The indigenous vascular flora of the forest domain of Anela (Sardinia, Italy)

Emmanuele Farris; Michele Carta; Salvatore Circosta; Salvatore Falchi; Guillaume Papuga; Peter de Lange

doi:10.3897/phytokeys.113.28681

Abstract

The importance of mountains for plant diversity and richness is underestimated, particularly when transition zones between different bioclimates are present along altitudinal gradients. Here we present the first floristic data for a mountain area in the island of Sardinia (Italy), which exhibits Mediterranean bioclimates at the bottom and temperate bioclimate at the top. We discovered a very high floristic richness, despite the fact that the number of endemic taxa is not high and the number of exclusive taxa is very low. Many of the detected taxa are at their range periphery and/or ecological margin. We conclude that climate transition zones in Mediterranean mountains and especially on islands are key areas regarding plant biodiversity and should be better investigated and protected.

Keywords

bioclimate, biodiversity, Mediterranean mountains, submediterranean, temperate

Introduction

Mountains are a critical landscape and ecosystem; they not only provide water for the lowlands but are a source of well-being and inspiration for numerous people (Korner 2004). The green ‘coat’ of the world’s mountains is composed of specialised biota, all nested in a great variety of microhabitats. Mountains biota are determined by a series of climatically different life zones over short elevational distances (Rahbek 1995, Korner 2000, Hemp 2002, Korner and Paulsen 2004), which often result in areas of high biodiversity of high conservation interest (Korner 2004). However, those areas are also under high threat regarding climate change, as it is expected that they experience drastic changes (Inouye 2008).

Mountain biodiversity can be studied at a multitude of scales in space, time and function (Molau 2004). Even though species richness is usually the focal component in nature conservation, genetic diversity within species is equally important. The small-scale distribution of species in the tropical Andes, as exemplified by the plant genera Calceolaria (Calceolariaceae) and Bartsia (Orobanchaceae), contrasts against the situation in high-latitude mountains, e.g. the Scandes, where species have wide ranges and many are circumpolar (Molau 2004). Several studies on alpine plants, based on molecular data, show that the intraspecific genetic diversity tends to increase with latitude, a situation brought about by glaciation cycles permitting repeated contraction-expansion episodes of species’ distributions (Abbott et al. 2000, Abbott and Brochmann 2003, Gamache et al. 2003, Holderegger and Abbott 2003, Lian et al. 2003, Abbott and Comes 2004). In tropical mountains, species distributions are geographically much narrower, often as a result of relatively recent, local speciation (Deshpande et al. 2001, Friar et al. 2001, Tremetsberger et al. 2003a, 2003b, Zhou et al. 2003). Thus, the classical decrease of genetic diversity observed from the equator toward the pole can eventually be blurred for mountain species. Actually, repeated contraction-expansion of species ranges has led rear edge populations to maintain some genetic diversity, therefore counterbalancing the effect of peripheral isolation (Hampe and Petit 2005). Conjointly, the high genetic differentiation between populations underlines the conservation relevance of those populations.

Mediterranean mountains represent an interesting case, because they often have a relic temperate-like bioclimate at their top (with no or little summer drought) in a context characterised by severe water deficit for at least two consecutive months at lower altitudes. Mediterranean mountains can therefore be considered as climatic islands, where plant diversity patterns are influenced by different factors (or in different ways) with respect to temperate and boreal mountains (Winkler et al. 2016). Furthermore, climatic and land-use changes have different effects on Mediterranean vs Boreal-Temperate mountains of Europe, being detrimental for the floral richness of the first and increasing the species richness of the second (Pauli et al. 2012). Considering that expected climatic trend is an increasing of temperature and a decrease of precipitation (mainly during spring) in Mediterranean mountains, whereas non-Mediterranean European mountains will not experience a reduction of annual and spring precipitation (Bravo et al. 2008), the urgency rises to monitor those mountains at the transition between Temperate and Mediterranean bioclimates. Moreover, before the middle of the century, the expected climatic changes will provoke the disappearance or strong reduction of a suitable habitat in the summit area, where most of the endemic and/or rare species are located (Benito et al. 2011). The most endangered habitats and species are those linked to water availability like streams, wet meadows and temporary ponds (Ghosn et al. 2010, Pérez-Luque et al. 2015). On islands, threats to mountain floras are even more acute compared to mainlands, because narrower spatial scales of habitats and the usually lower mountain altitudes (Vogiatzakis et al. 2016), led some species to have a relic distribution (Petit et al. 2005, Mayol et al. 2015, Fazan et al. 2017). Historical climatic fluctuations and associated ecological constraints are the basis of the fragmented distribution of Boreal-Temperate species on Mediterranean mountains (Mayol et al. 2015, Iszkulo et al. 2016) and determine the presence of plant refugia, climatically stable areas that constitute key areas for the long-term persistence of species and genetic diversity, especially at present and future decades given the threat posed by the extensive environmental change processes operating in the Mediterranean region. These refugia, including large Mediterranean islands, represent ‘phylogeographical hotspots’; that is, significant reservoirs of unique genetic diversity favourable to the evolutionary processes of Mediterranean plants (Médail and Diadema 2009).

The island of Sardinia, the second largest in the whole Mediterranean basin, was already known to have a prevalent Mediterranean bioclimate, with a temperate bioclimate in the two main massifs of the island, the Gennargentu (centre-eastern Sardinian, maximum elevation 1834 m a.s.l.) and the Limbara (north-eastern Sardinia, maximum elevation 1359 m a.s.l.) (Arrigoni 1968). Recent detailed bioclimate analysis (Canu et al. 2015) also showed that the only mountain chain of the island named Marghine-Goceano (located between the Limbara and the Gennargentu massifs, maximum elevation at Mt. Rasu 1259 m a.s.l.) is characterised by a temperate bioclimate (in the sub-Mediterranean variant) along the ranges summit. Although the mountain floras of the Gennargentu and Limbara are well documented (Veri and Bruno 1974, Arrigoni and Camarda 2015), floristic information about the Marghine-Goceano range is lacking (Valsecchi and Corrias 1966).

This paper goes some way to fill this knowledge gap by reporting on the indigenous flora of a forest domain located in the middle of the Marghine-Goceano range. Our aim was to provide a checklist of the flora located in this area to enable future characterisation of the biotic environment of this mountain area of Sardinia. This data will also allow the identification of target species to monitor and understand climate changes in the particular context of Mediterranean islands.

Methods

Study area

The forest domain of Anela is a public property since 1886, at present managed by the Sardinian regional agency Forestas (Fig. 1). The domain covers 1280 hectares of which 1200 ha fall in the municipality of Anela, 55 ha in that of Bultei (to the east) and 25 ha in that of Bono (to the west). The lowest altitude is about 600 m a.s.l. in locality Badu Edras whereas the summit point is at Punta Masiedda 1158 m a.s.l. The geographic coordinates of the forestry station headquarter are 40°27'14"N; 9°01'36"E. At present, the vegetation cover is mainly characterised by coppices and mature shrubs linked to sub-Mediterranean woods Glechomo sardoae-Quercetum congestae and Saniculo europaeae-Quercetum ilicis above 800 m a.s.l. and meso-Mediterranean Loncomelo pyrenaici-Quercetum ichnusae and Galio scabri-Quercetum ilicis below 800 m a.s.l., as described by Bacchetta et al. (2009). The 2004 forest census determined that 46% of this area was occupied by holm oak (Quercus ilex L.) woods, 2.7% by deciduous oak (Q. pubescens Willd.) woods, 23.4% by mixed woods of holm oak and deciduous oak, 0.8% by cork oak (Q. suber L.) woods, 2.8% by plantations with alien trees (Abies, Cedrus, Acer, Fagus, Pinus), 14.7% by shrub communities (with Erica arborea, Crataegus monogyna, Rubus ulmifolius), 6.2% by dwarf communities (with Helichrysum microphyllum subsp. tyrrhenicum, Thymus herba-barona, Genista desoleana), 0.3% by rocky places and the rest by human activities (including buildings, an artificial lake and firebreaks) (Sechi and Falchi 2013). It should be noted that a large fire destroyed 800 hectares of the domain on 31 July 1945, so the wooded area decreased from 72.4% in 1910 to less than 20% in the 50s (Sechi and Falchi 2013).

Figure 1.

The study area, Forest of Anela and its location in Sardinia (red rectangle on the inset map). Colours on the map represent different isobioclimates (derived from Canu et al. 2015). In the domain, we can recognise five different isobioclimates: Violet: upper mesotemperate (subMediterranean), lower humid, weak semi-continental; blue: lower supraMediterranean, lower humid, weak semi-continental; orange: upper mesoMediterranean, lower humid, weak semi-continental; lilac: upper mesoMediterranean, upper subhumid, weak semi-continental; pink: upper mesoMediterranean, lower subhumid, weak semi-continental. Thick black lines represent domain limits; thin black lines represent altitude intervals of 100 m.

In the ambit of the Sardinian-Corsican biogeographic province (as defined by Bacchetta et al. 2012), the study area falls in the Goceano-Logudorese sector (Fenu et al. 2014).

The geology of the study area comprises Palaeozoic granites and schists (Madrau 2013). The impermeable nature of these substrates has created a substantial aquifer evident by the presence of 39 springs (half perennial and half seasonal) in the study area (Farris 2013b).

Bioclimate analysis of 1971–2000 data (Canu et al. 2015) showed that 96.9% of the area falls in the Mediterranean Pluviseasonal Oceanic bioclimate, whereas 3.1% in the Temperate Oceanic bioclimate (submediterranean variant). A total of 64.6% of the area is included in the meso-Mediterranean thermotype, 32.3% in the supra-Mediterranean and 3.1% in the meso-Temperate.

Thermo-pluviometric data of the period 1951–1985 showed annual mean temperature of 11.2 °C and annual mean rainfall of 1040 mm; after the year 2000 temperatures did not vary significantly, whereas a reduction of ca. 30% in the annual rainfall was recorded. Late spring and summer rainfall (May-August) decreased even more (more than 50%, see Farris 2013a).

The study area is entirely included in the Natura 2000 site of community importance ITB 011102 ‘‘Catena del Marghine e Goceano’’, extended on 14,984 ha and is also nominated as a Protection Oasis for wildlife “Foresta Anela”, managed by the Province of Sassari.

Floristic research

Floristic research started in the year 2000 and was intensified in the years 2012–17 with regular monthly sampling. Each month, we made one day excursions, which covered three altitudinal ranges (< 800 m a.s.l.; 800–1000 m a.s.l.; > 1000 m a.s.l. on the third). For each excursion, we tried to visit as many habitats as possible in order to capture the highest environmental heterogeneity. Collected plants were stored at the Herbarium SS, where we also searched for specimens collected in previous decades (if present, they are reported in the floristic list).

Plant names were derived from the Euro+Med PlantBase (Euro+Med 2006–2018), except for: a) families not already included in this database for which we referred to the Checklists of Italian Flora (Conti et al. 2005; Bartolucci et al. 2018), APG IV (APG 2016); b) the family Orchidaceae (for which we follow GIROS (2016)); c) the genus Orobanche, for which we follow Domina and Arrigoni (2007); d) the genus Dianthus, for which Bacchetta et al. (2010) is followed; e) and the species Struthiopteris spicant which we use in preference to Blechnum spicant (Gasper et al. 2016); f) for endemics, we also consulted Arrigoni et al. (1976–1991) and Peruzzi et al. (2014). The Italian floras (Pignatti 1982, 2017–2018) and the Sardinian flora (Arrigoni 2006–2015) were also consulted. When other relevant literature was followed, it is specified in the text.

Plant authorities and names were further verified using ‘The Plant List’, ‘The World Checklist of Selected Plant Families’ and ‘The International Plant Names Index’ (IPNI). Herbarium acronyms follow Thiers (2018).

The taxonomic circumscription of orders and families, as well as their sequence in the list was derived from Smith et al. (2006) for Pteridophytes; and APG III (APG 2009), APG IV (APG 2016) and Haston et al. (2009) for Angiosperms. Within each family, genera, species and subspecies are listed in alphabetical order. Species and subspecies are numbered progressively.

For each taxon we report:

Progressive number Scientific name (with authority) Biological type, Chorologic type

Abundance (locality(ies) of collection is(are) specified only for uncommon or range restricted taxa): Habitat

Notes (eventual)

Biological types are in accordance to Raunkiær (1934) and were verified on the collected samples and also in Pignatti (1982, 2017–2018); chorologic types were determined following maps reported in the Euro+Med PlantBase (Euro+Med 2006–2018) and again verified in Pignatti (1982, 2017–2018) and the other bibliographic sources reported in the text.

Geographical abbreviations are:

Atl. Atlantic;

Cauc. Caucasian;

Circumbor. circum-boreal;

Cosmop. cosmopolitan;

Endem. endemic;

Euras. Eurasian;

Eurimedit. euri-Mediterranean;

Europ. European;

Eurosib. Euro-Siberian;

It Italy;

Itc central Italy;

Its northern Italy;

Macaron. Macaronesian;

Medit. Mediterranean;

Medit. Mont. Mediterranean montane;

S. Europ. Orof. Southern European Orophylous;

Paleotemp. paleo-temperate;

Paleotrop. paleo-Tropical;

Sib. Siberian;

Stenomedit. Steno-Mediterranean;

Subatl. sub-Atlantic;

Subcosmop. sub-cosmopolitan;

Submedit. sub-Mediterranean;

Subtrop. sub-Tropical;

Turan. Turanian.

Here we consider as endemics sensu stricto all taxa limited to the Corsican-Sardinian biogeographic province (sensuBacchetta et al. 2012), therefore including the Tuscan Archipelago. Other taxa are considered endemic sensu lato, which includes those present in western Mediterranean islands and continental areas – Calabria in Europe, Kabylies in Africa – as far as the Miocene part of the Hercynian chain (Hercynian endemics sensuMansion et al. 2008). Finally, other endemics sensu lato are ‘administrative endemics’, i.e. taxa confined within Italian national borders (Peruzzi et al. 2014). For endemics, geographic abbreviations are as follows:

Ag Algeria;

AT Tuscan Archipelago;

Bl Balearic Islands;

Co Corsica;

Hy Hyères islands;

Sa Sardinia;

Si Sicily.

Abundance is expressed on the basis of the following criteria:

RR range restricted: taxa present in only one locality within the study area or covering a surface not exceeding 1 hectare, i.e. Mentha requienii subsp. requienii;

U uncommon: taxa found in 2–5 localities within the study area, or covering a surface not exceeding 1 km², i.e. Arisarum vulgare;

L localised: taxa present in 6 or more localities within the study area, but covering less than 2.5 km², i.e. Agrostis capillaris;

C common: taxa covering more than 2.5 km², i.e. Quercus ilex.

Results

Floristic list

Lycopodiopsida

Isoetales

Isoetaceae

1 Isoetes histrix Bory G bulb, Stenomedit.-Atl.

U (Zuanne Cane Malu, near Mt. Masiennera): Temporary ponds

Selaginellales

Selaginellaceae

2 Selaginella denticulata (L.) Spring Ch rept, Stenomedit.

C: Woods, wet cliffs

Polypodiopsida

Osmundales

Osmundaceae

3 Osmunda regalis L. G rhiz, Subcosmop.

L: Alnus glutinosa woods, streams

Polypodiales

Dennstaedtiaceae

4 Pteridium aquilinum (L.) Kuhn subsp. aquilinum G rhiz, Cosmop.

C: Woods, meadows, fringes, garrigues, shrublands

Pteridaceae

5 Anogramma leptophylla (L.) Link T caesp, Cosmop.

L: Shady rocks and cliffs

Aspleniaceae

6 Asplenium adiantum-nigrum L. H ros, Paleotemp.

C: Shady rocks and cliffs, sometimes woods

Notes: since the taxon has been excluded from the Sardinian flora by Marchetti (2004), Arrigoni (2006–2015) and Bartolucci et al. (2018), here we consider it as new for the Sardinian flora.

7 Asplenium onopteris L. H ros, Subtrop.

C: Woods, sometimes cliffs

8 Asplenium ceterach L. subsp. ceterach H ros, Euras.

L: Walls

9 Asplenium foreziense Magnier H ros, NW-Medit.-Mont.

U (Badu Edras): Shady rocks and cliffs

Notes: the taxon has been excluded from the Sardinian flora by Marchetti (2004) and Bartolucci et al. (2018), but confirmed by Arrigoni (2006–2015).

10 Asplenium obovatum Viv. subsp. obovatum H ros, Stenomedit.

U (Mt. Masiennera): Crevices at the top of the mountain

11 Asplenium trichomanes subsp. quadrivalens D.E. Mey. H ros, Cosmop.

C: Shady rocks and cliffs

Woodsiaceae

12 Athyrium filix-femina (L.) Roth H Ros, Subcosmop.

L: Wet places, mainly Alnus glutinosa woods

Blechnaceae

13 Struthiopteris spicant (L.) F.W.Weiss H ros, Circumbor.

RR (Few individuals in a wet wood near Sos Sauccheddos spring): Alnus glutinosa wood

Dryopteridaceae

14 Polystichum setiferum (Forssk.) Woyn. G rhiz, Circumbor.

C: Woods

Polypodiaceae

15 Polypodium cambricum L. subsp. cambricum H ros, Eurimedit.

C: Rocks, big trees

16 Polypodium interjectum Shivas H ros, Paleotrop.

U (Bidighinzos): Shady rocks

Magnoliopsida

Alismatales

Araceae

17 Arisarum vulgare O. Targ. Tozz. G rhiz, Stenomedit.

U (Bonu Trau, Badde Cherchi, Badu Edras): Woods and shrubland (lower altutides)

18 Arum italicum Mill. subsp. italicum G rhiz, Stenomedit.

L: Fringes

19 Arum pictum L. f. G rhiz, Endem. Sa-Co-AT-Bl

RR (Su Pizzu Sa Pedra): at the base of a cliff

Notes: this taxon is not considered as an Italian endemic by Peruzzi et al. (2014)

20 Lemna gibba L. I nat, Subcosmop.

L: Wet places, standing water

21 Lemna minor L. I nat, Subcosmop.

RR (Su Francallossu spring): standing water

Dioscoreales

Dioscoreaceae

22 Dioscorea communis (L.) Caddick & Wilkin G rad, Eurimedit.

C: Woods

Liliales

Colchicaceae

23 Colchicum nanum K. Perss. G bulb, Endem. Sa-Co

L: Wet pastures and meadows

Smilacaceae

24 Smilax aspera L. NP, Subtrop.

C: Woods

Liliaceae

25 Gagea bohemica (Zauschn.) Schult. & Schult.f. G bulb, Eurimedit.

C: Pastures

Asparagales

Orchidaceae

26 Anacamptis laxiflora (Lam.) R. M. Bateman, Pridgeon & M. W. Chase G bulb, Eurimedit.

L: Wet meadows

Specimen examined (syn. Orchis laxiflora Lam.): Funtana Arile, Anela, 08 June 1980, B. Corrias, S. Diana (SS)

27 Anacamptis longicornu (Poir.) R. M. Bateman, Pridgeon & M. W. Chase G bulb, W-Stenomedit.

Not found in the field during this research

Specimen examined (syn. Orchis longicornu Poir.): S’Isfundadu, Anela, 13 May 1965, B. Corrias (SS)

28 Anacamptis papilionacea (L.) R. M. Bateman, Pridgeon & M. W. Chase G bulb, Eurimedit.

C: Dry grasslands

Specimen examined (syn. Orchis papilionacea L.): Funtana Arile, Anela, 08 June 1980, B. Corrias, S. Diana (SS)

29 Dactylorhiza insularis (Sommier) Landwehr G bulb, W-Stenomedit.

Not found in the field during this research

Specimen examined (syn. D. sambucina (L.) Soó): S’Isfundadu, Anela, 13 May 1965, B. Corrias (SS)

30 Limodorum abortivum (L.) Sw. G rhiz, Eurimedit.

U (Littu Majore and Minda ‘e Bassu - Minda ‘e Supra): Quercus ilex woods

31 Orchis provincialis Balb. ex Lam. & DC. G bulb, Stenomedit.

L: Clearings, fringes

Specimens examined: S’Isfundadu, Anela, 13 May 1965, B. Corrias (2 specimens, SS)

32 Serapias lingua L. G bulb, Stenomedit.

L: Wet meadows

Specimen examined: Funtana Arile, Anela, 08 June 1980, B. Corrias, S. Diana (SS)

33 Spiranthes spiralis (L.) Chevall. G rhiz, Europ.-Cauc.

U (Funtana Arile): Wet meadows

Iridaceae

34 Crocus minimus DC. G bulb, Endem. Sa-Co

C: Pastures

35 Iris pseudacorus L. G rhiz, Euras.

U (Su Pranu): Flooded meadows, ponds

36 Romulea columnae Sebast. & Mauri subsp. columnae G bulb, Stenomedit.

C: Pastures

37 Romulea requienii Parl. G bulb, Endem. Sa-Co

C: Pastures

Asphodelaceae

38 Asphodelus ramosus L. subsp. ramosus G rhiz, Stenomedit.

C: Perennial grasslands, pastures, garrigues

Amaryllidaceae

39 Allium chamaemoly L. subsp. chamaemoly G bulb, W-Stenomedit.

L: Annual grasslands (lower altitudes)

40 Allium guttatum subsp. sardoum (Moris) Stearn G bulb, Stenomedit.

C: Pastures, meadows

41 Allium parciflorum Viv. G bulb, Endem. Sa-Co

L: Garrigues, rocky habitats

42 Allium subhirsutum L. G bulb, W-Stenomedit.

C: Perennial grasslands

43 Allium triquetrum L. G bulb, W-Stenomedit.

C: Fringes, woods

44 Allium vineale L. G bulb, Eurimedit.

L: Perennial grasslands

45 Leucojum aestivum subsp. pulchellum (Salisb.) Briq. G bulb, Endem. Sa-Co-Bl

L: Wet meadows

Notes: This taxon is reported also in the Var (Southern France) (see: Tison and de Foucault 2014, Arrigoni 2006–2015; Pignatti 2017–2018) whereas the Euro+Med Plantbase considers it exclusive only in Sardinia, Corsica and the Balearic Islands.

46 Pancratium illyricum L. G bulb, Endem. Sa-Co-AT

L: Garrigues

Asparagaceae

47 Asparagus acutifolius L. G rhiz, Stenomedit.

L: Woods and shrubland (lower altitudes)

48 Drimia pancration (Steinh.) J. C. Manning & Goldblatt G bulb, W-Stenomedit.

L: Grasslands

49 Leopoldia comosa (L.) Parl. G bulb, Eurimedit.

C: Grasslands, pastures

50 Ornithogalum corsicum Jord. & Fourr. G bulb, Endem. Sa-Co

C: Pastures

51 Ornithogalum pyrenaicum L. G bulb, Eurimedit.

C: Deciduous woods

52 Prospero autumnale (L.) Speta G bulb, Eurimedit.

C: Grasslands, pastures

53 Ruscus aculeatus L. G rhiz, Eurimedit.

C: Woods

Poales

Typhaceae

54 Typha angustifolia L. G rhiz, Circumbor.

L: Artificial lake, flooded areas, streams

Juncaceae

55 Juncus articulatus L. G rhiz, Circumbor.

C: Wet meadows, temporary ponds

56 Juncus bufonius L. T caesp, Cosmop.

C: Temporary ponds, wet soils

57 Juncus capitatus Weigel T scap, Medit.-Atl.

C: Temporary ponds

58 Juncus effusus L. subsp. effusus H caesp, Cosmop.

C: Wet meadows, temporary ponds

59 Juncus hybridus Brot. T caesp, Medit.-Atl.

C: Temporary ponds

60 Luzula forsteri (Sm.) DC. H caesp, Eurimedit.

C: Woods

Cyperaceae

61 Carex caryophyllea Latourr. H scap, Euras.

C: Wet pastures and meadows

62 Carex distachya Desf. H caesp, Stenomedit.

C: Woods

63 Carex divisa Huds. G rhiz, Medit.-Atl.

C: Wet meadows and pastures, temporary ponds, ditches

64 Carex divulsa Stockes H caesp, Eurimedit.

C: Fringes

65 Carex microcarpa Moris He, Endem. Sa-Co-AT-Itc

L: Alnus glutinosa woods, riparian vegetation

66 Carex remota L. H caesp, Europ.-Cauc.

U (Badu Addes): Alnus glutinosa wood

67 Cyperus longus L. G rhiz, Paleotemp.

C: Wet meadows, riparian vegetation

Notes: some authors exclude the presence of this species from Sardinia (Desfayes 2004, Arrigoni 2006–2015, Bartolucci et al. 2018) and consider the presence of Cyperus badius Desf. instead. In the Euro+Med Plantbase, C. badius is considered a heterotypic synonym of C. longus.

68 Eleocharis palustris (L.) Roem. & Schult. subsp. palustris G rhiz, Subcosmop.

L: Wet meadows

Gramineae (nom. altr.Poaceae)

69 Aegilops geniculata Roth T scap, Stenomedit.-Turan.

L: Annual grasslands

70 Agrostis capillaris L. H caesp, Circumbor.

L: Wet pastures and meadows

Notes: this taxon is new for the Sardinian flora following Pignatti (1982), Conti et al. (2005), Arrigoni (2006–2015), Pignatti (2017–2018), Bartolucci et al. (2018) and the Euro+Med PlantBase.

71 Aira caryophyllea L. subsp. caryophyllea T scap, Subtrop.

C: Annual grasslands

72 Alopecurus bulbosus Gouan subsp. bulbosus H caesp, Eurimedit.-Subatl.

L: Wet pastures and meadows

73 Anisantha diandra (Roth) Tutin T scap, Eurimedit.

C: Annual grasslands

74 Anisantha madritensis (L.) Nevski subsp. madritensis T scap, Eurimedit.

C: Annual grasslands, pastures

75 Anthoxanthum odoratum L. H caesp, Euras.

C: Wet pastures and meadows

76 Arrhenatherum elatius subsp. sardoum (Em. Schmid) Gamisans H caesp, W-Stenomedit.

L: Garrigues, rocky habitats (higher altitudes)

77 Avena barbata Link subsp. barbata T scap, Eurimedit.

C: Annual grasslands

78 Brachypodium retusum (Pers.) P. Beauv. H caesp, W-Stenomedit.

C: Perennial grasslands on rocky or stony soils

79 Brachypodium sylvaticum (Huds.) P. Beauv. subsp. sylvaticum H caesp, Paleotemp.

C: Woods, fringes

80 Briza maxima L. T scap, Subtrop.

C: Annual grasslands, pastures

81 Briza minor L. T scap, Subcosmop.

U (near Mt. Masiennera): Wet pastures and meadows

82 Bromus hordeaceus L. subsp. hordeaceus T scap, Subcosmop.

C: Annual grasslands, pastures

83 Bromus scoparius L. T scap, Stenomedit.

U (Top of Mt. Masiennera): Annual grasslands

84 Catabrosa aquatica (L.) P. Beauv. G rhiz, Circumbor.

L: Wet soils

85 Cynodon dactylon (L.) Pers. G rhiz, Cosmop.

C: Wet pastures and meadows

86 Cynosurus cristatus L. H caesp, Europ.-Cauc.

C: Wet pastures and meadows

87 Cynosurus echinatus L. T scap, Eurimedit.

C: Annual grasslands, fringes

88 Cynosurus effusus Link T scap, Stenomedit.

C: Annual grasslands, fringes

89 Dactylis glomerata subsp. hispanica (Roth) Nyman H caesp, Stenomedit.

C: Perennial grasslands

90 Danthonia decumbens (L.) DC. subsp. decumbens H caesp, Europ.

L: Wet pastures and meadows

91 Dasypyrum villosum (L.) P. Candargy T Scap, Eurimedit.-Turan.

L: Annual grasslands

92 Festuca morisiana Parl. subsp. morisiana H caesp, Endem. Sa

L: Wet meadows and pastures

93 Glyceria notata Chevall. G rhiz, Subcosmop.

L: Wet habitats

94 Holcus lanatus L. subsp. lanatus H caesp, Circumbor.

C: Wet meadows

95 Hordeum geniculatum All. T scap, Stenomedit.

C: Wet meadows and pastures, temporary ponds

96 Lagurus ovatus L. subsp. ovatus T scap, Eurimedit.

C: Annual grasslands, pastures

97 Lolium perenne L. subsp. perenne H caesp, Euras.

C: Wet pastures

98 Lolium rigidum Gaudin subsp. rigidum T scap, Subtrop.

C: Pastures on arid soil

99 Melica ciliata L. subsp. ciliata H caesp, Eurimedit.

U (Mt. Masiennera): Rocky habitats

100 Melica minuta L. H caesp, Stenomedit.

C: Fringes

101 Melica uniflora Retz. H caesp, Paleotemp.

L: Deciduous woods, fringes

102 Neoschischkinia pourrettii (Willd.) Valdés & H. Scholz T scap, W-Stenomedit.

L: Temporary ponds

103 Piptatherum miliaceum (L.) Coss. subsp. miliaceum H caesp, Stenomedit.

L: Road edges (lower altitudes)

104 Poa annua L. subsp. annua T caesp, Cosmop.

C: Annual grasslands, pastures

105 Poa balbisii Parl. H caesp, Endem. Sa-Co

U (Mt. Masiennera): Garrigues, rocky habitats

106 Poa bulbosa L. subsp. bulbosa H caesp, Paleotemp.

C: Pastures

107 Poa infirma Kunth T caesp, Eurimedit.

C: Mud, wet soils

108 Poa nemoralis L. subsp. nemoralis H caesp, Circumbor.

C: Woods

109 Poa trivialis L. subsp. trivialis H caesp, Euras.

C: Wet meadows

110 Vulpia ligustica (All.) Link T caesp, Stenomedit.

C: Pastures

111 Vulpia myuros (L.) C. C. Gmel. subsp. myuros T caesp, Subcosmop.

C: Pastures

112 Vulpia sicula (C. Presl) Link H caesp, W-Medit.-Mont.

C: Pastures, grasslands

Ranunculales

Papaveraceae

113 Fumaria bastardii Boreau T scap, Subatl.

C: Annual grasslands, fringes

114 Fumaria officinalis L. subsp. officinalis T scap, Paleotemp.

C: Annual grasslands, fringes

115 Papaver rhoeas L. subsp. rhoeas T scap, E-Medit.

C: Pastures, grasslands

Ranunculaceae

116 Anemone hortensis L. subsp. hortensis G bulb, N-Medit.

RR (Su Tattharesu): Perennial grasslands

117 Clematis vitalba L. P lian, Europ.-Cauc.

C: Woods, mantles

118 Ficaria verna Huds. subsp. verna. G bulb, Euras.

C: Woods

119 Ranunculus bulbosus subsp. aleae (Willk.) Rouy & Foucaud H scap, Euras.

C: Grasslands, fringes, woods

120 Ranunculus bullatus L. subsp. bullatus H ros, Stenomedit.

C: Annual grasslands

121 Ranunculus cordiger Viv. subsp. cordiger H scap, Endem. Sa-Co

L: Wet meadows, temporary ponds

122 Ranunculus macrophyllus Desf. H scap, SW-Medit.

L: Wet meadows

123 Ranunculus muricatus L. T scap, Eurimedit.

C: Mud, wet meadows

124 Ranunculus ophioglossifolius Vill. T scap, Eurimedit.

L: Mud, temporary ponds

125 Ranunculus paludosus Poir. subsp. paludosus H scap, Stenomedit.

C: Pastures

126 Ranunculus sardous Crantz T scap, Eurimedit.

C: Mud, temporary ponds

Saxifragales

Paeoniaceae

127 Paeonia corsica Tausch G rhiz, Endem. Sa-Co

L: Woods, clearings

Saxifragaceae

128 Saxifraga tridactylites L. T scap, Eurimedit.

L: Annual grasslands

Crassulaceae

129 Sedum caeruleum L. T scap, SW-Medit.

C: Rocky habitats, annual grasslands

130 Sedum cepaea L. T scap, Submedit.-Subatl.

C: Rocky habitats, annual grasslands

131 Sedum rubens L. T scap, Eurimedit.-Subatl.

C: Rocky habitats, annual grasslands

132 Sedum stellatum L. T scap, Stenomedit.

C: Rocky habitats, annual grasslands

133 Sedum villosum subsp. glandulosum (Moris) P. Fourn. H scap, Endem. Sa-Ag

C: Rocky habitats, annual grasslands

134 Umbilicus rupestris (Salisb.) Dandy subsp. rupestris G bulb, Medit.-Atl.

C: Rocky habitats

Fabales

Leguminosae (nom. altr.Fabaceae)

135 Cytisus villosus Pourr. P caesp, W-Stenomedit.

C: Shrubland, mantles

136 Dorycnium rectum (L.) Ser. H scap, Stenomedit.

L: Wet habitats

137 Genista corsica (Loisel.) DC. NP, Endem. Sa-Co

L: Garrigues on rocky soils

138 Genista desoleana Vals. NP, Endem. Sa-Co-Its

C: Garrigues, dwarf shrubs

Specimens examined: Punta Chelchidores, Anela, 18 July 1972, F. Valsecchi (3 specimens, SS)

139 Lathyrus aphaca L. T scap, Eurimedit.

C: Pastures, fringes

140 Lathyrus sphaericus Retz. T Scap, Eurimedit.

L: Pastures

141 Lotus alpinus (DC.) Ramond H scap, Orof. S-Europ.

C: Wet pastures and meadows

142 Lotus angustissimus L. T scap, Eurimedit.

L: Temporary ponds

143 Lotus conimbricensis Brot. T scap, W- Stenomedit.

C: Annual grasslands

144 Lotus hispidus DC. T scap, W-Medit.

C: Annual grasslands

145 Lupinus angustifolius L. subsp. angustifolius T scap, Stenomedit.

C: Annual grasslands

146 Medicago polymorpha L. T scap, Eurimedit.

C: Pastures, annual grasslands

147 Ononis spinosa L. subsp. spinosa Ch suffr, Eurimedit.

C: Grasslands, pastures

148 Ornithopus compressus L. T scap, Eurimedit.

C: Annual grasslands

149 Ornithopus pinnatus (Mill.) Druce T Scap, Medit.-Atl.

L: Pastures

150 Trifolium angustifolium L. T scap, Eurimedit.

C: Annual grasslands

151 Trifolium arvense L. T scap, Paleotemp.

C: Pastures

152 Trifolium campestre Schreb. T scap, Paleotemp.

C: Annual grasslands

153 Trifolium glomeratum L. T Scap, Eurimedit.

L: Pastures

154 Trifolium incarnatum subsp. molinerii (Hornem.) Syme T scap, Eurimedit.

C: Grasslands, pastures

155 Trifolium micranthum Viv. T scap, Paleotemp.

C: Annual grasslands

156 Trifolium nigrescens Viv. subsp. nigrescens T scap, N-Medit.

C: Pastures

157 Trifolium pratense L. H scap, Eurosib.

C: Wet meadows and pastures

158 Trifolium repens subsp. prostratum Nyman H rept, Eurimedit.

C: Wet meadows and pastures

159 Trifolium spumosum L. T scap, Stenomedit.

C: Annual grasslands

160 Trifolium squarrosum L. T scap, Eurimedit.

L: Pastures

161 Trifolium stellatum L. T scap, Eurimedit.

C: Annual grasslands, pastures

162 Trifolium subterraneum subsp. yanninicum Katzn. & F. H. W. Morley T rept, E-Medit.

C: Pastures

163 Trifolium tomentosum L. T rept, Paleotemp.

C: Annual grasslands, pastures

164 Vicia cracca L. subsp. cracca H scap, Euras.

C: Fringes

165 Vicia lathyroides L. T scap, Eurimedit.

C: Fringes

166 Vicia lutea L. subsp. lutea T scap, Eurimedit.

C: Fringes

167 Vicia villosa subsp. ambigua (Guss.) Kerguélen H Scap, W-Stenomedit.

L: Fringes

168 Vicia villosa Roth subsp. villosa T scap, Eurimedit.

C: Fringes

Rosales

Rosaceae

169 Agrimonia eupatoria L. subsp. eupatoria H scap, Subcosmop.

C: Fringes

170 Crataegus monogyna Jacq. P caesp, Paleotemp.

C: Shrublands, woods, mantles

171 Fragaria vesca L. subsp. vesca H rept, Eurosib.

C: Deciduous woods, fringes

172 Geum urbanum L. H scap, Circumbor.

C: Deciduous woods, fringes

Specimen examined: Caserma Forestale Anela, sine die, Barba (SS)

173 Malus pumila Mill. P scap, CW-Euras.

L: Woods, mantles

Notes: in accordance with Bagella and Urbani (2006), this is the valid name for Malus domestica Borkh. (nom. illeg.), also reported in the Euro+Med PlantBase. Yet Galasso et al. (2018) call a taxon Malus domestica, considering it as a non-native species, while Camarda and Valsecchi (2008), Arrigoni (2006–2015) and Pignatti (2017–2018) still call it M. dasyphylla. Finally, Bartolucci et al. (2018) report the taxon M. sylvestris in Sardinia. Malus pumila is reported as a synonym of M. domestica by Galasso et al. (2018), it is excluded from the Sardinian flora by Arrigoni (2006–2015), finally, it was not mentioned by Camarda and Valsecchi (2008). In the Euro+Med Plantbase, Malus pumila Mill. is the valid name for Malus domestica Borkh. The populations we have examined in the Marghine-Goceano range (not only the forest domain of Anela) have the characters of Malus domestica, not M. sylvestris.

174 Potentilla reptans L. H ros, Paleotemp.

C: Wet meadows

175 Prunus avium (L.) L. P scap, Pontic

L: Woods

176 Prunus domestica subsp. insititia (L.) Bonnier & Layens P scap

U (Su Cantareddu): Mantles

177 Prunus spinosa L. subsp. spinosa P caesp, Europ.-Cauc.

C: Shrublands

178 Pyrus communis subsp. pyraster (L.) Ehrh. P scap, Euras.

L: Woods, mantles

179 Pyrus spinosa Forssk. P caesp, Stenomedit.

C: Shrublands, mantles, woods

180 Rosa canina L. NP, Paleotemp.

C: Shrublands

181 Rosa sempervirens L. NP, Stenomedit.

L: Woods, shrublands (lower altitudes)

182 Rosa subcanina (Christ) Vuk. NP, Europ.

C: Shrublands

183 Rubus ulmifolius Schott NP, Eurimedit.

C: Shrublands, woods

184 Sanguisorba minor subsp. balearica (Bourg. ex Nyman) Muñoz Garm. & C. Navarro H scap, Eurimedit.

C: Grasslands

Ulmaceae

185 Ulmus minor Mill. subsp. minor P caesp, Europ.-Cauc.

L: Woods

Cannabaceae

186 Celtis australis L. subsp. australis P scap, Eurimedit.

RR (Pedru Addes): Wood edge

Moraceae

187 Ficus carica L. subsp. carica P scap, Medit.-Turan.

U (Badu Edras): Riparian vegetation

Urticaceae

188 Parietaria lusitanica L. subsp. lusitanica T rept, Stenomedit.

C: Buildings, fringes

189 Urtica atrovirens Loisel. H scap, Endem. Sa-Co-Bl-AT-Itc

L: Ruderal vegetation

190 Urtica dioica L. subsp. dioica H scap, Subcosmop.

C: Ruderal vegetation

Fagales

Fagaceae

191 Quercus ilex L. P scap, Stenomedit.

C: Woods

192 Quercus pubescens Willd. agg. P caesp, SE-Europ.

C: Woods

Notes: There are many controversial treatments for describing the variation within Q. pubescens (Mossa et al. 1998, 1999). Until the various treatments are resolved, we prefer to treat this variation as a complex (or aggregate) within Q. pubescens s.l.

193 Quercus suber L. P scap, W-Eurimedit.

L: Woods

Betulaceae

194 Alnus glutinosa (L.) Gaertn. subsp. glutinosa P scap, Paleotemp.

L: Streams, wet places, springs

Oxalidales

Oxalidaceae

195 Oxalis corniculata L. subsp. corniculata H rept, Eurimedit.

L: Walls, buildings

Malpighiales

Guttiferae (nom. altr.Clusiaceae)

196 Hypericum androsaemum L. NP, W-Eurimedit.-Subatl.

L: Wet habitats, springs

197 Hypericum hircinum L. subsp. hircinum NP, Endem. Sa-Co-AT

L: Springs, streams, Alnus glutinosa woods

Notes: H. hircinum includes several subspecies, amongst which the subsp. hircinum is exclusive of Sardinia, Corsica and the Tuscan Archipelago (Carta and Peruzzi 2015)

198 Hypericum perforatum L. subsp. perforatum H scap, Paleotemp.

C: Fringes, road edges

Violaceae

199 Viola alba subsp. dehnhardtii (Ten.) W. Becker H ros, Eurimedit.

C: Woods, fringes

200 Viola reichenbachiana Jord. ex Boreau H scap, Eurosib.

C: Deciduous woods

Notes: it was excluded for the Sardinian flora by Arrigoni (2006–2015), but later confirmed by Mereu (2012) for the Gennargentu massif

Salicaceae

201 Salix cinerea subsp. oleifolia Macreight P caesp, W-Medit.-Atl.

L: Streams, springs

202 Salix purpurea L. P scap, Euras.

L: Ditches

Euphorbiaceae

203 Euphorbia characias L. subsp. characias NP, Stenomedit.

C: Woods, shrublands (lower altitudes)

204 Euphorbia helioscopia L. subsp. helioscopia T scap, Cosmop.

C: Annual grasslands

205 Euphorbia pithyusa subsp. cupanii (Guss. ex Bertol.) Radcl.-Sm. G rhiz, Endem. Sa-Co-Si

C: Perennial grasslands, pastures

206 Euphorbia semiperfoliata Viv. G rhiz, Endem. Sa-Co

L: Woods, fringes

Linaceae

207 Linum bienne Mill. H bienn, Eurimedit.

C: Annual grasslands

Geraniales

Geraniaceae

208 Erodium chium (L.) Willd. T scap, Eurimedit.

L: Pastures

209 Erodium ciconium (L.) L’Hér. T scap, Eurimedit.-Pontic

C: Pastures

210 Erodium cicutarium (L.) L’Hér. T scap, Subcosmop.

C: Pastures

211 Geranium purpureum Vill. T scap, Eurimedit.

C: Woods, fringes

212 Geranium robertianum L. T scap, Subcosmop.

C: Woods, fringes

213 Geranium rotundifolium L. T scap, Paleotemp.

C: Woods, fringes

Myrtales

Lythraceae

214 Lythrum portula (L.) D. A. Webb T rept, S-Europ.-S-Sib.

L: Temporary ponds

Onagraceae

215 Epilobium montanum L. H scap, Euras.

C: Woods

Sapindales

Sapindaceae

216 Acer monspessulanum L. subsp. monspessulanum P caesp, Eurimedit.

L: Woods and mantles

Malvales

Malvaceae

217 Althaea hirsuta L. T scap, Eurimedit.

L: Annual grasslands

218 Malva olbia (L.) Alef. P caesp, Stenomedit.

C: Shrublands on wet soils

219 Malva sylvestris L. H scap, Eurosib.

C: Grasslands, fringes

Cistaceae

220 Cistus monspeliensis L. NP, Stenomedit.

C: Garrigues (lower altitudes)

221 Cistus salviifolius L. NP, Stenomedit.

C: Garrigues

222 Tuberaria guttata (L.) Fourr. T scap, Eurimedit.

C: Annual grasslands

Brassicales

Resedaceae

223 Sesamoides purpurascens subsp. spathulata (Moris) Lambinon & Kerguélen H Scap, W-Medit.-Mont.

C: Dirty tracks, trampled places

Cruciferae (nom. altr.Brassicaceae)

224 Arabidopsis thaliana (L.) Heynh. T scap, Paleotemp.

C: Annual grasslands, pastures

225 Capsella bursa pastoris (L.) Medik. subsp. bursa-pastoris H bienn, Cosmop.

C: Annual grasslands, pastures

226 Cardamine flexuosa With. H scap, Circumbor.

C: Fringes

227 Cardamine hirsuta L. T scap, Cosmop.

C: Fringes

228 Draba muralis L. T scap, Circumbor.

L: Cliffs, road edges

229 Erophila verna subsp. praecox (Steven) Walters T scap, Eurimedit.

C: Annual grasslands

230 Morisia monanthos (Viv.) Asch. H ros, Endem. Sa-Co

U (Near Mt. Masiennera): Wet meadows

231 Nasturtium officinale (L.) R. Br. H scap, Cosmop.

L: Muds, streams

232 Raphanus raphanistrum L. subsp. raphanistrum T scap, Eurimedit.

C: Grasslands

233 Sisymbrium officinale (L.) Scop. T scap, Paleotemp.

C: Pastures

234 Teesdalia coronopifolia (J.P. Bergeret) Thell. T scap, Eurimedit.

C: Pastures

Santalales

Santalaceae

235 Osyris alba L. NP, Eurimedit.

L: Woods, clearings, rocky habitats

Caryophyllales

Plumbaginaceae

236 Armeria sardoa Spreng. subsp. sardoa Ch suffr, Endem. Sa

L: Garrigues, rocky habitats

Polygonaceae

237 Rumex bucephalophorus L. subsp. bucephalophorus T scap, Eurimedit.-Macaron.

C: Annual grasslands

238 Rumex crispus L. H scap, Subcosmop.

C: Wet meadows

239 Rumex pulcher L. subsp. pulcher H scap, Eurimedit.

C: Wet meadows

240 Rumex scutatus subsp. glaucescens (Guss.) Brullo, Scelsi & Spamp. H scap, Endem. Sa-Si

L: Rocky habitats

241 Rumex thyrsoides Desf. H scap, W-Medit.

C: Fringes

Caryophyllaceae

242 Arenaria balearica L. Ch suffr, Endem. Sa-Co-Bl-AT

L: Shady rocks and cliffs

Specimens examined: S’Isfundadu, Anela, 25 May 1966, B. Corrias (2 specimens, SS); S’Isfundadu, Anela, 18 June 1965, F. Valsecchi (1 specimen, SS).

243 Cerastium gibraltaricum Boiss. Ch suffr, Orof. W-Medit.

L: Garrigues

Notes: in the Euro+Med Plantbase, Cerastium boissierianum Greuter et Burdet is considered a synonym of C. gibraltaricum

244 Cerastium glomeratum Thuill. T scap, Eurimedit.

C: Pastures

245 Cerastium ligusticum subsp. palustre (Moris) P. D. Sell et Whitehead T scap, Endem. Sa-Co

RR (near Mt. Masiennera): Wet pastures and meadows

246 Corrigiola telephiifolia Pourr. H Ros, W-Medit.

L: Trampled sites, dirty roads

Specimen examined: Badu Addes, Anela, September 1962 (sine die), sine coll. (SS)

247 Dianthus ichnusae subsp. toddei Bacch., Brullo, Casti et Giusso H scap, Endem. Sa

L: Garrigues, rocky habitats

Notes: this taxon is exclusive for the Goceano mountain range (Bacchetta et al. 2010).

248 Moenchia erecta (L.) P. Gaertn., B. Mey. & Scherb. subsp. erecta T scap, Medit.-Atl.

C: Pastures

249 Petrorhagia dubia (Raf.) G. López & Romo T scap, S-Medit.

C: Pastures

250 Petrorhagia saxifraga (L.) Link H caesp, Eurimedit.

C: Garrigues, rocky habitats

251 Sagina apetala Ard. T scap, Eurimedit.

L: Annual grasslands, dirty tracks

252 Sagina procumbens L. H caesp, Subcosmop.

L: Wet places, springs

Specimen seen: Badu Addes, Anela, sine die, Barba (SS)

253 Sagina subulata (Sw.) C. Presl H caesp, Medit.-Atl.

L: Wet meadows, rocky habitats (higher altitudes)

Notes: for this taxon, recently the name S. alexandrae Iamonico has been proposed (Iamonico 2016)

254 Silene gallica L. T scap, Eurimedit.

C: Pastures

255 Silene laeta (Aiton) Godr. T scap, W-Stenomedit.

L: Muddy places, wet meadows, temporary ponds

256 Silene latifolia Poir. H bienn, Paleotemp.

C: Fringes

257 Silene vulgaris (Moench) Garcke subsp. vulgaris H scap, Paleotemp.

C: Fringes

258 Spergula arvensis L. T scap, Subcosmop.

C: Pastures

259 Stellaria media (L.) Cirillo subsp. media T rept, Cosmop.

C: Ruderal vegetation, woods, fringes

Amaranthaceae

260 Chenopodium album L. subsp. album T Scap, Subcosmop.

Not found in the field during this research

Specimens examined: Badu Addes, Anela, 09 September 1962, Barba (2 specimens, SS).

Portulacaceae

261 Montia fontana subsp. amporitana Sennen T scap, Medit-Mont. Subatl.

C: Mud, flooded soils

Ericales

Primulaceae

262 Anagallis arvensis L. T rept, Eurimedit.

C: Annual grasslands

263 Asterolinon linum-stellatum (L.) Duby T Scap, Stenomedit.

L: Annual grasslands, pastures

264 Cyclamen repandum Sibth. & Sm. subsp. repandum G bulb, NW-Stenomedit.

C: Woods

Ericaceae

265 Arbutus unedo L. P caesp, Stenomedit.

RR (Littu Majore): Wood

266 Erica arborea L. P caesp, Stenomedit.

C: Shrublands, woods

Gentianales

Rubiaceae

267 Cruciata glabra (L.) Ehrend. H scap, Euras.

C: Grasslands, pastures

Specimen examined: Badu Addes, Anela, 18 July 1972, B. Corrias, S. Diana, F. Valsecchi (SS).

268 Galium aparine L. subsp. aparine T scap, Euras.

C: Fringes

269 Galium corsicum Spreng. H scap, Endem. Sa-Co

L: Rocky habitats

270 Galium debile Desv. H scap, Eurimedit.

L: Wet habitats

271 Galium rotundifolium L. H scap, Orof.-W-Euras.

L: Woods (higher altitudes)

272 Rubia peregrina L. subsp. peregrina P lian, Stenomedit.-Macaron.

C: Woods

273 Sherardia arvensis L. T scap, Eurimedit.

C: Pastures, annual grasslands

274 Theligonum cynocrambe L. T scap, Stenomedit.

C: Annual grasslands, fringes

Gentianaceae

275 Exaculum pusillum (Lam.) Caruel T scap, W-Eurimedit.

RR (Minda ‘e Bassu): Temporary pond

Boraginales

Boraginaceae

276 Anchusa hybrida Ten. H scap, Stenomedit.

Not found in the field during this research

Specimens examined: Badu Addes, Anela, 22 October 1963, F. Valsecchi, Barba (3 specimens, SS).

277 Cynoglossum creticum Mill. H bienn, Eurimedit.

L: Fringes

278 Echium plantagineum L. T Scap, Eurimedit.

C: Pastures, grasslands

279 Myosotis arvensis (L.) Hill subsp. arvensis T scap, Europ.-W-Asian

C: Annual grasslands, pastures

280 Myosotis sicula Guss. T scap, N-Eurimedit.

L: Wet meadows, temporary ponds

Convolvulaceae

281 Convolvulus althaeoides L. H scand, Stenomedit.

C: Perennial grasslands

282 Convolvulus arvensis L. G rhiz, Paleotemp.

C: Perennial grasslands

283 Cuscuta epithymum subsp. corsicana (Yunck.) Lambinon T par, Endem. Sa-Co

L: Garrigues (mainly parasite on Genista desoleana)

Solanales

Solanaceae

284 Solanum dulcamara L. NP, Paleotemp.

U (Su Pranu): Riparian vegetation

Lamiales

Oleaceae

285 Phillyrea latifolia L. P caesp, Stenomedit.

C: Woods, shrubland (lower altitude)

Plantaginaceae

286 Callitriche stagnalis Scop. I rad, Euras.

L: Temporary ponds, springs, muddy soils

287 Cymbalaria aequitriloba (Viv.) A. Chev. subsp. aequitriloba Ch rept, Endem. Sa-Co-Bl-AT

L: Shady rocks and cliffs

288 Digitalis purpurea L. subsp. purpurea H scap, W-Eurimedit.

C: Fringes, clearings

289 Linaria arvensis (L.) Desf. T scap, Submedit.-Subatl.

C: Annual grasslands

290 Linaria pelisseriana (L.) Mill. T scap, Medit.-Atl.

C: Pastures

291 Plantago coronopus L. T scap, Eurimedit.

C: Grasslands, pastures

292 Plantago lagopus L. subsp. lagopus T scap, Stenomedit.

C: Annual grasslands, pastures

293 Plantago lanceolata L. H ros, Euras.

C: Grasslands

294 Plantago major L. subsp. major H ros, Euras.

L: Wet meadows

295 Plantago weldenii Rchb. T scap, Stenomedit.

C: Annual grasslands

296 Veronica anagallis-aquatica L. subsp. anagallis-aquatica H scap, Cosmop.

L: Mud, springs, ditches

Specimen examined: Punta Chelchidores est, Anela, 18 July 1972, B. Corrias, S. Diana, F. Valsecchi (SS)

297 Veronica arvensis L. T scap, Subcosmop.

C: nitrophilous vegetation

298 Veronica hederifolia L. subsp. hederifolia T scap, Euras.

C: Woods, fringes

299 Veronica verna subsp. brevistyla (Moris) Rouy T scap, Endem. Sa-Co

L: Pastures (higher altitudes)

Scrophulariaceae

300 Scrophularia trifoliata L. H caesp, Endem. Sa-Co-AT

L: Rocky habitats

Specimen examined: Badu Addes, Anela, 18 July 1972, F. Valsecchi (SS)

301 Scrophularia umbrosa Dumort. subsp. umbrosa H Scap, Euras.

Not found in the field during this research

Specimens examined: Badu Addes, Anela, 18 July 1973, F. Valsecchi (3 specimens, SS)

302 Verbascum pulverulentum Vill. H bienn, Europ.

C: Clearings, fringes

Labiatae (nom. altr.Lamiaceae)

303 Clinopodium nepeta subsp. glandulosum (Req.) Govaert H scap, Stenomedit.

C: Fringes

304 Clinopodium vulgare subsp. orientale Bothmer H scap, E-Stenomedit.

C: Fringes

Notes: The Italian Flora Checklists (Conti et al. 2005, Bartolucci et al. 2018) consider the subsp. arundanum (Boiss.) Nyman as present in Sardinia, whereas, the Euro+Med PlantBase considers subsp. arundanum absent from the island (and the whole Italian peninsula) and that, instead, subsp. orientale is present. Our specimens fit well with the diagnostic characters of subsp. orientale as described by Bothmer (1967).

305 Glechoma sardoa (Bég.) Bég. H rept, Endem. Sa

L: Woods, fringes

306 Lamium maculatum (L.) L. H scap, Euras.

U: Forest near forestry headquarters, under Quercus ilex

Notes: according to Arrigoni (2006–2015), this taxon was not found in Sardinia in recent years

307 Lamium purpureum L. T scap, Euras.

C: Fringes

308 Lavandula stoechas L. subsp. stoechas NP, Stenomedit.

C: Garrigues

309 Mentha aquatica L. H scap, Paleotemp.

L: Wet meadows

310 Mentha pulegium L. subsp. pulegium H scap, Eurimedit.

C: Wet meadows, temporary ponds

311 Mentha requienii Benth. subsp. requienii H rept, Endem. Sa-Co

RR (Su Cantareddu spring): Wet rocks, spring

312 Mentha suaveolens subsp. insularis (Req. ex Gren. & Godr.) Greuter H scap, Endem. Sa-Co-AT-Bl

U (Funtana Arile spring): Fringes

313 Micromeria graeca (L.) Benth. subsp. graeca Ch suffr, Stenomedit.

C: Garrigues

314 Prunella vulgaris L. subsp. vulgaris H scap, Circumbor.

C: Wet meadows, fringes, clearings

315 Salvia verbenaca L. H scap, Medit.-Atl.

C: Grasslands

Notes: following the Euro+Med PlantBase, in this taxon we include ecotypes referred to Salvia clandestina L.

316 Stachys arvensis (L.) L. T scap, Europ.

L: Annual grasslands, pastures

317 Stachys corsica Pers. H rept, Endem. Sa-Co

L: Shady rocks and cliffs

Specimens examined: S’Isfundadu, Anela, 18 June 1965, F. Valsecchi (SS); Badu Addes, Anela, 18 July 1972, B. Corrias, S. Diana, F. Valsecchi (SS)

318 Stachys glutinosa L. Ch frut, Endem. Sa-Co-AT

L: Garrigues, rocky habitats

319 Teucrium chamaedrys L. subsp. chamaedrys Ch suffr, Eurimedit.

U (near the helicopter base): Pastures, grasslands

320 Thymus herba-barona Loisel. Ch rept, Endem. Sa-Co-Bl

C: Garrigues

Orobanchaceae

321 Orobanche hederae Duby T par, Eurimedit.

C: Woods

322 Orobanche minor Sm. T par, Paleotemp.

C: Grasslands, pastures

323 Orobanche nana (Reut.) Beck T par, Medit.-Macaron.

L: Grasslands, pastures

324 Orobanche ramosa L. T par, Paleotemp.

L: Road sides, pastures

325 Orobanche rapum-genistae Thuill. T par, Subatl.

L: Garrigues with Genista sp.

326 Orobanche rigens Loisel. T par, Endem. Sa-Co

L: Garrigues with Genista sp.

327 Parentucellia latifolia (L.) Caruel subsp. latifolia T scap, Eurimedit.

C: Pastures

328 Parentucellia viscosa (L.) Caruel T scap, Medit.-Atl.

C: Annual grasslands

Aquifoliales

Aquifoliaceae

329 Ilex aquifolium L. P caesp, Submedit.-Subatl.

C: Woods

Asterales

Campanulaceae

330 Jasione montana L. H scap, Europ.-Cauc.

C: Pastures and rocky habitats

Compositae (nom. altr.Asteraceae)

331 Achillea ligustica All. H scap, W-Stenomedit.

C: Fringes

332 Anthemis arvensis L. subsp. arvensis T scap, Stenomedit.

C: Pastures

333 Arctium minus (Hill) Bernh. H bienn, Eurimedit.

C: Fringes, clearings

334 Bellis annua L. subsp. annua T scap, Stenomedit.

C: Annual grasslands on wet soils

335 Bellis perennis L. H ros, Europ.-Cauc.

C: Wet meadows

336 Bellis sylvestris Cirillo H ros, Stenomedit.

L: Perennial grasslands (lower altutides)

337 Bellium bellidioides L. H ros, Endem. Sa-Co-Bl-AT

C: Temporary ponds, wet soils

338 Carlina corymbosa L. H scap, Stenomedit.

C: Pastures

339 Carthamus lanatus L. subsp. lanatus T scap, Eurimedit.

C: Pastures, nitrophilous vegetation near sheep pens

340 Centaurea calcitrapa L. subsp. calcitrapa H bienn, Eurimedit.

C: Pastures

341 Chamaemelum fuscatum (Brot.) Vasc. T scap, W-Stenomedit.

L: Temporary ponds

342 Chondrilla juncea L. H scap, S-Europ.-S-Sib.

C: Pastures

343 Cichorium intybus L. subsp. intybus H scap, Paleotemp.

L: Perennial grasslands

344 Cirsium scabrum (Poir.) Bonnet & Barratte H scap, SW-Medit.

L: Fringes, road edges (lower altitudes)

345 Cirsium vulgare subsp. silvaticum (Tausch) Arènes H bienn, Eurimedit.

C: Fringes, road edges

346 Crepis bellidifolia Loisel. T scap, W-Stenomedit.

L: Pastures

347 Crepis leontodontoides All. H ros, W-Medit.-Mont.

C: Pastures

348 Crepis vesicaria L. subsp. vesicaria T scap, Submedit.-Subatl.

C: Pastures

349 Crupina vulgaris Cass. T scap, S-Sib.-Eurimedit.

L: Pastures, perennial grasslands

350 Filago gallica L. T scap, Eurimedit.

C: Annual grasslands

351 Filago germanica (L.) Huds. T scap, Paleotemp.

U (S. Giorgio): Annual grasslands

352 Galactites tomentosus Moench H bienn, Stenomedit.

C: Pastures

353 Glebionis coronaria (L.) Spach. T scap, Stenomedit.

L: Pastures, annual grasslands (lower altitudes)

354 Helichrysum italicum subsp. tyrrhenicum (Bacch., Brullo et Giusso) Herrando, J.M. Blanco, L. Sáez & Galbany Ch frut., Endem. Sa-Co-Bl

C: Garrigues

Notes: for this taxon, we follow Herrando-Moraira et al. (2016)

355 Hyoseris radiata L. H ros, Stenomedit.

C: Pastures, meadows

356 Hypochaeris achyrophorus L. T scap, Stenomedit.

C: Annual grasslands

357 Hypochaeris cretensis (L.) Bory & Chaub. H scap, NE-Medit.-Mont.

L: Dry pastures and rocky habitats

358 Hypochaeris glabra L. T scap, Eurimedit.

C: Pastures, meadows

359 Hypochaeris radicata L. subsp. radicata H ros, Europ.-Cauc.

C: Pastures, meadows

360 Hypochaeris robertia (Sch. Bip.) Fiori H ros, Endem. Sa-Co-Si-It

L: Wet rocks and cliffs

361 Lactuca muralis (L.) Gaertn. H scap, Europ.-Cauc.

C: Woods, fringes

362 Leontodon tuberosus L. H ros, Stenomedit.

C: Grasslands, pastures

363 Pilosella ziziana (Tausch) F. W. Schultz & Sch. Bip. H scap, Europ. (?)

L: Grasslands

364 Ptilostemon casabonae (L.) Greuter H scap, Endem. Sa-Co-AT-Hy

U (Entrance of the Domain): Road edge

365 Pulicaria odora (L.) Rchb. H scap, Eurimedit.

C: Woods, fringes (lower altitude)

366 Reichardia picroides (L.) Roth H scap, Stenomedit.

L: Rocky habitats (lower altitudes)

367 Rhagadiolus stellatus (L.) Gaertn. T scap, Eurimedit.

C: Annual grasslands

368 Scolymus hispanicus L. subsp. hispanicus H bienn, Eurimedit.

C: Pastures

369 Senecio vulgaris L. subsp. vulgaris T scap, Eurimedit.

C: Pastures, ruderal vegetation

370 Sonchus asper (L.) Hill subsp. asper T scap, Euras.

C: Ruderal vegetation

371 Sonchus oleraceus L. T scap, Euras.

C: Ruderal vegetation

372 Silybum marianum (L.) Gaertn. H bienn, Medit.-Turan.

C: Ruderal vegetation, pastures

373 Taraxacum sect. Erythrosperma (H. Lindb.) Dahlst. or Taraxacum sect. Scariosa Hand.-Mazz. H ros, Circumbor.

C: Wet meadows

374 Urospermum dalechampii (L.) F.W. Schmidt H scap, Eurimedit.

C: Grasslands

Dipsacales

Adoxaceae

375 Sambucus ebulus L. G rhiz, Eurimedit.

L: Streams

376 Sambucus nigra L. P caesp, Europ.-Cauc.

C: Woods, shrublands

Caprifoliaceae

377 Dipsacus ferox Loisel. H bienn, Endem. Sa-Co-Itc

C: Pastures

378 Valerianella eriocarpa Desv. T scap, Stenomedit.

C: Annual grasslands

Apiales

Araliaceae

379 Hedera helix L. P lian, Eurimedit.

C: Woods

Umbelliferae (nom. altr.Apiaceae)

380 Bunium corydalinum DC. G bulb, Endem. Sa-Co

C: Garrigues, rocky habitats

381 Chaerophyllum temulum L. T scap, Euras.

L: Woods, fringes

382 Eryngium campestre L. H scap, Eurimedit.

C: Pastures

383 Ferula communis L. subsp. communis H scap, S-Eurimedit.

L: Pastures, clearings (lower altitudes)

384 Oenanthe crocata L. H scap, Medit.-Atl.

L: Alnus glutinosa woods, streams

385 Oenanthe lisae Moris H scap, Endem. Sa

U (Funtana Arile spring): Wet meadows

Specimen examined: Funtana Arile, Anela, 08 June 1980, B. Corrias, S. Diana (SS)

386 Oenanthe pimpinelloides L. H scap, Medit.-Atl.

C: Woods, fringes

387 Sanicula europaea L. H scap, Paleotemp.

C: Woods, fringes

388 Smyrnium perfoliatum subsp. rotundifolium (Mill.) Bonnier & Layens H bienn, Stenomedit.

C: Fringes, woods

389 Thapsia garganica L. subsp. garganica H scap, S-Medit.

C: Pastures, grasslands

390 Torilis africana Spreng. T scap, Medit.-Macaron.

C: Pastures, annual grasslands

391 Torilis nodosa (L.) Gaertn. T scap, Medit.-Turan.

C: Pastures, annual grasslands

Ecological and biogeographical analysis of the indigenous flora of Anela

Here we assess the presence in the forest domain of Anela of 391 taxa, belonging to 32 orders and 74 families.

Of the listed taxa, 5 (Anacamptis longicornu (Orchidaceae), Anchusa hybrida (Boraginaceae), Chenopodium album subsp. album (Amaranthaceae), Dactylorhiza insularis (Orchidaceae), Scrophularia umbrosa (Scrophulariaceae)) were not found during our investigation. Excluding these species, then we recorded a total of 386 indigenous taxa within the domain. Two species are new for the Sardinian flora (Agrostis capillaris, Asplenium adiantum-nigrum) and, for 17 taxa, our findings determine an important enlargement of their known range on the island (Arrhenatherum elatius subsp. sardoum, Asplenium foreziense, Clinopodium vulgare subsp. orientale, Colchicum nanum, Danthonia decumbens subsp. decumbens, Euphorbia semiperfoliata, Exaculum pusillum, Festuca morisiana subsp. morisiana, Lamium maculatum, Mentha requienii subsp. requienii, Morisia monanthos, Poa balbisii, Prunus domestica subsp. insititia, Ranunculus cordiger subsp. cordiger, Rosa subcanina, Veronica verna subsp. brevistyla, Viola reichenbachiana).

Overall, we found 141 hemicryptophytes (36.1%), 137 therophytes (35.0%), 56 geophytes (14.3%), 27 phanaerophytes (6.9%), 15 nano-phanaerophytes (3.8%), 11 chamaephytes (2.8%), 3 hydrophytes (0.8%), and 1 helophyte (0.3%).

A total of 239 taxa belong to the Mediterranean element (61.1%), 53 are Eurasian sensu lato (including the true Eurasian, plus European, Euro-Siberian, Euro-Caucasian and Pontic district: overall 13.6%), 42 are Boreal-Temperate taxa (paleotemperate + circumboreal: 10.7%), 36 are widespread (cosmopolitan, sub-cosmopolitan and sub-tropical: 9.2%) and 19 are Atlantic (4.9%). We were not able to assign a geographical category to Prunus domestica subsp. insititia.

Hemicryptophytes dominate within the Boreal-Temperate and the Eurasian components; annual species prevail within the widespread and the Mediterranean-Atlantic groups. The Mediterranean component hosts similar percentages of annuals and hemicryptophytes (Fig. 2).

The Mediterranean component is dominated by the euri-Mediterranean sub-element (94 taxa, 24.0% of the whole flora), followed by the steno-Mediterranean (77 taxa, 19.7%) and the endemics (45 entities, 11.5%). A total of 23 Mediterranean taxa belonged to other chorotypes (mountain-Mediterranean, Mediterranean-Turanian, Mediterranen-Macaronesian).

Figure 2.

Percentage of biological types for each chorologic element detected in the vascular flora of Anela (390 taxa). boreo.temp = Boreal-temperate taxa; med.atlan = Mediterranean-Atlantic taxa; med = Mediterranean; nd = not determined.

The endemic component of the flora of Anela is dominated by those of the Sardinian-Corsican biogeographic province (sensuBacchetta et al. 2012) accounting for 28 taxa (endemics sensu stricto, 7.4%), of which 19 taxa are Sardinian-Corsican (42.2% of the endemic component), followed by Sardinian entities (5, 11.1%) and those present on Sardinia, Corsica and the Tuscan Archipelago (4, 8.9%). Tyrrhenian or Hercynian endemics (those present in Sardinia, Corsica, Tuscan Archipelago, the Balearic and Hyeres Islands and Sicily) account 12 (26.7%) and, finally, 11.1% is constituted by 5 entities with larger ranges including some continental areas (Sardinia and northern Africa or Sardinia and Italy).

On the basis of our criteria, 241 taxa (61.6%) can be considered common at the local level, 113 (28.9%) are localised, 23 (5.9%) are uncommon, 9 (2.3%) are range restricted and 5 (1.3%) are locally extinct in the last 50 years. Common taxa are the dominant category in all the geographic groups, whereas range restricted taxa are found only in the widespread, Boreal-Temperate and the Mediterranean groups (Fig. 3).

A total of 176 out of 387 taxa were found mainly in grasslands habitats (45.5%) including dry pastures (61 taxa), annual and perennial grasslands (52 and 31 taxa, respectively) and wet pastures and meadows (32 taxa). Woodland habitats hosted 97 taxa (25.1%), comprising woods (57 taxa), fringes and clearings (30 taxa) and shrubs (10 taxa). Wet habitats (including Alnus glutinosa woods, springs, temporary ponds, ditches, muds, streams) hosted 53 taxa (13.7%). Rocky habitats (cliffs, rocks, screes) harbour 24 taxa (6.2%), then the garrigues hosted 21 taxa (5.4%) and finally the anthropogenic habitats (ruderal vegetation, buildings, walls, trampled sites, road edges) were the main habitat for 15 taxa (3.9%).

Figure 3.

Percentage of abundance categories for each chorologic element detected in the vascular flora of Anela (390 taxa). c = common; l = localized; u = uncommon; rr = range restricted; ex = extinct. ; boreo.temp = Boreal-temperate taxa; med.atlan = Mediterranean-Atlantic taxa; med = Mediterranean; nd = not determined.

Discussion

Biogeographical description of the mountain

Our research discovered a high species density at the study area (30.6 taxa km^-2), that is one of the highest ever documented in the Sardinian mountain floras (Table 1). Even if there is a clear inverse relationship between the area investigated and species’ density, we should note that, for areas having a comparable surface (~ 10 km²), the floristic density recorded at our study area is second only to the Mt. Gonare complex (Camarda 1984a, 1984b). It is noteworthy that the summit area of Sardinia (> 1500 m a.s.l.), having a surface of 16.8 km², hosts “only” 214 taxa of which 66 are considered endemics (Arrigoni and Camarda 2015). So we can argue that areas at the edge between the Mediterranean and the temperate bioclimates, like Foresta Demaniale Anela and Mt. Gonare, host floristic components from both the two bioclimatic – biogeographic regions, having therefore more abundant floras than areas located in coastal or summit zones.

The hemicryptophytes/therophytes (H/T) ratio, as previously noted by Arrigoni and Camarda (2015), underlines the co-presence of two main elements, the perennial and the annual herbs, having very different life-cycles and summing 71.1% of our flora.The H/T ratio, that in Sardinia peaks at 2.5 at the summit of Gennargentu (Arrigoni and Camarda 2015), but decreases to 0.74 as the regional average, is at Anela 1.03. Limestone mountains like Mt. Albo, with a karst geology and consequently a pronounced summer drought, have a H/T ratio even lower than the regional average, whereas mountain complexes with impermeable substrates (plutonic, volcanic, metamorphic) approaching 1000 m a.s.l. have a H/T ratio ~ 1 gradually increasing with elevation (Table 1). This means that at 1000 m a.s.l., the co-presence of two large groups of non-woody plants, having an annual or perennial life cycle, has been detected: the annuals have a greater prevalence at lower altitudes, the perennials at higher altitudes and their ratio ~ 1 at 1000 m a.s.l. underlines the transition character of this altimetric level in Sardinia.

Important differences with the regional (Sardinian) value (Pignatti 1995) have also been detected for the Mediterranean floristic component, particularly the steno-Mediterranean taxa having a 28.9% regional percentage and 19.7% at the Anela forest domain; contrarily, the euri-Mediterranean component has 16.1% regional average but increases to 24% at our study area, the same percentage (24.3%) reached by the sum of the Boreal-Temperate and the Eurasian floristic components. Whereas lower altitude floras have a dominant steno-Mediterranean component and the floras at the summit of Mediterranean mountains show the prevalence of southern-European and Mediterranean orophytes and narrow endemics (Cañadas et al. 2014; Arrigoni and Camarda 2015), our flora is a good example of transition areas, having the 80% of taxa quite equally distributed amongst steno-Mediterranean, euri-Mediterranean, Boreal-Temperate and Eurasian and the endemic contingents. High species density, H/T ratio ~ 1, balance amongst different chorologic groups and endemic percentage ~ 10% can be considered characteristic features of mountain areas at the transition between the Mediterranean and the temperate bioclimates.

The composition of the flora of the Forest Domain of Anela is also peculiar because it is one of the few examples, not only in Sardinia but in the whole Mediterranean area, with no native Gymnosperms. Junipers (Juniperus phoenicea subsp. turbinata (Guss.) Nym. and J. oxycedrus subsp. macrocarpa (Sibth. & Sm.) Neilr.) in NW Sardinia are mainly confined in coastal areas (Farris et al. 2017), but Yew (Taxus baccata L.) and Prikly Juniper (Juniperus oxycedrus L. subsp. oxycedrus) are usually present in high hills and mountains. However junipers are not present in NW Sardinia inland areas (Farris et al. 2017), but the Yew is occurring in all the massifs and mountain ranges, including the two forest domains bordering Anela, the Fiorentini Forest Domain to the east (municipality of Bultei) and the Mt. Pisanu Forest Domain to the west (municipality of Bono, see Farris and Filigheddu 2008). The total absence of Gymnosperms in the native flora of the Anela forest domain is therefore surprising, most probably anomalous and it seems likely to be linked to the management history of the area rather than a natural pattern (Sechi and Falchi 2013).

Despite the fact that in 2004 (last forest census) 90.4% of the domain area was covered by forest or shrub communities (Sechi and Falchi 2013), it is striking that the 45% of the detected taxa were linked mainly to herbaceous habitats (annual and perennial grasslands, dry and wet pastures and meadows), already described for their peculiar and original floristic composition (Farris et al. 2013). Traditional grazing, particularly ovine pastoralism characterised by low flock density and transhumance, has been proven to be beneficial for the plant biodiversity of Mediterranean silvo-pastoral systems, whereas abandonment is detrimental even at short temporal scales (Farris et al. 2010a). The forest domain of Anela is a typical case where ovine stocks had a dramatic decrease in a short period: between 1990 and 2007, a decrease from 0.77 sheep ha^-1 to 0.13 sheep ha^-1 has been recorded (-83%, Farris et al. 2010a), whereas wood and shrub communities linked to potential natural vegetation (sensuFarris et al. 2010b) are recovering very fast, following a trend common to all Italy (Falcucci et al. 2007) and particularly to Sardinia (Puddu et al. 2012).

Table 1.

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Synthetic data on mountain floras from Sardinia and the regional flora, based on different sources (see notes below).

Site	Altitudinal interval	Area (km²)	No. taxa	Taxa / km²	H/T	No. endemics	% endemics	Source
Anela forest domain	600–1158	12.8	391	30.6	1.03	45	11.5	This work
Gennargentu	1500–1834	16.8	214	12.7	2.5	66	30.8	Arrigoni and Camarda 2015
Gennargentu	1000–1834	240	675	2.8	1.25	105	15.6	Arrigoni and Camarda 2015
Gennargentu	1000–1834	500	897^†	1.8	1.03^‡	n.d.	28^§	Bacchetta et al. 2013
Supramontes	0–1463	335	n.d.	n.d.	n.d.	138	30 ^§	Fenu et al. 2010
Mt. Albo	900–1127	68	659	9.7	0.61	48	7.3	Camarda 1984a
Mt. Gonare	538–1083	10	520	52	0.85	23	4.4	Camarda 1984b
Mt. Limbara	160–1359	166.24	923	5.5	0.75	80	8.7	Calvia and Ruggero unpublished
Mt. Limbara	800–1359	49.46	687	13.9	0.84	72	10.5	Calvia and Ruggero unpublished
Mt. Limbara	500–1359	n.r.	506	n.d.	1.18	55	10.9	Veri and Bruno 1974
Sardinia	0–1834	24090	2028	0.084	0.70	n.d.	7.1	Pignatti 1995
Sardinia	0–1834	24090	2400	0.099	n.d.	n.d.	n.d.	Arrigoni (2006–15)
Sardinia	0–1834	24090	2408^\|	0.1	0.74^¶	290^#	12	Various (see notes)
Sardinia	0–1834	24090	2149	0.09	n.r.	290	13.5	Médail 2017, table 2
Sardinia	0–1834	24090	2301	0.095	n.r.	331	14.4	Bartolucci et al. 2018

^†Bacchetta et al. (2013) list 948 entities, including 10 varieties, 3 hybrids and 38 aliens: here we therefore consider 897 native taxa;

^‡calculated by Arrigoni and Camarda 2015;

^§Cañadas et al. 2014;

^|Conti et al. 2005;

^¶Arrigoni and Camarda 2015;

^#Fenu et al. 2014; n.r. not reported; n.d. not determined.

Conservation issues of this Flora

Even if rarity is not always linked to threat (de Lange and Norton 1998, Bacchetta et al. 2012), it is an important feature to consider when setting conservation priorities within long lists of taxa (Bacchetta et al. 2012, Le Berre et al. 2018), as in the case of the flora of the Anela forest domain. Additionally, 14 out of 32 uncommon and range-restricted taxa found in this flora are linked to wet habitats: some belong to the Mediterranean and endemic contingents (Cerastium ligusticum subsp. palustre, Exaculum pusillum, Isoetes hystrix, Mentha suaveolens subsp. insularis, Mentha requienii subsp. requienii, Morisia monanthos, Oenanthe lisae), others to the Eurasian and Boreal-Temperate contingents (Struthiopteris spicant, Carex remota, Iris pseudacorus, Solanum dulcamara, Spiranthes spiralis). Those habitats are supposed to be highly vulnerable (Filipe et al. 2013), as changes in land use and modification of water balance (because of climate change or human use) are amongst the most important threats to wetlands. Moreover, little is known about the resilience of associated plant communities, a threat increased by the high spatial isolation of such places within a Mediterranean context. At the study site, we detected several species having a contraction of range or local extinctions caused by the capture of surface or underground water for human use, as for example Struthiopteris spicant, Cerastium ligusticum subsp. palustre, Mentha requienii subsp. requienii and the localized fern Osmunda regalis for which we documented a local decrease > 50% in the last 20 years. Other species had a decrease directly caused by drainage of temporary ponds (Exaculum pusillum, Isoetes hystrix, Morisia monanthos). Water management in a climatic changing scenario is and will increasingly be a key issue for the conservation of biodiversity in the Mediterranean basin (Casazza et al. 2014), a climatic change hotspot at the global scale (Giorgi 2006, Giorgi and Lionello 2008), where wet habitats and the species linked are amongst the most threatened (Ghosn et al. 2010, Pérez-Luque et al. 2015).

The 5 taxa, locally extinct, have no relationship with a particular habitat or human use from which they are (were) dependent for their survival in the area, with the exception of Chenopodium album whose disappearance could be explained with the above-mentioned abandonment of pastoral activities, as it is a nitrophilous species. Their disappearance in the last decades, inferred from herbarium records, can be therefore a normal turnover in the composition of the local indigenous flora or an artifact derived from our sampling method (in the sense that these taxa are maybe still present in the area but we were not able to find them during our monthly sampling excursions).

Amongst the flora we inventoried, it is worth mentioning that several populations represent peripheral populations regarding the overall distribution of the taxa. First, a group of uncommon or range restricted species in the domain, are common plants in the Mediterranean bioclimate areas of Sardinia and sometimes in the whole basin. They are here confined to warm niches in the mountain area under study (Anemone hortensis, Arbutus unedo, Arisarum vulgare, Arum pictum, Celtis australis, Ficus carica, Ptilostemon casabonae), places relatively scattered through this mountain landscape. Oppositely, several Boreal-Temperate and Eurasian taxa confined in this sub-Mediterranean bioclimate island represent peripheral populations isolated sometimes by over 1000 km of their northern range. Those constitute rear edge populations (Hampe and Petit 2005) which may contain unique genetic variation, inherited from ancient species distribution and particular ecological conditions. These two contrasted situations have been highlighted several times within the Mediterranean flora (Lavergne et al. 2005, 2006) and are characteristic of those climatic transition areas. These plants all share the characteristic of occurring as fragmented, disjunct and often highly isolated populations, which restrain gene flow with central population (Pironon et al. 2017) and enhance amongst-population differentiation (Papuga et al. 2018). Thus, the relative isolation associated with potentially marginal ecological conditions highlight their evolutionary potential (Thompson 1999, Anacker and Strauss 2014), as it has recently been shown in Sardinia and Corsica for some marginal and peripheral populations of Cyclamen repandum (Thompson et al. 2018). Additionally, these groups of taxa are often found in different macro-habitats which have very different links with human activities, therefore leading to different threats and management issues (Lavergne et al. 2006). Thus, conservation policies need to integrate such complex entities within their framework (Lesica and Allendorf 1995, Brunnell et al. 2004, Leppig and White 2006). Finally, those transition areas also contain numerous endemics, which render those places original and of high value for conservation.

Even if biodiversity hot-spots definition at multiple spatial scales is commonly based on the presence, density and distribution of endemic taxa (Myers et al. 2000, Cañadas et al. 2014), the data here presented support that other parameters should also be taken into account to more precisely define priority areas for conservation, as taxonomic complexity (Ennos et al. 2005) of floras and evolutionary potential of populations (Thompson et al. 2010), detected within continuous schemes of biodiversity monitoring (Marignani et al. 2014). This is particularly urgent in southern European mountains, whose biodiversity is threatened by both climate and land use changes (Bravo et al. 2008, Benito et al. 2011, Pauli et al. 2012, Vogiatzakis et al. 2016).

Acknowledgements

E.F. acknowledges the Amministrazione Provinciale di Sassari – Settore Ambiente e Agricoltura Nord Ovest, for funding the project “Monitoraggio di flora, vegetazione e habitat delle oasi di protezione faunistica del Goceano per la gestione delle popolazioni di animali selvatici” for the years 2014–17.

P.d.L. and E.F. thank the project Visiting Professor (Sardinian regional laws 3/2008 and 2/2009) for funding P.d.L. journey to Sardinia in 2013 (Protocol No. 13574 of the 07.05.2012 of the University of Sassari).

Giacomo Calvia and Alessandro Ruggero kindly offered their unpublished quantitative data on the flora of Mt. Limbara for Table 1.

Authors thank all the workers and staff of the forest station of Foresta Demaniale Anela belonging to the Sardinian regional agency Forestas for their help and genuine hospitality given throughout the entire research period.

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