the locus classicus for 49 species and subspecies (Peruzzi et al. 2015, 2019). The Park administration established and maintains two botanical gardens, a seed bank, and a herbarium for preserving and studying the flora of the Park (Di Martino et al. 2016a).

For the ex-situ conservation of plant biodiversity, in implementation of art. 9 of the Rio de Janeiro Convention on Biological Diversity, the Park Authority manages two botanical gardens (the “Michele Tenore” at Lama dei Peligni and the “Daniela Brescia” at Sant’Eufemia a Maiella) and the Majella Seed Bank (MSB). These structures aim to collect, study, and conserve seeds of wild species to preserve their genetic heritage.

The “Michele Tenore” Botanical Garden was created in 1995 and currently covers an area of 9,000 m². It harbours 433 plant species and is dedicated to the Neapolitan botanist Michele Tenore who visited the area in 1831 and described several species. The “Daniela Brescia” Botanical Garden is located at 900 m a.s.l. in the MNP. It was set up in 2001 and currently includes 545 plant species over a surface area of more than 40,000 m². The Garden was designed to contain reproductions of some mountain environments of the central Apennines, like high-altitude cliffs and screes , while an area is dedicated to demonstrative didactic sections, such as the field-showcase of agricultural biodiversity.

The MSB is situated in the “Michele Tenore” Botanical Garden. Its main aim is to preserve particularly rare and/or endangered wild or cultivated species. Seed banks are generally considered a strong and effective tool for long-term biodiversity conservation (Williams et al. 2003; Mattana et al. 2005) and an important potential seed source for the restoration of plant communities (Bakker and Berendse 1999). Furthermore, the ability to store a large diversity in a small space makes seed banking a practical and attractive tool for plant conservation (Liu et al. 2018).

The MSB was established in 2005 to mark the founding of R.I.B.E.S., the Italian Germplasm Bank Network (Rossi et al. 2006; Bonomi et al. 2008), whose founding members include the Park and 15 seed banks throughout Italy. Currently, the MSB is a reference at a Regional level and is a tangible example of the interaction between ex-situ and in-situ conservation, in the context of Italian national parks. The work of the MSB is inspired by specific international conservation conventions and strategies (Convention on Biological Diversity, CBD 1992; Global Strategy for Plant Conservation, GSPC 2012; European Plant Conservation Strategy, EPCS 2002). The MSB ex-situ conservation activities mainly focus on rare and/or endangered or endemic alpine belt species, many of which are in the IUCN lists and/or protected by international conventions (Cites, Bern, etc.), the European Community “Habitat” Directive 43/92, Regional Law no. 45/1979 for the protection of Flora in Abruzzo and Red Lists of Italian Plants (Rossi et al. 2013; Orsenigo et al. 2018, 2020). Furthermore, activities relating to restocking projects on plant species included in annex II of the Habitats Directive are underway. The fact that the MSB is managed by a national park is a unique case in Italy.

The Botanical Garden at Sant’Eufemia a Maiella also incorporates the “Native Plants Nursery”, aimed exclusively at reproducing (from seed or cuttings) native plant species, particularly the most endangered ones, and/or growing plants for use in the Botanical Gardens or the Park itself. Since 2011, the Sant’Eufemia nursery has been authorised to operate as a commercial nursery (pursuant to D.Lgs 214/2005) and is officially registered in the Producers’ Register. In this nursery, 140 species are grown, including perennial herbs, shrubs, trees, and medicinal plants. The nursery’s high-quality native plant production is a valid support for public administrations and other public and private botanical gardens (Di Martino et al. 2020).

The Sant’Eufemia Botanical Garden hosts the herbarium which, to date, boasts more than 3,000 specimens, including the critical groups and most endangered species of the Majella massif.

The Park is the coordinator for the FLORANET project, together with the Apennine Flora Research Centre, Legambiente, and other protected areas in Abruzzo. As part of the European Union’s LIFE programme, this project aims to preserve the plant species in annex II of the European Directive 43/92/EEC, the “Habitats Directive” (Di Martino et al. 2016a).

Several research activities carried out by the Park support specific in-situ conservation measures: management plans for the Natura 2000 sites, monitoring of exclusive endemic flora, a census of monumental trees, monitoring natural populations of native Pinus nigra J.F.Arnold subsp. nigra var. italica Hochst. and Betula pendula Roth, and controlling the spread of the invasive alien species Senecio inaequidens DC.

The plants grown in the Garden’s nursery can be used to (i) reintroduce species that are extinct in the protected areas, (ii) limit the collection of material from the wild in order to maintain the Garden’s own collection; (iii) reinforce very small populations of rare species, thereby improving their chances of survival; (iv) encourage the use of native plants for ornamental purposes; (v) encourage the cultivation of native medicinal herbs and traditional fruit trees, thus boosting the use of local species or ecotypes to restore and improve degraded environments.

In Table 1, Red List taxa cultivated in the Botanical Gardens of the MNP are listed, divided by conservation status. At present, nine threatened taxa (CR, EN) and ten Near Threatened (NT) taxa are being cultivated.

A list of endemic taxa cultivated in the two Botanical Gardens is reported in Table 2.

Currently, 52 endemic taxa are being cultivated in the Botanical Gardens, while 226 accessions of wild species are preserved for long-term storage; they were collected over a period of 15 years (from 2005 to 2019). Figure 2 shows the 171 different taxa (species and subspecies) stored in the MSB divided into families. The results highlight the achievement of target 8 for the percentage of species available for restocking, but the Park plans to reach the goal of 75% of the threatened species in the near future.

Figure 2. 

Number of species and subspecies per family stored in the Majella Seed Bank.

The most represented families are Asteraceae with 21 taxa, Ranunculaceae with 12, Brassicaceae with 11, Caryophyllaceae and Fabaceae with 10 taxa. Forty-nine taxa are included in the Italian Red List (Rossi et al. 2013, Orsenigo et al. 2018, 2020), and among them, two are Critically endangered (CR): Bubon macedonicum L. and Genista pulchella Vis. subsp. aquilana F.Conti & Manzi; four are Endangered (EN): Adonis distorta Ten., Astragalus aquilanus Anzal., Typha minima Funk ex Hoppe, and Pinguicula fiorii Tammaro & Pace; one is Vulnerable (VU): Oxytropis ocrensis F.Conti & Bartolucci; 11 are Near Threatened (NT): Aquilegia magellensis F.Conti & Soldano, Carex microcarpa Bertol. ex Moris, Crepis magellensis F.Conti & Uzunov, Fritillaria montana Hoppe ex W.D.J.Koch, Gentiana lutea L. subsp. lutea, Iris marsica I.Ricci & Colas., Jacobaea vulgaris subsp. gotlandica (Neuman) B.Nord., Leontopodium nivale (Ten.) Huet ex Hand.-Mazz., Phyllolepidum rupestre (Sweet) Trinajstić, Saxifraga italica D.A.Webb, Soldanella minima Hoppe subsp. samnitica Cristof. & Pignatti; 28 are Least Concern (LC); three are Data Deficient (DD).

Five taxa are listed in annex II of the Habitats Directive (1992) in Majella and they are stored in the seed bank (each with a Natura 2000 species code): Adonis distorta Ten. (1479), Androsace mathildae Levier (1630), Astragalus aquilanus Anzal. (priority – 1558), Cypripedium calceolus L. (1902), Himantoglossum adriaticum H.Baumann (4104). These five species are all present in the directive of the Park’s territory, therefore 100% of the species listed in annex II are preserved in the seed bank. Amongst these, Astragalus aquilanus and Androsace mathildae are narrow endemics to the Abruzzo Region, Adonis distorta is endemic to the central Apennines, while the two Orchidaceae, although extremely localized (especially Cypripedium calceolus), show a wider distribution.

During the past years, many species have been studied and several scientific articles published by the seed bank working group; these were inherent to the ecology of germination, the morphometry of seeds, seed dispersal mechanisms, etc. (Frattaroli et al. 2013; Di Martino et al. 2014, 2015; Di Cecco et al. 2017a, 2017c, 2018, 2019; Di Musciano et al. 2018).

These studies are carried out in order to support the preservation of plant germplasm and in-situ conservation activities. One of the main projects undertaken by the MSB is the EU-funded Life Floranet that aims to preserve species and ecosystems within the Natura 2000 network (Di Martino et al. 2016a). For this project, which is still in progress, 10 restockings has been scheduled, divided by species and action (Table 3). All the plant material used for restocking came from the MSB and the Sant’Eufemia nursery (Figure 3).

Figure 3. 

Plants of Iris marsica grown in the Majella National Park nursery).

The applied strategies of the MNP are essential to perform concrete actions for species conservation. Such actions go from the collection of seeds in nature, in-vitro propagation, seed reproduction long-term storage, in-situ restocking up to the creation of new populations. The functionality and operation of the structures, that are already well established for ex-situ conservation, have been tested on restocking actions.

The restocking actions were carried out not only within the boundary of the MNP, but were also extended to other areas of central Italy. In this context, considering that from the 67,620 seed accessions of native plants stored in European seed banks (ENSCOBASE) only 64 (0.09%) were used in translocation programs (Abeli and Dixon 2016), the MNP plays a crucial role in carrying out these strategies and in monitoring endemic or rare species, thereby avoiding the risk of extinction (Dalrymple and Abeli 2019). To date, eight accessions of the MSB have been used for restocking actions. The total number of accessions preserved in MSB is 256, consequently, the percentage of accessions used for restocking is 3.125%, which is significantly higher than the percentage indicated by Abeli and Dixon (2016). Such actions provide exceptionally strong support to the MNP by making use of a truly integrated conservation strategy through the concerted action of its in-situ / ex-situ facilities.

The amount of seeds stored in the MSB is growing constantly, both in terms of number of species and the area covered. At a Regional level, this represents a way to preserve the genetic resources of both wild species (including trees) and cultivated species of agricultural, culinary, and ornamental interest. The MSB is a landmark in Italy as an example of integration between ex-situ and in-situ conservation.

The strong effort for ex-situ conservation made by the MNP is confirmed by the high number of conserved species. Indeed one-third of the Italian endemic species of the Park (68 species out of 201) and one-third of the species present in the Red Lists (65 species out of 195) are conserved at the Botanical Gardens and in the seed bank. Future collection programmes will be focused on the missing endangered and endemic taxa. The main aim is to achieve the conservation and/or cultivation of at least 75% of the threatened and endemic taxa. This will be done essentially within the Park’s territory without, however, excluding species from the Abruzzo Administrative Region and throughout the Apennines.

As regards the agricultural heritage, more than 150 Crop Wild Relatives (CWR; Di Martino et al. 2016c) and 61 native agricultural varieties (Di Santo and Di Cecco 2015) have been identified within the MNP. Conservation plans for the CWR and landraces are made in accordance with the FAO Treaty and the Nagoya International Protocol on access to genetic resources and the fair and equitable sharing of benefits arising from their utilization as per the convention on biological diversity. Several CWR species are cultivated and conserved in both Botanical Gardens. For both wild and cultivated taxa, in vivo cultivation and reproduction is a fundamental step towards conservation planning within protected areas (Heywood 2019). On the other hand, seeds are stored in the MSB, also for studies relating to reproductive biology (Di Cecco et al. 2019). For some species, such as Secale strictum (C.Presl) C.Presl subsp. strictum, Apium graveolens L., and some species of Lathyrus, the number of individuals, and the status of populations are being monitored. Moreover, germination tests were carried out to increase knowledge about seed storage and germination niche (Di Cecco et al. 2017b).

The seed bank also stores seed from native local landraces recovered as part of the “Let’s Grow Diversity” project. Conserving these landraces is important because they are the result of a long, balanced co-evolution over centuries between Man and nature (Hawkes et al. 2012). The most significant landraces include “Solina” and “Marzuolo” wheats and “Caffè” and “Socere e Nore” beans.

All these actions suggest as an integrated strategy from the seed bank to field cultivation and restoking action is crucial for biodiversity conservation. Furthermore, the collaborations with other botanical gardens and germplasm banks are always desirable to exchange knowledge, procedures, and techniques and duplicate collections.