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Phytosociological features of Adonis distorta and Trifolium noricum subsp. praetutianum, two endemics of the Apennines (peninsular Italy)

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We present plant communities in which two endemic taxa of the Apennine mountain chain, Adonis distorta Ten. and Trifolium noricum Wulfen subsp. praetutianum (Savi) Arcang are distributed. Although these taxa occur sporadically in some other Apennine massifs, it is only in the Majella mountain range where these species are physiognomi-cally dominant. Trifolium noricum subsp praetutianum behaves as a differential element in Helianthemo-Festucetum italicae, where it characterizes a new edapho-mesophilous subassociation named Helianthemo alpestris-Festucetum italicae trifolietosum praetutia-ni. Adonis distorta is linked to specific geo-morphotypes characterised by an alternation of strips of detritus and fine soil particles as well as to distinguishable communities de-scribed in this paper as a new association named Ranunculo seguierii-Adonidetum distortae.
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Acta Bot. Croat. 67 (2), 175–200, 2008 CODEN: ABCRA 25
ISSN 0365–0588
Phytosociological features of Adonis distorta and
Trifolium noricum subsp. praetutianum, two endemics
of the Apennines (peninsular Italy)
ROMEO DIPIETRO1*, GIOVANNI PELINO2,ANGELA STANISCI2,CARLO BLASI3
1Department ITACA, University of Rome La Sapienza, Via Flaminia 70,
I-00196 Rome, Italy
2Department of Science and Technologies for the Environment and Territory,
University of Molise, Contrada Fonte Lappone, I-86170 Pesche (Isernia), Italy
3Department of Plant Biology, University of Rome La Sapienza, Piazzale Aldo Moro 5,
I-00185, Roma, Italy
We present plant communities in which two endemic taxa of the Apennine mountain
chain, Adonis distorta Ten . an d Trifolium noricum Wulfen subsp. praetutianum (Savi)
Arcang are distributed. Although these taxa occur sporadically in some other Apennine
massifs, it is only in the Majella mountain range where these species are physiognomi-
cally dominant. Trifolium noricum subsp praetutianum behaves as a differential element
in Helianthemo-Festucetum italicae, where it characterizes a new edapho-mesophilous
subassociation named Helianthemo alpestris-Festucetum italicae trifolietosum praetutia-
ni.Adonis distorta is linked to specific geo-morphotypes characterised by an alternation
of strips of detritus and fine soil particles as well as to distinguishable communities de-
scribed in this paper as a new association named Ranunculo seguierii-Adonidetum
distortae.
Keywords: Adonis,Trifolium, alpine, vegetation, synecology, syntaxonomy, Italy
Introduction
The basic features of the vegetation found on the Majella massif above the timberline
have been partially described in previous studies (MIGLIACCIO 1966, 1970; FEOLI-CHIA-
PELLA and FEOLI 1977; FEOLI-CHIAPELLA 1983; BIONDI et al. 1988; PETRICCIONE 1988;
STANISCI 1994, PETRICCIONE and PERSIA 1995; STANISCI 1997; BLASI et al. 2003, 2005).
Nonetheless, the extension of the alpine and subalpine belts, which cover relatively wide
areas of the Majella massif, and the difficulty of reaching the high altitude zones have pre-
cluded the carrying out of thorough studies of these environments. This is especially the
case as regards the central and eastern sectors of the massif, which are characterized by ex-
tensive plateaus, such as Piano Amaro and Sella di Grotta Canosa.
ACTA BOT. CROAT. 67 (2), 2008 175
* Corresponding author, e-mail: romeodipietro@hotmail.com
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This paper reports two new plant communities that have been found in these areas,
growing in the alpine bioclimatic belt, at altitudes ranging between 2500 and 2600 m a.s.l.
These communities are dominated by Adonis distorta Ten. and Trifolium noricum Wulfen
subsp. praetutianum (Savi) Arcang respectively, two taxa of high biogeographical impor-
tance for the Italian flora but extremely sporadic in the rest of the Apennines. Although
well known to botanists because of their biogeographical significance, the role of Adonis
distorta and Trifolium noricum subsp. praetutianum has never previously been studied
from a phytosociological and syntaxonomical point of view. There was a tendency to clas-
sify them as »companions« (less often as differentials) within communities dominated by
other species, rather than as dominant taxa of specific plant communities at the coenolo-
gical, phytosociological and nomenclatural level.
Study Area
The limestone massif of Majella (2974 m.) is one of the most imposing mountain mas-
sifs in the Apennine chain. It consists of a mountain ridge about 30 km in length, running in
a north-south direction between 42°12’00’’north and the 42nd parallel south, 32.5 km W of
the Adriatic Sea coast (Fig. 2). A morphological peculiarity of the Majella is its shape,
which is that of a dome flattened at the top; its very steep sides are cut by deep gorges of
glacial origin (DEMANGEOT 1965; JAURAND 1994; GIRAUDI 1998). Our study area was lo-
cated on the flattish areas occurring between 2450 and 2650 m a.s.l. (Piano Amaro, Sella
Grotta Canosa, Cima dell’Altare and M. Macellaro). The climate of this area is extremely
cold, with a mean minimum temperature in the coldest month lower than –5° and with
176 ACTA BOT. CROAT. 67 (2), 2008
DIPIETRO R., PELINO G., STANISCI A., BLASI C.
Fig. 1. Adonis distorta Ten (A), and typical stand of Helianthemo-Festucetum trifolietosum with
Trifolium noricum subsp. Praetutianum (B).
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snow cover for 7–8 months a year. From a bioclimatic point of view the area belongs to the
cold axeric subregion of the temperate region (RIVAS-MARTINEZ 1995) and to the cryo-oro-
temperate thermotype (BLASI et al. 2003).
The Majella massif is one of the most interesting floristic and vegetational areas in the
whole Italian peninsula. As in the rest of the Apennines, beech woods dominate the entire
montane belt up to 1800 m, while in the subalpine belt well-developed Pinus mugo wood-
lands extend up to 2300–2400 m. This is the only area where Pinus mugo stands are to be
found in the entire Apennine range, where Juniperus alpina and Arctostaphylos uva-ursi
tend to dominate on limestone and Vaccinium myrtillus on sandstones.
The karst plateaus of the alpine belt in the Majella massif are covered by an almost con-
tinuous layer of calcareous debris due to periglacial phenomena that have caused the break-
ing up of the rocky substrate into flattish clasts, small (3–6 cm) and sharp-edged (DRAMIS
and KOTARBA 1992). On these plateaus, in the absence of steep slopes, sliding movements
in the upper part of the substrate are practically non-existent. However, the prolonged win-
ter cold causes the formation of ice needles (pipkrakes) perpendicular to the surface, which
lift small stones, topsoil and plant remains from the ground. The soil is moreover subject to
the selective action of the wind, which blows away finer particles. The heavier material
which is left is then subjected to the weight of the snow cover and the action of flowing wa-
ter (both rainwater and melting snow water), which lines stones up with their longer sides
parallel and smaller end surface pointing upwards (GENTILESCHI 1967; JAURAND 1994). Un-
der these edaphic conditions, only a specialized flora, such as that characterizing the
xerophilous component of Thlaspietea rotundifolii (Saxifrago-Papaveretum julici, Cre-
pido-Leontodontetum montani) can survive.
In sheltered spots on the windy ridges, where small accumulations of finer material can
build up, it is possible to find Elyno-Seslerietea communities: Leontopodio-Elynetum and
Leontopodio-Seslerietum, at higher and lower altitudes, respectively. Instead, on the slight-
ACTA BOT. CROAT. 67 (2), 2008 177
TWO ENDEMICS OF THE APENNINES
Fig. 2. Schematic map of the geographical position of the study area.
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ly sloping stretch connecting the lower part of the slope to the bottom of the karst basins
Helianthemo-Festucetum italicae can be found. In similar environmental conditions to
these, communities belonging to the meso-hygrophilous fringe of Thlaspietea rotundifolii
(e.g. Carici-Salicetum retusae), floristically richer and less fragmented in structure, also
occur. Nardetea strictae communities (Gnaphalio-Plantaginetum atratae,Taraxaco-Trifo-
lietum thalii) are rare and only found in dolinas.
Investigated taxa
Adonis distorta is a steno-endemic species defined as »italic tertiary relic« (MONTE-
LLUCCI 1971). Its distribution area, restricted to the limestone massifs of the Central
Apennines such as Majella, Gran Sasso, Sirente, Vettore (Sibillini mountains), Velino and
Monti della Duchessa, has its southern limit at Mount Morrone in the southern Abruzzo ad-
ministrative region (TENORE and GUSSONE 1842, DEL GROSSO and POGLIANI 1971, TAM-
MARO 1971, FRATTAROLI and FRIZZI 1988, TAMMARO 2000) (Fig. 3A).
The relatively more widespread Trifolium noricum subsp. praetutianum is found in an
area ranging from the Sibillini Mountains (Umbria administrative region) to the Matese
massif (Campania administrative region) (Fig. 3B). Compared to Adonis distorta, which is
a strictly basiphilous species, Trifolium noricum subsp. praetutianum is edaphically less
demanding, and can also grow on acid and subacid soils (MAURI et al. 1830, FALQUI 1898,
LUCCHESE and DESIMONE 2000, TAMMARO 2000).
178 ACTA BOT. CROAT. 67 (2), 2008
DIPIETRO R., PELINO G., STANISCI A., BLASI C.
Tab. 1. Life form spectra calculated: on the percentage of species belonging to the different life
forms in the phytosociological tables 3 and 4 (n); weighted, based on the frequency of the
different life forms (f); weighted, based on the specific cover index (i.r.s.) of the different life
forms (c).
Life forms
Ranunculo-Adonitetum Helianthemo-Festucetum
trifolietosum
n% f% c% n% f% c%
Tot. Hemicryptophytes 65,0 63,9 71,4 63,0 66,5 75,0
Tot. Chamaephytes 27,5 30,1 22,0 30,4 31,2 23,3
Tot. Geophytes 5,0 5,3 6,5 4,3 1,2 1,6
Tot. Therophytes 2,5 0,8 0,1 2,2 1,2 0,1
H scapose 35,0 36,1 45,5 30,4 31,2 18,8
H rosulate 22,5 19,5 19,4 15,2 15,3 9,8
H caespitose 7,5 8,3 6,5 15,2 19,4 46,3
H biennis 0,0 0,0 0,0 2,2 0,6 0,1
Ch suffruticose 22,5 25,6 20,4 17,4 18,2 15,4
Ch succulent 0,0 0,0 0,0 4,3 2,4 0,6
Ch reptant 2,5 3,8 1,5 4,3 5,3 4,1
Ch pulvinate 2,5 0,8 0,1 4,3 5,3 3,2
T scapose 2,5 0,8 0,1 2,2 1,2 0,1
G rhizomatose 5,0 5,3 6,5 4,3 1,2 1,6
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Materials and methods
The field work was carried out during 2002–2003, with the establishment of 15 vegeta-
tion relevés using the Braun-Blanquet phytosociological approach (BRAUN-BLANQUET 1964).
Synoptic tables were also compiled in order to verify the coenological and syntaxonomical
autonomy of the plant communities identified in this paper.
Nomenclature of the species follows CONTI et al. (2005), while phytosociological no-
menclature follows the rules of ICPN (WEBER et al. 2000). Chorological elements and life
forms were drawn from PIGNATTI (1982). In order to obtain more complete information
ACTA BOT. CROAT. 67 (2), 2008 179
TWO ENDEMICS OF THE APENNINES
Fig. 3. Updated distribution map of Adonis distorta (A) and Trifolium noricum subsp. praetutianum
(B). The distribution maps were drawn up using the topographic map of Italy (UTM system)
of the Italian Geographic and Military Institute, superimposed by a grid of square elements
at the scale of 1:50.000 with dimensions of 20’ in longitude and 12’in latitude.
Tab. 2. Chorotypes spectrum calculated: on the percentage of species belonging to the different
chorotypes in the phytosociological tables 3 and 4 (n); weighted, based on the frequency of
the different chorotypes (f); weighted, based on the specific cover index (i.r.s.) of the differ-
ent chorotypes (c).
chorotypes
Ranunculo-Adonitetum Helianthemo-Festuc.
trifolietosum
n% f% c% n% f% c%
Endem. Central Apennines 17,5 21,1 38,1 8,7 7,6 2,8
Endem. Centr.-South. Apennines 7,5 7,5 3,5 10,9 15,9 36,5
Endemic Italian 10 6,0 1,1 10,9 11,2 5,2
SE-European Orophytes s.l. 17,5 23,3 25,0 17,4 17,6 16,6
European Orophytes 22,5 20,3 17,2 30,4 22,9 13,3
Mediterranean-Montane 7,5 7,5 6,5 2,2 2,9 0,9
Euroasiatic 7,5 6,8 4,8 10,9 12,9 15,2
Circumboreal and Artic-Alpine 7,5 6,8 3,7 8,7 8,8 9,5
Cosmopolite 2,5 0,8 0,1 0,0 0,0 0,0
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about the structure and chorology of the plant communities we analysed the percentages of
the different life forms and chorotypes in the phytosociological tables (normal spectrum
»n«), the frequency of occurrence in the phytosociological tables (frequency spectrum,
»f«), and the specific cover index of each chorotype in the phytosociological tables (I.R.S.
spectrum). The specific cover index (I.R.S.) for each species makes reference to BRAUN-
-BLANQUET (1964); it is the ratio (multiplied by 100) between the sum of the medium value
of the Braun-Blanquet dominance-abundance index (e.g. 5 = 87,5, 4 = 62,5...) and the num-
ber of relevés in the phytosociological table. In the synoptic tables the frequency percent-
ages of the species in the original phytosociological tables (see caption of Tab. 5 and 6) are
reported. The synoptic tables report the syntaxonomical scheme according to BLASI et al.
(2003, 2005) (Tabs. 8, 9).
Results
Ranunculo seguierii-Adonidetum distortae ass. nov. hoc loco
(Holotypus Tab. 3, rel. 3)
The plant community dominated by Adonis distorta and Ranunculus segueri subsp.
seguieri grows in the form of scattered stands of limited extent (about 50 m2), usually lo-
cated on very gentle slopes (from 0° to 7°), where the substrate consists of a continuous
layer of calcareous detritus made up of small-sized (2–5 cm) pebbles (Fig. 4A). The slight
slope of the ground causes rain and snow melt-water to flow slowly towards the nearest
180 ACTA BOT. CROAT. 67 (2), 2008
DIPIETRO R., PELINO G., STANISCI A., BLASI C.
Fig. 4. Physiognomical expression of Ranunculo-Adonidetum distortae in the alpine belt of Majella
(A), and Helianthemo-Festucetum italicae typicum (Ba) with Helianthemo-Festucetum
trifolietosum (Bb).
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karstic swallow-hole, in the process creating alternating strips of detritus (white colored)
and fine soil particles (dark colored). These are the so-called »striped soils« which are
fairly common in the flattish areas of the eastern part of the Majella massif (SACCO 1908,
SEGRE 1947).
The main physiognomic role in the community is played by Adonis distorta (specific
cover index = 3300), but Leontodon montanus subsp. montanus,Ranunculus seguieri
subsp. seguieri and Cerastium thomasii are found frequently and in abundance as well (Fig.
4A).
In the present paper we propose a new association which belongs to the endemic
Apennine alliance Linario-Festucion dimorphae (Thlaspietea rotundifolii), to be named
Ranunculo seguierii-Adonidetum distortae. Character species of this community are Ado-
nis distorta,Ranunculus seguieri and Crepis bithynica. The choice of this specific charac-
ter component follows a biogeographical criterion. In fact, these species are not exclusive
to the Majella massif (the species occurs also in Gran Sasso massif). However Crepis
bithynica exhibits the highest frequency and abundance in the Majella massif, in the associ-
ation Ranunculo-Adonidetum. From the spatial-pattern point of view, Ranunculo-Adoni-
detum comes into contact with pioneer vegetation types such as Crepidi-Leontodontetum
(unstable screes), and Galio-Silenetum acaulis (dwarf cushions of alpine tundra). Where
the clayey component of the soil is higher, Ranunculo-Adonidetum comes into spatial con-
tact with Helianthemo-Festucetum italicae (Elyno-Seslerietea).
Helianthemo alpestris-Festucetum italicae trifolietosum praetutiani
subass. nova hoc loco (Holotypus Tab. 4, rel. 12)
The communities of Trifolium noricum subsp. praetutianum form thick, dark-green
patches of limited extent (about 50 square metres) within the wider areas occupied by
Helianthemo alpestris-Festucetum italicae (Fig. 4B), which is an association described for
the karst-tectonic basins of lower altitude of the Majella massif but which can also be found
at higher altitudes. The occurrence and the dominance of Trifolium noricum subsp.
praetutianum in the range of the Helianthemo-Festucetum alpestris communities appear to
be linked to slight variations in the edaphic and micro-geomorphological features of the
substrates. It is found where the slope of the ground is less steep (about 5–10° C) and the
upper soil layer is deeper and richer in fine material, with a consequent reduction in its ba-
sicity. In an environment such as this, Trifolium noricum subsp. praetutianum is very com-
petitive, forming its typical continuous carpets among surrounding vegetation types having
a lower degree of cover, and thus reducing the possibilities of growth for other plants (Fig.
4B). Thermal factors would appear to play a role, too, since Trifolium noricum subsp.
praetutianum communties are much more widespread on the warmer slopes of the massif,
being found more frequently in those areas subject to the mitigating influence of the Adri-
atic Sea breezes, on the eastern slopes, and the mostly south-facing plateaus of Majella
higher zones (STANISCI et al. 2005). On the other hand, on the north facing slopes or where
the microclimatic conditions are too cold, Trifolium noricum subsp. praetutianum stands
are never found. In fact, the ecological optimum of Trifolium noricum subsp. praetutianum
occurs at altitudes lower than those recorded in the present study. For this reason the
Trifolium noricum subsp. praetutianum grassland type we found on the Majella alpine belt
is considered to be a sub-association (edapho-mesophilous) of the association Helian-
ACTA BOT. CROAT. 67 (2), 2008 181
TWO ENDEMICS OF THE APENNINES
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182 ACTA BOT. CROAT. 67 (2), 2008
DIPIETRO R., PELINO G., STANISCI A., BLASI C.
Tab. 3. New association of Ranunculo seguierii-Adonidetum distortae. Place and date of the relevés: Rel. 1: Taranta Peligna valley, 5/8/2003, 42° 3’
49,8’’N 14° 6’ 40’’ E; Rel. 2-3-6-7-8 : Piano Amaro, 9/8/2003, 42°4’23,1’’N 14°6’40,4’’E (rel.2), 42°4’22,8’’N 14°6’48,7’’E (rel.3), 42°4’30,5’N
14°6’29,3’’E (rel.6), 42°4’52’’N 14°7’4,4’’E (rel.7), 42°4’35,3’’N 14°6’45,1’’E (rel.8); Rel. 4 : Piano Amaro-Sella Grotta Canosa, 6/8/2003, 42°4
29,7N14°6’34,3E;Rel.5:Sella Grotta Canosa, 9/8/2003, 42°4’30,5’’N 14°6’29,3’’E.
relevée no. 12 345678
altitude m a. s. l. (x10) 243 255 256 255 255 255 252 254
aspect WWNWSW0 0W
slope° 72 23,54001
rochyness (% of stable rocks
occurring in the relevé area)
00 000000
detritus% (% of mobile detritus occurring in
the relevé area)
100 95 100 100 100 100 100 100
area m2 60 50 50 70 100 16 20 20
cover % 55 55 45 60 60 45 45 45
number of species per relevée 19 27 23 14 16 14 9 11
Typ.
Char.species of Ranunculo seguierii-Adonitetum distortae freq.% i.r.s.
Adonis distorta 333332321003300
Ranunculus seguieri subsp. seguieri 2 3 2 . . 2 2 2 75 1600
Crepis bithynica .+++....3838
Char. species of Thlaspienion stylosi
Cerastium thomasii 1223212+1001413
Linaria alpina . . . + . +1 15050
Thlaspi stylosum ..111...3838
Viola magellensis ..1....+2525
Alyssum cuneifolium subsp. cuneifolium .+.....+2525
Char. species of Linario-Festucion (Al), Thlaspietalia rotundifolii (Or), Thlaspietea rotundifolii (Cl)
Cl Leontodon montanus subsp. montanus 132331111001700
Al Ranunculus brevifolius +11231. .75750
Or Arenaria grandiflora subsp. grandiflora 2 1 1 . + 2 . 2 75 713
Al Achillea barrelieri subsp. barrelieri +1112+. .75288
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ACTA BOT. CROAT. 67 (2), 2008 183
TWO ENDEMICS OF THE APENNINES
relevée no. 12 345678
Al Galium magellense . 1 1 . . . 1 1 50 50
Or Iberis saxatilis subsp. saxatilis +....1..2525
Al Valeriana saliunca .++.....2525
Al Androsace vitaliana subsp. praetutiana ..+...+.2525
Al Myosotis ambigens .+......1313
Seslerion apenninae (Al), Seslerietalia tenuifoliae (Or), Elyno-Seslerietea (Cl)
Or Anthyllis vulneraria subsp. pulchella 1++11+.+8888
Cl Minuartia verna subsp. verna ++1+1+..7575
Or Androsace villosa subsp. villosa ++1. .1.16363
Cl Potentilla crantzii subsp. crantzii . + + . . . + . 38 38
Cl Draba aizoides subsp. aizoides +11.....3838
Or Acinos alpinus subsp. alpinus . + + . + . . . 38 38
Al Pedicularis elegans .++..+..3838
Cl Campanula scheuchzeri +2......25238
Or Edrajanthus graminifolius subsp. graminifolius +.+.....2525
Cl Silene acaulis s.l. + .......1313
Al Avenula praetutiana +.......1313
Al Gnaphalium hoppeanum subsp. magellense .+......1313
Or Armeria majellensis s.l. . + ......1313
Cl Erigeron epiroticus .+......1313
Cl Sedum atratum subsp. atratum ....+...1313
Other species
Poa molinerii 1.112. +.63275
Poa alpina subsp. alpina 11112. . .63275
Taraxacum apenninum (group) . + . + 1 . . . 38 38
Leontodon hispidus 3...+...25488
Plantago atrata subsp. atrata .+..+...2525
Viola eugeniae subsp. eugeniae .+......1313
Botrychium lunaria ...+....1313
Leontodon crispus subsp. crispus .....+..1313
Tab. 3. – continued
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184 ACTA BOT. CROAT. 67 (2), 2008
DIPIETRO R., PELINO G., STANISCI A., BLASI C.
Tab. 4. New subassociation of Helianthemo alpestris-Festucetum italicae trifolietosum praetutiani. Place and date of the relevés: Rel. 9-10-11-12 : Piano
Amaro, 9/8/2003, 42°4’48’’N 14°5’56,6’’E (rel.9), 42°4’29,7’’N 14°6’55,2’’E (rel.10), 42°4’21,7’’N 14°7’1’’E (rel.11), 42°4’20,4’’N 14°6’52,2’’E
(rel.12); Rel.13 slope S-E of the Macellaro Mt., 5/8/2003, 42°3’9’’N 14°6’29,4’’E; Rel. 14: Piano Amaro, 29/7/2003, 42°4’37,5’’N 14°7’4,7’’E;
Rel. 15: Piano Amaro-Cima dell’Altare, 29/7/2003, 42°4’42,3’’N 14°7’4,4’’E.
relevée no. 9101112131415
altitude m a.s.l. (x10) 260 255 253 256 253 255 254
aspect ENE E ESE E E N NNE
slope° 9,5 3 5 6 17 7,5 8
rochyness (% of stable rocks occurring in the relevé area) 5000015
detritus% (% of mobile detritus occurring in the relevé area) 45 30 30 40 40 50 20
area sq. mt. 25 100 70 36 20 20 20
cover % 85 90 75 80 85 85 80
number of species per relevée 24 19 24 21 31 26 22
Char. species of Helianthemo alpestris-Festucetum italicae Typ. freq.% i.r.s
Helianthemum oelandicum subsp. alpestre 3.33111861843
Festuca violacea subsp. italica 22222221001800
Poa molinerii +2221111001000
Leontopodium nivale .11111.71357
Sempervivum arachnoideum . . . + . + 1 43 100
trifolietosum praetutianii subass. nova
Trifolium noricum subsp. praetutianum 34444331005229
Char. species of Leontopodio-Elynenion
Potentilla crantzii subsp. crantzii 21221111001057
Silene acaulis s.l. + 1 1 2 1 . 1 86 557
Oxytropis campestris 111111+100443
Sedum atratum subsp. atratum ..++...2929
Char. species of Seslerion apenninae (Al), Seslerietalia tenuifoliae (Or), Elyno-Seslerietea (Cl)
Or Carex kitaibeliana subsp. kitaibeliana 22112211001243
Or Armeria majellensis s.l. 22112111001057
Cl Thymus praecox subsp. polytrichus 211.21186800
Cl Minuartia verna subsp. verna 1+1111 .86371
Or Anthyllis vulneraria subsp. pulchella 2 . . + 2 1 . 57 600
Al Trinia dalechampii 111111+100443
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ACTA BOT. CROAT. 67 (2), 2008 185
TWO ENDEMICS OF THE APENNINES
relevée no. 9101112131415
Or Edrajanthus graminifolius subsp. graminifolius 1111+11100443
Cl Draba aizoides subsp. aizoides 1111+11100443
Al Pedicularis elegans + . 1 1 . . 1 57 229
Al Achillea barrelieri subsp. barrelieri +. . +++.5757
Al Avenula praetutiana +...11.43157
Or Androsace villosa subsp. villosa ...+++.4343
Cl Erigeron epiroticus +...+..2929
Cl Gentiana verna subsp. verna .+..+..2929
Cl Campanula scheuchzeri 1......1471
Cl Bistorta vivipara 2......14257
Cl Phyteuma orbiculare ....2..14257
Cl Astragalus depressus subsp. depressus .....+.1414
Cl Acinos alpinus subsp. alpinus ......+1414
Char. species of Ranuncolo-Nardion (Al), Nardetalia (Or), Nardetea (Cl)
Al Viola eugeniae subsp. eugeniae 1+. . 1+.57171
Cl Hieracium pilosella 1 . +. +. .43100
Cl Plantago atrata subsp. atrata ....1.129143
Cl Phleum alpinum subsp. rhaeticum ..+....1414
Char.species of Linario-Festucion (Al), Thlaspietalia rotundifolii (Or), Thlaspietea rotundifolii (Cl)
Or Arenaria grandiflora subsp. grandiflora . 1 + + + . 1 71 186
Al Myosotis ambigens . + 1 . + 1 + 71 186
Al Androsace vitaliana subsp. praetutiana ..1..1+43157
Cl Saxifraga oppositifolia subsp. oppositifolia . + . . + 1 . 43 100
Al Cerastium thomasii .....+12986
Cl Salix retusa 1......1471
Al Artemisia umbelliformis subsp. eriantha ..+....1414
Al Ranunculus brevifolius ....1..1471
Other species
Thesium parnassi ..+....1414
Poa alpina subsp. alpina ....1..1471
Cerastium arvense subsp. strictum ....+..1414
Sedum acre .....+.1414
Saxifraga adscendens subsp. adscendens ......+1414
Tab. 4. – continued
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themo-Festucetum italicae, and it is codified here with the epithet »trifolietosum praetu-
tiani« (subass. nova). This sub-association is included in the Leontopodio-Elynenion
sub-alliance and the Seslerion apenninae alliance.
Structural and chorological analysis of the communities
The life form spectrum shows a marked dominance of Hemicryptophytes and, to a
lesser degree, of Chamaephytes. The presence of Geophytes, on the other hand, is negligi-
ble, and Therophytes are only represented by Sedum atratum subsp. atratum. Phanero-
phytes are absent, and, in any case, at these altitudes could only be represented by isolated
stations of Pinus mugo or Juniperus communis subsp. nana (Tab. 1). Among the hemi-
cryptophytes the »scapose« growth form is dominant but also »rosulate« forms are also
well represented. The chamaephyte life form is almost exclusively composed of suffru-
ticose species (Ch suffr.). The Hemicryptophytes/Chamaephytes ratio is similar in the two
communities identified, ranging between 2.1 and 3.2.
From a chorological point of view, the communities investigated present a strong en-
demic component, especially those restricted to the central Apennines and to the cen-
tral-southern Apennines (Tab. 2). The high percentage of endemics is strictly connected to
the highly fragmentary character of the alpine bioclimatic belt in the Apennines, which has
led to species isolation and consequent interruption of genic flow.
The central Apennines endemic component is higher in Ranunculo-Adonidetum than in
Helianthemo-Festucetum trifolietosum. This is due to the frequent and abundant presence,
apart from Adonis distorta, of species such as Cerastium thomasii, Galium magellense and
Thlaspi stylosum, whose development is favoured by the higher percentages of detritus in
the substrate.
Helianthemo-Festucetum trifolietosum praetutiani shows a high percentage of the en-
demic central-south Apennines chorotype, whose values increase sharply when passing
from the normal spectrum (11%) to the cover spectrum (36.5%) as a consequence of the
coenological dominance of Trifolium noricum subsp. Praetutianum in the community.
Also important is the role of the south-European Orophytes species, which attain per-
centages of 16.6% for Helianthemo-Festucetum trifolietosum and 25% for Ranunculo-
-Adonidetum. Circumboreal and Arctic-Alpine species exhibit relatively low values, com-
pared, that is, to what might be expected for the average altitude of the study area, ranging
fromjust3.7%to9.5%ofthecoverspectrum.
Discussion
The presence of Adonis distorta has been recorded on various massifs (ANZALONE
1947, STEINBERG 1952, TAMMARO 1971), but it is only on the Majella and, to a lesser de-
gree, on the Velino massif that its cover percentages are such as to make it a characteristic
feature of the high altitude landscape. Indeed, its presence is only sporadic on the Gran
Sasso, Vettore, and in the Abruzzo, Lazio and Molise National Park (TAMMARO 1971,
BALLELLI 1999).
On the Majella massif Adonis distorta has been considered a differential species in a
variant of Crepidi-Leontodontetum montani (FEOLI-CHIAPELLA and FEOLI 1977, FEOLI-CHIA-
186 ACTA BOT. CROAT. 67 (2), 2008
DIPIETRO R., PELINO G., STANISCI A., BLASI C.
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PELLA 1983). On the Velino, on the other hand, AVENA and BLASI (1980) consider it a char-
acteristic species of Crepido-Isatidetum allionii (nom. inv. Art. 2). On the basis of our ob-
servations, Adonis distorta tends to form its own communities, which have a precisely
defined location within the micro-geosigmetum of the alpine belt of the Majella. These
communities, in fact, grow only on those special forms of the detritus component which
give rise to alternating strips of broad detritus and fine soil particles.
A preference for this kind of environment, however, does not prevent Adonis distorta
from also playing a role of »differential species« within Crepidi-Leontodontetum, whose
substrate scree-layer is steeper and less stable. Because of the heterogeneity of the original
table of Crepidi-Leontodontetum, we indicate here the lectotypification of the association:
Lectotypus hoc loco design. rel. 44 in Tab. 4 from FEOLI-CHIAPELLAand FEOLI (1977, page
34). The community Crepidi-leontodontetum, more than any other community of the
high-altitude scree communities of the central Apennines, is most floristically similar to
Ranunculo-Adonidetum (Tab. 5). This similarity is essentially due to the common presence
of Adonis distorta (25% in Crepidi-leontodontetum) and to the high frequency and abun-
dance of Leontodon montanus. Nevertheless, the ecological features of these two commu-
nities remain very different, and these differences are reflected, floristically, in the absence
in Crepidi-Leontodontetum of the other two character species of Ranunculo-Adonidetum
(Crepis bithynica and Ranunculus seguieri), as well as that of several other species which
are frequent in Ranunculo-Adonidetum, such as Arenaria grandiflora, Androsace villosa,
Anthyllis vulneraria subsp. pulchella, Minuartia verna, Poa molinerii). At the same time in
Crepido-Leontodontetum there is a higher number of species such as Crepis pygmaea,
Arabis alpina, Cymbalaria pallida, Isatis apennina, Festuca dimorpha, Arenaria berto-
lonii (etc.) which are more strictly linked to scree environments and which do not occur in
Ranunculo-Adonidetum.
Concerning the phytosociological literature references on this argument, it is likely that
the diagnosis of the Isatis apennina and Adonis distorta variant of Crepidi-Leontodontetum
proposed in FEOLI-CHIAPELLA and FEOLI (1977) would also be suitable for representing the
communities with Adonis distorta found on some scree environments of Mount Velino
(AVENAand BLASI 1980) and described as Crepidi-Isatidetum allionii. At present, however,
such a comparison cannot be performed, because in the paper of AVENA and BLASI (1980)
there is no phytosociological table for Crepidi-Isatidetum stands. Moreover, Crepidi-
-Isatidetum is not mentioned at all in a further paper giving an overview of the entire flora
and vegetation of Mount Velino (PETRICCIONE 1993).
In the past Trifolium noricum subsp. praetutianum was considered a characteristic spe-
cies of the association Carici-Salicetum retusae found on the Gran Sasso (BIONDI et al.
1999) and a subassociation differential (PETRICCIONE and PERSIA 1995) of Pedicularido-
-Seslerietum apenninae caricetosum ericetori. Such attributions are in line with the ecol-
ogy of this taxon which, although broadly linked to the dry grasslands of the subalpine belt,
abounds only where the layer of soil is relatively deep and the degree of plant cover in the
relevés ranges between 80 and 100%. From a syntaxonomical point of view neither
Carici-Salicetum or Pedicularido-Seslerietum are suitable for use as references for the
Majella Trifolium noricum subsp. praetutianum stands described in this paper, due, on the
one hand, to the extremely sporadic presence of Salix retusa and, on the other hand, to the
complete lack of Sesleria juncifolia. Instead, the similarities between the Helianthemo-
ACTA BOT. CROAT. 67 (2), 2008 187
TWO ENDEMICS OF THE APENNINES
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188 ACTA BOT. CROAT. 67 (2), 2008
DIPIETRO R., PELINO G., STANISCI A., BLASI C.
Tab. 5. Synoptic table including all the scree associations described at present for the alpine belt of the
central Apennines. 1 – Ranunculo seguirii-Adonidetum distortae ass. nova; 2 – Saxifrago-
-Papaveretum julici from Feoli-Chiapella and Feoli, 1977; 3 – Saxifrago-Papaveretum julici
from BLASI et al. (2005);4–Saxifrago-Papaveretum julici androsacetosum villosae from
BLASI et al. (2005); 5 – Saxifrago speciosae-Silenetum cenisiae from PETRICCIONE (1993);
6–Arabido alpinae-Cerastietum thomasii from (BIONDI et al. 2000); 7 – Crepido-Leontodon-
tetum montanii from FEOLI-CHIAPELLA and FEOLI (1983); 8 – Isatido -Thlaspietum stylosi
from BIONDI et al. (2000); 9 – Achilleo-Saxifragetum azoidis from DIPIETRO et al. (2001).
column no. 172345689
average altitude m a.s.l. (x 10) 253 254 267 270 265 255 280 225 242
total number of relevés 8161468891010
Ranunculo-Adonidetum distortae
Adonis distorta 10025.......
Crepis bythinica 38........
Ranunculus seguieri subsp. seguieri 75.7.13....
char. spesies of Crepido-Leontodontetum montani
Leontodon montanus subsp. montanus 100 81 43 17 63 75 . 30 10
Crepis pygmaea subsp. pygmaea .6214.25. .80.
Senecio squalidus .50. . . . .6010
char. spesies of Saxifrago-Papaveretum julici
Papaver alpinum subsp. ernersti-mayeri .319310088.3340.
Arenaria grandiflora subsp. grandiflora 75.9383100....
Androsace vitaliana subsp. praetutiana 25 12 79 67 88 50 . . .
Saxifraga exarata subsp. ampullacea . . 64 50 75 . 33 . .
char. spesies of Saxifrago speciosae-Silenetum cenisiae
Ranunculus brevifolius 756 . . .87.20.
Valeriana saliunca 25.50.8850. . .
Festuca alfrediana . . . . .8733.80
Achillea barrelieri subsp. barrelieri 75 75 50 17 75 75 33 30 .
Potentilla crantzii subsp. crantzii 38 12 14 . 25 75 . . .
Veronica aphylla . . . . .50.10.
char. spesies of Arabido alpinae-Cerastietum thomasii
Cerastium thomasii 100 75 93 100 88 . 100 . .
Hornungia alpina ......7710.
Arabis alpina . 50 14 33 . . 100 20 .
Draba aspera .25. . . .55. .
char. spesies of Isatido-Thlaspietum stylosi
Isatis apennina .44. . . . .90.
Thlaspi stylosum 38 37 . . . . . 10 .
char. spesies of Achilleo-Saxifragetum azoidis
Acinos alpinus subsp. alpinus 3813......100
Saxifraga aizoides ........100
Achillea barrelieri subsp. mucronulata ........100
Senecio scopolii ........80
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ACTA BOT. CROAT. 67 (2), 2008 189
TWO ENDEMICS OF THE APENNINES
column no. 172345689
average altitude m a.s.l. (x 10) 253 254 267 270 265 255 280 225 242
total number of relevés 8161468891010
char. spesies of Linario-Festucion dimorphae
Galium magellense 50 37 86 83 88 . 44 50 100
Viola magellensis 25 37 71 83 63 . 11 . .
Myosotis ambigens 13757910063....
Alyssum cuneifolium subsp. cuneifolium 25377110050....
Arenaria bertolonii .6717....40
Artemisia umbelliformis subsp. eriantha ..141713....
Festuca dimorpha .6.....4020
Doronicum columnae .18. . . . .30.
Carduus chrysacanthus subsp. chrysacanthus .6......80
Robertia taraxacoides .......2080
Leucanthemum coronopifolium subsp. tenuifolium .6.......
Erysimum majellense .6.......
Cymbalaria pallida .6.......
Heracleum sphondylium subsp. orsinii .......50.
char. spesies of Thlaspietalia rotundifolii and Thlaspietea rotundifolii
Saxifraga oppositifolia subsp. oppositifolia . 31 86 67 100 87 44 10 80
Linaria alpina 50 62 64 67 63 . . 30 .
Salix retusa .13291738....
Iberis saxatilis subsp. saxatilis 25.29.50....
Saxifraga paniculata ..14.25....
Scrophularia hoppii .6.....20.
Campanula cochlearifolia ......11..
Saxifraga caesia ......11..
Saxifraga sedoides subsp. sedoides .6.......
Bellidiastrum michelii .......10.
Rumex scutatus .......20.
char. spesies of Seslerion apenninae and Seslerietalia tenuifoliae
Armeria majellensis 13 13 29 17 38 . 22 . 30
Edrajanthus graminifolius subsp. graminifolius 25.361750. .10.
Carex kitaibeliana subsp. kitaibeliana . . 43 17 63 62 . . .
Andosace villosa subsp. villosa 63.7133100....
Anthyllis vulneraria subsp. pulchella 88.36.63. . .10
Avenula praetutiana 1367.13....
Sesleria juncifolia subsp. juncifolia ..14.25...20
Pedicularis elegans 38.7.13....
Poa molinerii 63.7.13....
Cerastium tomentosum .6717.....
Leontopodium nivale ..21.38....
Festuca violacea subsp. italica ..57.100....
Trinia dalechampii .6.......
Tab. 5. – continued
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190 ACTA BOT. CROAT. 67 (2), 2008
DIPIETRO R., PELINO G., STANISCI A., BLASI C.
column no. 172345689
average altitude m a.s.l. (x 10) 253 254 267 270 265 255 280 225 242
total number of relevés 8161468891010
char. spesies of Elyno myosuroidis-Seslerietea caeruleae
Thymus praecox subsp. polytrichus . 6 7 . 13 62 . . 50
Erigeron epiroticus 131921.38....
Draba aizoides subsp. aizoides 38.21.38. .10.
Pulsatilla alpina subsp. alpina . . 7 .1362.10.
Campanula scheuchzeri s.l. 25 . 7 . 13 . . . 10
Minuartia verna subsp. verna 75.29.50....
Helianthemum oelandicum subsp alpestre ..7.13....
Gentiana verna subsp. verna ..7.13....
Asperula cynanchica .....12...
Phyteuma orbiculare .6.......
Carum flexuosum .6.......
Myosotis alpestris .......20.
Ranunculus pollinensis .......10.
Char. species of Carici-Kobresietea
Silene acaulis s.l. 13 19 64 17 100 87 11 . .
Sedum atratum subsp. atratum 13 13 21 17 25 . . 20 10
Bistorta vivipara ..431763....
Oxytropis campestris ..14.25....
Dryas octopetala subsp. octopetala ..7.13....
Carex capillaris subsp. capillaris ..717.....
Oxytropis neglecta .....12...
Kobresia myosuroides .6.......
Anemone narcissiflora subsp. narcissiflora .......10.
Other species
Poa alpina subsp. alpina 63 87 57 67 50 . 100 20 90
Taraxacum apenninum (group) 38 13 7 . 13 . . . 10
Helianthemum nummularium subsp. obscurum ..7.13....
Euphrasia salisburgensis ..7.1325...
Leontodon hispidus 25.7.13....
Percentages of the sporadic species not included in the table. col. 1: Botrychium lunaria (13);
Plantago atrata subsp. atrata (25); Viola eugeniae subsp. eugeniae (13); Gnaphalium hoppeanum
subsp. magellense (13); Leontodon crispus subsp. crispus (13); col. 2: Campanula tanfanii (21); col.
4: Campanula tanfanii (38); col. 7: Festuca gr. alpina (13); Saxifraga glabella (6); Sedum mage-
llense subsp. magellense, (6; Hypochaeris cretesi,s, 12; Alyssum montanum subsp. montanum, 6;
Alchemilla alpina (6); Saxifraga adscendens subsp. parnassica (6); Cerastium arvense subsp.
strictum (19); col. 8: Festuca circumediterranea (10); Valeriana montana (10); Rumex nebroides
(20); Cerastium arvense subsp. strictum (20); col. 9: Trifolium thalii (10); Crepis aurea subsp.
glabrescens (40); Valeriana montana (10); Cerastium arvense subsp. suffruticosum (?)
Tab. 5. – continued
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-Festucetum trifolietosum praetutiani (described in this paper) and the Helianthemetum
alpestris described about forty years ago in this same area (MIGLIACCIO 1970) are quite evi-
dent. Both communities are physiognomically dominated by Helianthemum oelandicum
subsp. alpestre and Trifolium noricum subsp. praetutianum. Nonetheless, the original de-
scription of Helianthemetum alpestris indicated that this association was the most wide-
spread within the entire Majella alpine belt, whereas in reality Trifolium noricum subsp.
praetutianum is not at all ubiquitous within this belt, its presence being rather sporadic
(even if it tends to be accompanied by high cover indexes where it does occur). In ecologi-
cal terms Trifolium noricum subsp. praetutianum is strongly linked to the microdepres-
sions in the substrate on flattish slopes. It would therefore appear more consistent to con-
sider Trifolium noricum subsp. praetutianum grasslands as a peculiar subassociation of the
more widespread Helianthemo-Festucetum, rather than to generalize and describe Tri -
folium noricum subsp. praetutianum as dominant throughout the Majella alpine belt (as
was done for Helianthemetum alpestris).
Doubts concerning the nomenclatural validity of Helianthemetum alpestris have al-
ready been expressed by BLASI et al. (2003, 2005). Indeed, strict observance of ICPN rules
(Art. 7) leads us to regard this name as invalid. Thus we have chosen to keep Heli-
anthemo-Festucetum as the reference association, and to consider only the subassociation
trifolietosum praetutianii as a syntaxonomical synonym of the invalid Helianthemetum
alpestris of MIGLIACCIO (1970). The original diagnosis of Helianthemetum alpestris con-
tains only one synoptic column, which is incomplete because it lacks the entire component
of companion species. Furthermore this name has not been validated by other authors in
further publications.
Given the dominant physiognomical role of Trifolium noricum subsp. praetutianum in
this new subassociation, and its almost complete absence from the Helianthemo-Festu-
cetum typicum (Tab. 6), we have decided that Trifolium noricum subsp. praetutianum
could be used as a differential at sub-association level only.
As for the higher rank syntaxa, Adonis distorta can be considered a good characteristic
species of Thlaspienion stylosi, the microthermic high altitude sub-alliance of the Apen-
nine endemic alliance Linario-Festucion dimorphae. Therefore we have included Ranun-
culo-Adonidetum in Thlaspietea rotundifolii class and Thlaspietalia rotundifolii order. In-
stead the best position for Helianthemo-Festucetum trifolietosum praetutiani is Seslerion
apenninae, where it is representative of the edapho-mesophilous fringe of this alliance.
Trifolium noricum subsp. praetutianum can nonetheless be present as »ingressive species«
in communities belonging to Arabidion caeruleae or Ranunculo-Nardion.
In some cases the H/Ch ratio has been used as a discriminating criterion in distinguish-
ing the scree communities of Thlaspietea rotundifolii (which show a notable presence of
Chamaephytes) from the dry grasslands of Elyno-Seslerietea (where Hemicryptophytes
are more clearly dominant) (BLASI et al. 2005). For the communities in issue, while the in-
clusion of Trifolium noricum subsp. praetituanum stands in Elyno-Seslerietea is implicit,
the inclusion of Ranunculo-Adonidetum in Thlaspietea is not, because of the high percent-
ages of Hemycryptophytes. It is likely, therefore, that the high H/Ch ratio of Ranun-
culo-Adonidetum is a consequence of the relative flatness of the ground, which has pre-
vented the finer part of the stony substrate from being washed away and has therefore
favoured the development of various Hemicryptophytes.
ACTA BOT. CROAT. 67 (2), 2008 191
TWO ENDEMICS OF THE APENNINES
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192 ACTA BOT. CROAT. 67 (2), 2008
DIPIETRO R., PELINO G., STANISCI A., BLASI C.
Tab. 6. Synoptic table including all the plant communities occurring in the alpine and subalpine belts
of the central Apennines which exhibit floristic similarities with Helianthemo-Festucetum
trifolietosum praetutiani. 1: Helianthemo alpestris-Festucetum italicae from BLASI et al.
2005; Helianthemo alpestris -Festucetum italicae trifolietosum praetutiani subass. nova (2);
Plantago atrata and Leontodon montanus comm. from BLASI et al. 2005 (3); Luzulo
italicae-Festucetum macratherae, from PETRICCIONE and PERSIA (1995)(4); Caricetum
kitaibelianae from MIGLIACCO (1970) (in the original table of MIGLIACCIO (1970), only the
frequency classes were given)(5); Leontopodio nivalis-Seslerietum juncifoliae from BLASI
et al. (2005)(6); Pedicularido-Seslerietum(caricetosum ericetorii) from PETRICCIONE and
PERSIA (1995) (7).
columnno. 1234567
average altitude m a.s.l. (x 10) 238 255 248 221 200 240 234
total numbers of relevés 13 7 9 18 20 13 20
char. species of Helianthemo alpestris-Festucetum italicae
Festuca violacea subsp. italica 100 100 67 94 V 77 25
Leontopodium nivale 100 71 11 . . 100 .
Helianthemum oelandicum subsp alpestre 92 86 . 6 . 85 95
Sempervivum arachnoideum 85 43 . . . 85 .
Poa molinerii 62 100 11 . . 62 .
char. species of Helianthemo alpestris-Festucetum italicae trifolietosum praetutiani
Trifolium noricum subsp. praetutianum .100....8
char. species of Plantago atrata and Leontodon montanus comm.
Leontodon montanus subsp. montanus 8.100....
Achillea barrelieri subsp. barrelieri 15 57 89 . . . 5
Ranunculus brevifolius 231478....
char. species of Luzulo italicae-Festucetum macratherae
Luzula spicata subsp. italica 8..89IV.20
Plantago atrata subsp.atrata 15 29 89 94 V 8 15
char. species of Caricetum kitaibelianae
Polygala alpestris 15 . . 72 V 8 10
Potentilla crantzii subsp.crantzii 62 100 78 . V 38 .
Phyteuma orbiculare 15 14 . 11 IV . 45
Hypericum richeri subsp. richeri ....IV..
Trifolium montanum subsp. montanum ....III..
Erigeron epiroticus 62 29 11 72 III 46 35
Saxifraga adscendens subsp. adscendens 15 14 11 28 III 31 .
Alchemilla colorata ....I..
char. species of Leontopodio-Seslerietum juncifoliae
Sesleria juncifolia subsp. junicifolia 54 . . 6 . 100 95
Carex humilis .....77.
Aster alpinus subsp. alpinus 46....8055
Ranunculus breyninus 46....8565
char. species of Pedicularido-Seslerietum caricetosum ericetori
Pedicularis elegans 54 57 44 6 . 92 75
Carex ericetorum ...17..65
char. species of Seslerion apenninae and Seslerietalia tenuifoliae
Carex kitaibeliana subsp. kitaibeliana 100 100 67 78 . 85 95
Anthyllis vulneraria subsp. pulchella 100 57 67 . V 92 .
Trinia dalechampii 85 100 56 61 . 85 75
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ACTA BOT. CROAT. 67 (2), 2008 193
TWO ENDEMICS OF THE APENNINES
columnno. 1234567
average altitude m a.s.l. (x 10) 238 255 248 221 200 240 234
total numbers of relevés 13 7 9 18 20 13 20
Andosace villosa subsp. villosa 85 43 33 . . 54 40
Edrajanthus graminifolius subsp. graminifolius 62 100 33 11 . 92 80
Armeria majellensis s.l. 69 100 89 83 . 46 80
Avenula praetutiana 85 43 11 44 . 69 .
Carduus carlinifolius subsp. carlinifolius ...33...
Cerastium tomentosum ...47...
Astrantia pauciflora subsp. tenorei ......30
Cynoglossum magellense ......5
char. species of Elyno myosuroidis-Seslerietea caeruleae
Minuartia verna subsp. verna 85 86 89 61 V 69 85
Acinos alpinus subsp. alpinus .145611I . 5
Campanula scheuchzeri 31 14 44 61 . 8 15
Thymus praecox subsp. polytrichus 85 86 44 83 . 46 90
Draba aizoides subsp.aizoides 85 100 44 . V 85 .
Gentiana verna subsp.verna 46 29 33 61 II 31 55
Pulsatilla alpina subsp. alpina 85.2222.3130
Juncus trifidus subsp. monanthos . . . 6 I 23 30
Thesium parnassi .14.6.8.
Astragalus depressus subsp. depressus .1411....
Festuca laevigata subsp. crassifolia ...6.8.
Asperula cynanchica . ..6..15
Anthyllis vulneraria subsp. nana ...33..75
Galium anisophyllon ...28..45
Linum alpinum ......5
char. species of Carici-Kobresietea
Sedum atratum subsp. atratum 31 29 78 . . . 30
Silene acaulis s. l. 77 86 67 39 . 69 65
Oxytropis campestris 77 100 . 6 . 85 55
Bistorta vivipara 814 . 11 . 850
Kobresia myosuroides ...17.1545
Oxytropis neglecta 8......
Gentiana nivalis ....II..
Dryas octopetala subsp. octopetala .....15.
Anemone narcissiflora subsp. narcissiflora ......15
char. species of Linario-Festucion dimorphae, Thlaspietalia rotundifolii, Thlaspietea rotundifolii
Myosotis ambigens 23 71 44 61 . 23 45
Saxifraga oppositifolia subsp. oppositifolia 843226 . 2315
Saxifraga paniculata 69 . . 6 . 85 55
Salix retusa 15 14 . . . 3 15
Arenaria grandiflora subsp. grandiflora 871...315
Galium magellense ..446.8.
Thlaspi stylosum 8.33....
Cerastium thomasii .2978....
Iberis saxatilis subsp. saxatilis 23....92.
Valeriana saliunca 8....15.
Androsace vitaliana subsp. praetutiana .43....5
Artemisia umbelliformis subsp. eriantha .14....5
Tab. 6. – continued
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194 ACTA BOT. CROAT. 67 (2), 2008
DIPIETRO R., PELINO G., STANISCI A., BLASI C.
columnno. 1234567
average altitude m a.s.l. (x 10) 238 255 248 221 200 240 234
total numbers of relevés 13 7 9 18 20 13 20
Ranunculus magellensis ..22....
Carduus chrysacanthus subsp. chrysacanthus ..22....
Stachys alopecuros ...6...
Bellidiastrum michelii ...6...
Alyssum diffusum ...6...
Erysimum majellense ......5
Saxifraga caesia ......10
char. species of Ranunculo Nardion, Nardetalia strictae Nardetea strictae
Viola eugeniae subsp. eugeniae 31 57 67 72 . 15 20
Gentianella columnae . . . 17 . 8 20
Hieracium pilosella . 436717 . . .
Ranunculus pollinensis . . 56 83 IV . .
Botrychium lunaria ...61III.25
Gentiana lutea subsp. lutea ...6..5
Luzula multiflora ...28..5
Rumex nebroides ...22..15
Leucanthemum tridactylites 62......
Crepis aurea subsp. glabrescens ..11....
Alchemilla glaucescens ...87...
Nardus stricta ...22...
Coeloglossum viride ......25
Other species
Poa alpina subsp. alpina 46 14 100 100 . 54 80
Hieracium lactucella 31 . . 83 IV 8 25
Trifolium pratense subsp. semipurpureum 23.1161. .20
Cerastium arvense subsp. strictum .14.83. .50
Gnaphalium hoppeanum subsp. magellense 62 . 100 44 . . 5
Euphrasia salisburgensis 8 . . 22 III 23 30
Trifolium thalii 23.11. II . .
Cerastium arvense subsp. suffruticosum 69 . . . IV 62 .
Percentages of the sporadic species not included in the table. Col. 1: Pinus mugo subsp. mugo(8);
Col. 2: Phleum alpinum subsp. rhaeticum (14); Sedum acre (14); Col. 3: Taraxacum apenninum (33);
Taraxacum glaciale (11); Leontodon crispus subsp. crispus (11); Col. 4: Dianthus sylvestris s.l. (6);
Festuca circumeditteranea (11); Potentilla rigoana (89); Koeleria lobata (11); Rhinantus wettsteinii
(11); Brachypodium genuense (6); Juniperus communis (6); Scabiosa holosericea (6); Silene ciliata
subsp. graefferi (44); Col. 5: Phleum alpinum subsp. rhaeticum, I; Sagina saginoides subsp.
saginoides,I;Carduus defloratus subsp. tridentinus, III; Crepis lucida,I;Helianthemum nummu-
larium subsp. obscurum,IV;Col. 6: Helianthemum nummularium subsp. obscurum (8); Pinus mugo
subsp. mugo (8); Potentilla apennina subsp. apennina (8); Anthyllis montana subsp atropurpurea
(15); Col. 7: Anthyllis montana subsp atropurpurea (35); Scabiosa holosericea (25); Juniperus
communis (20); Potentilla rigoana (30); Festuca circumeditteranea (50); Dianthus sylvestris s.l.
(55); Sedum acre (5); Astragalus sempervirens (10); Biscutella levigata subsp. levigata (5); Cyanus
triumfetti (10); Hieracium amplexicaule (25); Linum capitatum subsp. serrulatum (10); Senecio
doronicum (10); Gentiana dinarica (5); Globularia meridionalis (5); Pedicularis comosa subsp.
comosa (15)
Tab. 6. – continued
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As is the case with the majority of dry grassland communities of the subalpine and al-
pine belts of the central Apennines (BRUNO and FURNARI 1966, LAKU[I] 1969, PETRICCIONE
and PERSIA 1995, BLASI et al. 2003), Helianthemo-Festucetum italicae trifolietosum prae-
tutiani and Ranunculo-Adonidetum distortae, too, exhibit strong floristic similarities with
the parallel communities occurring in the high-altitude zones of the Balkan peninsula
(Carex kitaibeliana, Edrajanthus graminifolius, Thesium parnassi, Anthyllis vulneraria
subsp. pulchella).
The biogeographical links existing between the Central Apennines and the Balkans
(RIVAS-MARTÍNEZ et al., 2001) are confirmed by the chorological spectra, which clearly
show a higher number of Orophilous south-European species than Circumboreal and Arc-
tic-Alpine species. This is probably a consequence of the last ice age (20.000–18.000 years
ago) when the Majella, unlike the other massifs of large dimensions of the Apennines, was
covered by an Icelandic-type glacier of about 30 square kilometres (GIRAUDI 1998). This
thick ice cap, together with the gentle and regular shape of the massif, are likely to have
prevented nunataks from emerging, or to have at least limited their presence. This, in turn,
would have deprived microthermal species native to cold regions of their refuge, and in this
way hindered their re-colonization of the area during the subsequent interglacial period
(CATONICA and MANZI 2002).
A summary of the syntaxa identified so far in the alpine belt of the Majella can be found
in table 7. Due to endemism of Adonis distorta and Trifolium noricum subsp. Pratetutia-
num in the Apennines, there are no associations physiognomically similar to those occur-
ring in the Majella massif on the other side of the Adriatic sea. As far as the high-rank
syntaxa are concerned, the Balkan communities that substitute Helianthemo-Festucetum
italicae trifolietosum preatutiani are likely to be included in orders such as Seslerietalia
tenuifoliae (Seslerion juncifoliae)orCrepidetalia dinaricae (Oxitropidion dinaricae). At
association level Helianthemo-Festucetum italicae is most closely related to Carici kitai-
ACTA BOT. CROAT. 67 (2), 2008 195
TWO ENDEMICS OF THE APENNINES
Tab. 7. Summary of the alpine belt vegetation of the Majella massif.
Crepidi pygmaeae-Leontodontetum montani Vegetation of the unstable talus slopes
Galio – Silenetum acaulis alyssetosum cuneifolii
Carici kitaibelianae – Salicetum retusae
Carici kitaibelianae – Salicetum retusae elynetosum
Vegetation of the stable scree and
relatively steep slopes
Saxifrago speciosae – Papaveretum julici
Saxifrago – Papaveretum julici androsacetosum villosae
Vegetation of the stable scree on mild
slopes of the very summit areas
Ranunculo seguierii-Adonidetum distortae Vegetation of the unstable scree on
mild slopes and »soil with stripes«
Plantago atrata and Leontodon montanus comm.
Helianthemo alpestris – Festucetum italicae
Helianthemo– Festucetum italicae trifolietosum praetutiani
Vegetation of the mild and stable
gravelly substrates
Leontopodio nivalis – Elynetum myosuroidis
Saxifrago speciosae – Papaveretum julici typicum
Leontopodio nivalis – Seslerietum juncifoliae
Vegetation of the windy ridges and
steeper slopes
Gnaphalio magellensis – Plantaginetum atratae
Taraxaco apennini – Trifolietum thalii gnaphalietosum
Luzulo italicae – Nardetum strictae
Vegetation of snowbeds and dolinas
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196 ACTA BOT. CROAT. 67 (2), 2008
DIPIETRO R., PELINO G., STANISCI A., BLASI C.
Tab. 8. Syntaxonomical scheme.
ELYNO-SESLERIETEA Br.-Bl. 1948
Seslerietalia tenuifoliae Horvat 1930
Seslerion apenninae Furnari in Bruno et Furnari 1966
Leontopodio nivalis-Elynenion myosuroidis Blasi et Di Pietro in Blasi, Di Pietro, Fortini et
Catonica 2003
Helianthemo alpestris-Festucetum italicae Blasi, Di Pietro et Pelino 2005
Helianthemo alpestris-Festucetum italicae trifolietosum praetutiani Di Pietro, Pelino,
Stanisci & Blasi subass. nova hoc loco
THLASPIETEA ROTUNDIFOLII Br.-Bl. 1948
Thlaspietalia rotundifolii Br.-Bl. in Br.-Bl. et Jenny 1926
Linario-Festucion dimorphae Avena et Bruno 1975
Thlaspienion stylosi Avena et Bruno 1975
Ranunculo seguierii-Adonidetum distortae Di Pietro, Pelino, Stanisci et Blasi ass. nova
hoc loco
Tab. 9. Complete list of the syntaxa quoted in the text and in the synoptic tables.
Achilleo mucronulatae-Saxifragetum aizoidis Di Pietro, Conti, Vannicelli-Casoni 2001;
Arabido alpinae-Cerastietum thomasii Biondi, Ballelli, Allegrezza et Taffetani 2000;
Arabidion caeruleae Br.-Bl. in Br.-Bl. et Jenny 1926;
Bunion alpini Laku{i~ (1968) 1970;
Caricetum kitaibelianae Migliaccio 1970;
Carici-Crepidetum dinaricae trifolietosum norici Laku{i~ 1964;
Carici kitaibelianae-Helianthemetum alpestris Horvat 1930 (= Laevi-Helianthemetum alpestris);
Carici rupestris-Kobresietea bellardii Ohba 1974;
Edrajantho-Helianthemetum alpestris Horvat 1935;
Carici kitaibelianae-Salicetum retusae Biondi, Ballelli, Allegrezza, Taffetani, Frattaroli, Guitian et
Zuccarello 1999;
Carici kitaibelianae-Salicetum retusae elynetosum Blasi, Di Pietro, Fortini et Catonica 2003;
Crepidetalia dinaricae Laku{i~ 1966;
Crepidi-Leontodontetum montani Feoli-Chiapella et Feoli 1977;
Elyno myosuroidis-Seslerietea caeruleae Br.-Bl. 1948;
Galio magellensis-Silenetum acaulis Blasi, Di Pietro, Fortini et Catonica 2003;
Helianthemetum alpestris Migliaccio 1970;
Helianthemo alpestris-Festucetum italicae Blasi, Di Pietro et Pelino 2005;
Gnaphalio magellensis-Plantaginetum atratae Feoli-Chiapella et Feoli 1977;
Leontodontetum montani Feoli Chiapella et Feoli 1977;
Isatido-Thlaspietum stylosi Migliaccio 1970 corr. Feoli-Chiapella 1983;
Luzulo italicae-Festucetum macratherae (violaceae) Bonin 1978;
Luzulo italicae-Nardetum strictae Biondi, Ballelli, Allegrezza, Frattaroli et Taffetani 1992;
Leontopodio nivalis-Elynenion myosuroidis Blasi et Di Pietro in Blasi, Di Pietro, Fortini et
Catonica 2003;
Leontopodio nivalis-Seslerietum juncifoliae Blasi, Di Pietro et Pelino 2005;
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belianae-Helianthemetum alpestris of the northern Dinarides and to Edrajantho-Helian-
themetum alpestris of the southern Dinarides, even if some floristic similarities to Carici-
-Crepidetum dinaricae are also apparent (HORVAT et al. 1974, REDZIC 2003, SURINA and
DASKOBLER 2005). As far as Ranunculo-Adonidetum distortae is concerned, major similar-
ities with the communities of the alliance Bunion alpini are identifiable,whichintheBal-
kan peninsula behaves as vicariant alliance of the central Apennines Thlaspienion stylosi.
Less evident is the relationship with Saxifragion prenjae communities.
Conclusions
Apart from providing new data on the ecological and coenological aspects of two
Apennine endemisms, Adonis distorta and Trifolium noricum subsp. praetutianum,thispa
-
per also makes a significant contribution to the mapping of the intricate vegetational mo-
saic of the Apennine alpine belt, a mosaic whose most complete expression is to be found
precisely on the Majella. In fact, the massif is a major node within the »European system of
high mountains for the study of global change« (the G.L.O.R.I.A. project), and it has long
been known as an important centre of plant speciation in the south of Europe (the epithets
»magellense« or »majellensis« are very common among the central Apennines endemic
floristic component). Recently, more detailed phytosociological studies have revealed that
the high number of species is matched by a high coenological diversity, which becomes
manifest in the complex mosaic of microgeosigmeta (BLASI et al. 2005). The identification
of two new syntaxonomical types inside the alpine bioclimatic belt of the Apennines results
ACTA BOT. CROAT. 67 (2), 2008 197
TWO ENDEMICS OF THE APENNINES
Linario-Festucion dimorphae Avena et Bruno 1975;
Luzulo italicae-Festucetum macratherae Bonin 1978;
Nardetea strictae Riv. Goday et Borja C. 1961;
Nardetalia strictae Oberd. ex Preisg. 1949;
Oxytropidion dinaricae Laku{i} 1966;
Pedicularido elegantis-Seslerietum apenninae caricetosum ericetori (Petriccione 1991)
Petriccione et Persia 1995;
Ranunculo-Nardion Bonin 1972;
Saxifragion prenjae Laku{i} 1966;
Saxifrago speciosae-Papaveretum julici Feoli-Chiapella et Feoli 1977;
Saxifrago speciosae-Papaveretum julici androsacetosum villosae Feoli-Chiapella et Feoli ex Blasi,
Di Pietro et Pelino 2004;
Saxifrago speciosae-Silenetum cenisiae Petriccione 1993;
Seslerion apenninae Furnari in Bruno et Furnari 1966;
Seslerietalia tenuifoliae Horvat 1930;
Seslerion juncifoliae Horvat 1930;
Taraxaco apennini-Trifolietum thalii gnaphalietosum magellensis Blasi, Di Pietro et Pelino 2004;
Thlaspienion stylosi Avena et Bruno 1975;
Thlaspietalia rotundifolii Br.-Bl. in Br.-Bl. et Jenny 1926;
Thlaspietea rotundifolii Br.-Bl. 1948.
Tab. 9. – continued
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in an increase in the degree of »biodiversity«, at the level of both community and land-
scape. Table 7 provides a summary of the syntaxa identified so far in the alpine belt of the
Majella.
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200 ACTA BOT. CROAT. 67 (2), 2008
DIPIETRO R., PELINO G., STANISCI A., BLASI C.
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... Further important contributions and many new floristic records and newly described or re-evaluated taxa were published in recent times (Conti 1988a, Conti & Pellegrini 1989, Marcantonio 2000, Bruschi et al. 2006a, 2006b, Conti & Uzunov 2011, Bartolucci & Walter 2015. Moreover, several published vegetation reports contain relevant information on species occurrence and distribution particularly those by Migliaccio (1966Migliaccio ( , 1970, Feoli & Feoli-Chiapella (1975, 1977), Feoli Chiapella (1983, Bonin (1978), Pirone (1987aPirone ( , 1987bPirone ( , 1997Pirone ( , 1998Pirone ( , 2016, Biondi et al. (1988), Tammaro & Pirone (1995, Stanisci (1997), , Pirone & Cutini (2002), Pirone & De Nuntiis (2002), Pirone et al. (2005Pirone et al. ( , 2010Pirone et al. ( , 2014, Blasi et al. (2005), Di Fabrizio et al. (2006), Frattaroli et al. (2006), Di Pietro et al. (2008) and Ciaschetti et al. (2016b). ...
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This paper deals with the occurrence of the main plant community types occurring within the alpine bioclimatic belt in the Central Apennines. The study area was represented by three mountain groups, the Gran Sasso, Maiella and Laga mountains. These three mountain chains are those in which, out of the entire Apennine area, alpine vegetation is best expressed. The vegetation has been investigated following the Braun-Blanquet phytosociological approach. During the field work 135 relevés were collected. The raw data were further treated with multivariate analysis procedures to find objective clusters on a statistical basis. Several plant community types, belonging to different phytosociological classes, such as Elyno-Seslerietea, Salicetea herbaceae, Thlaspietea rotundifolii, Asplenietea trichomanis, were identified. In particular two new phytosociological associations, Galio-Silenetum acaulis and Potentillo-Festucetum alfredianae, and several new subassociations are described in this paper. In order to express the occurrence and autonomy of the Apennine alpine bioclimatic belt also in coenological terms, a new Seslerion apenninae sub-alliance, named Leontopodio-Elynenion, is here proposed.
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vgr . LAVORI DELLA SOCIETA ITALIANA DI BIOGEOGRAFIA NUOVA SEBIE — VOL. II 1971 J» r.» Dz'rett07'e responsabile e reclattore: B. BACCETTI (Sieua) Consulentz‘ e(lz'toriali: R. AGOSTINI (Napoli), E. GIANNINI (Siena), H. JANETSCHEK (Innsbruck), M. LA GRECA (Ca,tam'a.), R. P1cH1 SERMOLLI (Ge-nova), S. RUFFO (Verona), S. L. TUXEN (Copenaghen), P. ZANGHERI (Forli) IL POPOLAMENTO ANIMALE E VEGETALE DELL’ APPENNINO CENTRALE TIPOGRAFIA VALBONESI - FORIJ Amvo 1971
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High mountain vegetation on calcareous geological foundation is prevalent in the alpine and subalpine belts of mountains on the Balkan peninsula. The class Elyno-Seslerietea is represented in alpine and subalpine communities of meadows and pastures in these communities, species from Dinaric, Balkan, Illyrian, and the south-eastern European floral element play dominant role. In the investigation area, the class is differentiated into the following orders: Seslerietalia juncifoliae (=tenuifoliae) Horvat 1930 with alliances Seslerion juncifoliae (=tenuifoliae) Horvat 1930 in alpine belts and Festucion bosniacae (=pungentis) Horvat 1930 in the subalpine belts and Festucion paniculatae calcicolum (Redzic, Lakusic et al., 1984) Redzic in North-western part; Onobrychydo-Seslerietalia Horvat 1949 with alliances Onobrycho-Festucion Horvat 1936, Festuco-Knaution longifoliae Horvat 1949, Edraiantho-Seslerion Horvat 1949, Seslerion rigidae Zolyomy 1939 and Seslerio-Festucion xanthinae Horvat 1949 in the eastern and southern part; Order Crepidetalia dinaricae (=urumovii) Lakusic 1966 with alliances Campanulion linifolii Lakusic 1964, Festucion pseudoxanthinae Lakusic 1968, Oxytropidion dinaricae (=urumovii) Lakusic 1966 in the southeastern Dinaric Alps; order Edraintho-Seslerietalia robustae Ordo nov prov. with alliances Seslerion robustae (Lakusic et al. 1982) and Seslerio-Edraianthion pumilii Redzic all. nova prov. in the coastal side of oro-mediterranean region of Balkans. © 2003, Universita degli Studi di Roma 'La Sapienza'. All rights reserved.
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Relict rock glaciers have been identified in the Maiella Mountain massif, Central Apennines (42° N). They are located at elevations of 2500–2300 m a.s.l. and probably mark the uppermost and southernmost limit of Pleistocene discontinuous alpine permafrost in the Apennine peninsula. Des restes d'anciens glaciers rocheux ont été identifiés dans le massif montagneux de Maiella, au centre de Appennins (42° N). Ils sont localisés à des altitudes comprises entre 2500 et 2300 m et marquent probablement la limite inférieure et méridionale du pergélisol pléistocène discontinu dans la péninsule italienne.