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Syntaxonomic survey of the class Peganoharmalae‐Salsoletea vermiculatae Br.‐Bl. & O. Bolos1958 in ItalyS. Brullo a , G. Giusso del Galdo a , R. Guarino b , P. Minissale a , S. Sciandrello a & G.Spampinato ca Department of Biological, Geological and Environmental Sciences, University of Catania,Via A. Longo 19, Catania, 95125, Italyb Department of Environmental Biology and Biodiversity, University of Palermo, via Archirafi38, Palermo, 90123, Italyc Department S.T.A.F.A, University of Reggio Calabria, P.zza San Francesco 4, Gallina (RC),89061, Italy
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To cite this article: S. Brullo, G. Giusso del Galdo, R. Guarino, P. Minissale, S. Sciandrello & G. Spampinato (): Syntaxonomicsurvey of the class Pegano harmalae‐Salsoletea vermiculatae Br.‐Bl. & O. Bolos 1958 in Italy, Plant Biosystems - AnInternational Journal Dealing with all Aspects of Plant Biology: Official Journal of the Societa Botanica Italiana,DOI:10.1080/11263504.2012.717544
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Syntaxonomic survey of the class Pegano harmalae-Salsoletea
vermiculatae Br.-Bl. & O. Bolos 1958 in Italy
S. BRULLO1, G. GIUSSO DEL GALDO1, R. GUARINO2, P. MINISSALE1,
S. SCIANDRELLO1, & G. SPAMPINATO3
1Department of Biological, Geological and Environmental Sciences, University of Catania, Via A. Longo 19, 95125 Catania,
Italy; 2Department of Environmental Biology and Biodiversity, University of Palermo, via Archirafi 38, 90123 Palermo, Italy
and 3Department S.T.A.F.A., University of Reggio Calabria, P.zza San Francesco 4, 89061 Gallina (RC), Italy
AbstractAfter a general outline on the syntaxonomical framework of the class Pegano harmalae-Salsoletea vermiculatae Br.-Bl. & O.Bolos 1958, the occurrence of this vegetation in the Italian territory is examined. In Italy, this vegetation is mostly found onclayish or marly substrata, particularly if slightly enriched with nitrates and chlorides. With reference to the Rivas-Martınezbioclimatic classification, the Italian stands have an infra- to thermo-Mediterranean thermotype (marginally up to the meso-Mediterranean one) with dry or arid ombrotype. In particular, this vegetation is well represented in Sicily, while in Sardiniaand along the Italian Peninsula, it progressively becomes more rare and localized. Altogether, 21 associations are recognized,11 of which are described here for the first time. The surveyed associations are ascribed to the following alliances: Salsolooppositifoliae-Suaedion mollis (linked to less disturbed and more thermo-xeric habitats) and Artemision arborescentis (moredisturbed and less thermo-xeric habitats). Finally, some conservation issues are discussed.
Keywords: Phytosociology, Pegano-Salsoletea, Italy, conservation, habitat
Introduction
The class Pegano-Salsoletea Br.-Bl & O. Bolos 1958
groups halo-nitrophilous scrublands colonizing
marly to clayey substrata and evaporitic sediments
in the Mediterranean, Saharan-Atlantic and Macar-
onesic Regions, particularly under thermo-xeric
climatic conditions (Rivas-Martınez et al. 2002).
This vegetation is characterized by succulent Cheno-
podiaceae with a chamaephytic to nanophanerophytic
habit, such as Salsola (S. oppositifolia, S. vermiculata),
Suaeda (S. vera, S. pruinosa), Atriplex (A. halimus, A.
glauca), frequently found together with other repre-
sentative species, belonging to the genera Peganum
(P. harmala), Artemisia (A. arborescens, A. herba-alba,
A. campestris), Anabasis (A. articulata), Haloxilon (H.
articulatum), and Limonium sp.pl.
The nomenclatural vicissitudes of the class being
studied are rather complicated: it was first described
for the Ebro Valley (Spain) by Braun-Blanquet and
Bolos (1958) to emphasize the floristic autonomy of
the order Salsolo. vermiculatae-Peganetalia harmalae
and the alliance Salsolo-Peganion, previously framed
by the same authors (Braun-Blanquet & Bolos 1954)
into the class Chenopodietea Br.-Bl in Br.-Bl et al. 1952.
Further researches widened the range of the class
to a great deal of the Iberian Peninsula and of the
southern Mediterranean territories (Rivas Goday &
Rivas-Martınez 1963; Bolos et al. 1970; Rigual
Magallon 1972; Peinado & Martinez Parras 1984;
Brullo et al. 1985; Peinado et al. 1986, 1989; Gehu
& Biondi 1986; Biondi 1988; Alcaraz et al. 1991;
Rivas-Martınez et al. 1993, 2002; Costa et al. 2000;
Alonso & De la Torre 2002).
In the meantime, several new orders and alliances
were added to the class (Appendix 1): the first
addition was proposed by Rivas Goday and Rigual
Magallon (1958), who described the Salsolo-Fagonion
creticae, replacing the Salsolo-Peganion in the Alicante
Province. Successively, Rivas Goday and Rivas-
Martınez (1963) included new orders in the
class two: the Onopordo-Salsoletalia Rivas Goday &
Correspondence: S. Brullo, Department of Biological, Geological and Environmental Sciences, University of Catania, Via A. Longo 19, 95125 Catania, Italy.
Email: [email protected]
Plant Biosystems, 2012; 1–26, iFirst article
ISSN 1126-3504 print/ISSN 1724-5575 online ª 2012 Societa Botanica Italiana
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Rivas-Martınez 1963 (with the alliances Salsolo-
Peganion and Onopordion arabici Br.-Bl. & O. Bolos
1958) and the Atriplicetalia glaucae Rivas Goday &
Rivas-Martınez 1963 (with the new alliances Salsolo-
Carthamion Rivas Goday & Rivas-Martınez 1963 and
Haloxilo-Atriplicion Rivas Goday & Rivas-Martınez
1963). All these syntaxa were reconsidered and
validated by Rigual Magallon (1972), who included
into them several associations and proposed the new
alliance Salsolo-Suaedion.
In later years, some more syntaxa have been
proposed for the Iberian Peninsula: the two alliances
Salsolo-Artemision and Atriplicion glaucae were de-
scribed as nomina nuda by Folch (1981); the
Medicagini citrinae-Lavaterion arboreae, proposed by
Bolos and Vigo (1984); the order Helichryso-Santo-
linetalia squarrosae, proposed by Peinado and Martı-
nez Parras (1984) to group the alliances Artemisio
glutinosae-Santolinion rosmarinifoliae Costa 1975 and
Santolinion pectinato-canescentis Peinado & Martınez-
Parras 1984, that, in the opinion of the same authors
and of Peinado et al. (1986), are closely related to the
glareicolous vegetation of the Andryaletalia ragusinae
Rivas Goday ex Rivas Goday & Esteve 1972.
For the Canary Islands, the endemic order
Chenoleoidetalia tomentosae and the alliance Cheno-
leoidion tomentosae were described by Sunding
(1972). His proposal has been followed by Rivas-
Martınez et al. (1993), who proposed, in addition,
the new endemic order Forsskaoleo angustifoliae-
Rumicetalia lunariae, with the two alliances Launaeo
arborescentis-Schizogynion sericeae and Artemision thus-
culae-Rumicion lunariae.
Also, some synanthropic plant communities char-
acterized by pantropical chorotypes was ascribed to the
class Pegano-Salsoletea: this is the case of the order
Nicotiano glaucae-Ricinetalia communis, with the alli-
ance Nicotiano glaucae-Ricinion communis, described
for the Canary Islands by Rivas-Martınez et al. (1999).
This is the case, as well, of the Lycio europaei-Ipomoeion
purpureae, proposed by Bolos (1988) for the Mediter-
ranean Region and framed into the Ipomoeetalia
purpureae Oberd. ex O.Bolos 1988, order of the class
Ruderali-Manihotetea utilissimae Leonard in Taton
1949. These last two syntaxa have been synonymized
by Rivas-Martınez et al. (2002) with the Salsolo-
Peganetalia and Pegano-Salsoletea, respectively, but the
alliance Lycio europaei-Ipomoeion purpureae has been
kept valid. A syntaxonomical survey of the class at
issue in the Iberian and Macaronesian territories was
presented by Rivas-Martınez et al. (2001, 2002).
For the semi-deserts of Cyrenaica (Libya), the
order Hammado scopariae-Anabasietalia articulatae
was described by Brullo and Furnari (1996) and
framed into the class Pegano-Salsoletea with the
alliance Limonion tubiflori, a Libyan vicariant of the
North-Egyptian Thymelaeion hirsutae Tadros & Atta
1958. Furthermore, the Artemision arborescentis has
been described by Gehu and Biondi (1986) for the
central Mediterranean territories and the Argiranthe-
mo succulenti-Calendulion maderensis for the Madeira
Islands by Capelo et al. (2000).
At the centre of the Mediterranean Basin, the
Pegano-Salsoletea vegetation has been found in the
southern part of the Italian Peninsula (Biondi 1988;
Biondi et al. 1992, 1994a; Pirone 1995; Brullo et al.
2001), in Sardinia (Biondi et al. 1988; Biondi &
Mossa 1992; Biondi et al. 1994b), Corsica (Gehu &
Biondi 1986; Gehu et al. 1986, 1988), and Sicily
(Brullo et al. 1980, 1985, 1988, 2000; Brullo &
Siracusa 2000). In all these sites, the vegetation
hitherto described belongs to the order Salsolo
vermiculatae-Peganetalia harmalae, even if the occur-
rence of stands ascribable to the Nicotiano glaucae-
Ricinetalia communis cannot be excluded. Aim of this
study is to make a point about the consistence of the
Pegano-Salsoletea vegetation in Italy, basing on
literature and original data.
Material and methods
Our survey is based on 263 phytosociological releves,
partially taken from literature. On the whole, 21
associations have been recognized, 11 of which are
new. For the identification of the new associations,
besides the structural and floristic homogeneity,
particular care was paid to the dominant species,
often denoting slightly different edaphic and micro-
climatic conditions in the vegetation at issue. The
numerical analysis has been performed on the cover
data of 72 selected releves (Van der Maarel 1979),
three for each plant community (Table S1), pro-
cessed by the software package Syn-Tax 2000
(Podani 2001). The releves selected for the numerical
analysis are those whose ratio between the total
number of species and the character ones was
minimized (Westhoff & Van der Maarel 1973; Noest
et al. 1989). The Euclidean distance algorithm was
used to produce the dissimilarity matrix, and the
simple average criterion was adopted to produce the
classification dendrogram. Vegetation data were inter-
preted in terms of syntaxonomical classification, based
on cover and floristic affinities, following the Zurich-
Montpellier approach (Braun-Blanquet 1964).
In the list commented below, the associations of
each alliance are alphabetically reported. The struc-
ture of the entries mostly follows the proposal of
Dengler et al. (2003), as well as the recommenda-
tions of the International Code of Phytosociological
Nomenclature (Weber et al. 2000). The bioclimatic
classification follows Rivas-Martınez (1997) and
Rivas-Martınez et al. (2004a, 2004b). For the taxa
nomenclature, some vascular plant checklists have
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been followed (Conti et al. 2005; Giardina et al.
2007; Greuter 2008).
Results
The Pegano-Salsoletea vegetation of Italy is chiefly
found on clayish and marly soils, particularly if
slightly enriched with nitrates and chlorides, in
places with infra- or thermo-Mediterranean thermo-
type (marginally up to the meso-Mediterranean one)
with dry or arid ombrotype. This vegetation is well
represented in Sicily, while in Sardinia and southern
Italy, it becomes more rare and localized. The
following differential/character species of Pegano-
Salsoletea and Salsolo. vermiculatae-Peganetalia har-
malae are frequently found in Italy: Lycium intrica-
tum, Atriplex halimus, Capparis sicula, Moricandia
arvensis, and Asparagus stipularis.
According to our surveys (Table I), the vegetation
can be ascribed to the following two alliances
(Appendix 2):
(A) Salsolo. oppositifoliae-Suaedion mollis Rigual
1972: vegetation of natural habitats, sometimes
favoured by human activities, in remarkable
xeric conditions. The associations ascribed to
this alliance have their primary stands on clayey
badlands (locally named ‘‘calanchi’’), coastal
slopes and along the borders of salt marshes
and coastal lagoons, slightly above the water-
table. They can be considered as permanent
edapho-xerophilous plant communities domi-
nated by relatively few highly specialized
species. In former contributions, these associa-
tions were ascribed to the Salsolo vermiculatae-
Peganion harmalae (see Brullo et al. 1980, 1985,
2000, 2001; Brullo & Siracusa 2000; Bartolo
et al. 1982, 1990). According to Rivas-Martı-
nez et al. (2002), the latter name applies to
plant communities linked to meso-supramedi-
terranean semiarid and lower-dry semiconti-
nental bioclimates of the inner Iberian
Peninsula, while those of Italy, for their ecology
and floristic settlement, should be ascribed to
the Salsolo oppositifoliae-Suaedion mollis (Rivas-
Martınez 2011). Indeed, the Italian commu-
nities are well characterized by the occurrence
of Salsola oppositifolia, Salsola vermiculata and
Suaeda vera, all linked to remarkable thermo-
xeric habitats falling in the infra- and thermo-
Mediterranean bioclimatic belt.
(B) Artemision arborescentis Gehu & Biondi in Gehu
et al. 1986: vegetation of disturbed habitats,
particularly frequent on clayish slopes next to
human settlements, roadsides, and folds, on
soils enriched with nitrogen and phosphates. It
is found in the thermo- and, marginally,
meso-Mediterranean bioclimatic belt, under
some more mesic conditions than the phyto-
coenoses ascribed to the previous alliance.
Differential species are Artemisia arborescens
and Anagyris foetida, with the former one often
dominant in such vegetation.
Also, the numerical analysis strongly supports the
recognition of two groups, clearly corresponding to
the two alliances occurring in Italy (Figure 1).
Salsolo oppositifoliae-Suaedion mollis
Suaedo verae-Atriplicetum halimi Biondi
1988 – Table I (21–22)
Holotypus: rel.2, tab.4, Biondi (1988).
Character species: Suaeda vera.
Structure and ecology: Markedly, halo-nitrophilous
association colonizing coastal rocky habitats (marls,
conglomerates, limestone and sandstone) were di-
rectly influenced by the sea-spray and dunged by the
seabirds. The physiognomy is given by the dense
bushes of Suaeda vera and Atriplex halimus, normally
achieving high cover values. According to Biondi
(1988), several different sub-associations can be
identified, depending on the type of substratum
and on the ecotonal contacts with other plant
communities (see Biondi 1988).
Distribution: Central and southern Italy (Figure 2,
online Appendix), along the Adriatic coast, and
Tremiti Islands (Biondi 1988; Pirone 1995).
Camphorosmo monspeliacae-Atriplicetum ha-
limi Biondi, Ballelli & Taffetani 1992 – Table I
(18)
Holotypus: rel.2, tab.2, Biondi et al. (1992).
Character species: Camphorosma monspeliaca.
Structure and ecology: This association is found
inland, where it colonizes the steepest part of clayey
badlands at 200 to 600 m a.s.l., within the meso-
Mediterranean subhumid bioclimatic belt (Biondi
et al. 1992). The structure is given by a prostrate
layer of Camphorosma monspeliaca mixed with scat-
tered bushes of Atriplex halimus. On less sloping sites,
this vegetation comes into contact with the perennial
dry grasslands of Camphorosmo monspeliaci-Lygeetum
sparti Biondi et al. 1992.
Distribution: Basilicata, in the valley of Basento River
(Figure 2 online Appendix).
Atriplici halimi-Polygonetum tenoreani Biondi,
Ballelli & Taffetani 1992 – Table I (19)
Holotypus: rel.2, tab.3, Biondi et al. (1992).
Character species: Polygonum tenoreanum, endemic to
southern Italy.
Structure and ecology: This association replaces the
Camphorosmo monspeliacae-Atriplicetum halimi on
Pegano-Salsoletea in Italy 3
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Tab
leI.
Syn
op
tic
tab
leo
fth
eP
egano-
Sals
olet
ave
get
atio
nin
Ital
y.
(con
tinued
)
4 S. Brullo et al.
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Tab
leI.
(Con
tinued
).
1-
Cappari
do
sicu
lae-
Sals
olet
um
oppos
itifol
iae
(SE
Sic
ily
(aft
erB
arto
loet
al.
19
82
,T
ab.
24
,su
bS
uaed
o-S
als
olet
um
oppos
itifol
iae)
.
2-
Cappari
do
sicu
lae-
Sals
olet
um
oppos
itifol
iae
(Sic
ily,
Ad
ran
o;
Bru
llo
&S
irac
usa
20
00
,T
ab.1
7,
sub
Suaed
o-S
als
olet
um
oppos
itifol
iae)
.
3-
Cappari
do
sicu
lae-
Sals
olet
um
oppos
itifol
iae
(Sic
ily,
Po
rto
Em
ped
ocl
e,T
ab.4
B).
4-
Asp
ara
goalb
i-S
als
olet
um
oppos
itifol
iae
(Sic
ily
and
SC
alab
ria;
Tab
.4A
).
5-
Thapsi
opel
agi
cae-
Sals
olet
um
oppos
itifol
iae
(Lam
ped
usa
;B
arto
loet
al.
19
90
,T
ab.1
0,
sub
Suaed
o-S
als
olet
um
oppos
itifol
iae)
.
6-
Lim
onio
opule
nti
-Sals
olet
um
oppos
itifol
iae
(Sic
ily,
Po
rto
Em
ped
ocl
e;B
rullo
etal
.1
98
0,
Tab
.1).
7-
Lim
onio
cata
nzaro
i-S
als
olet
um
oppos
itifol
iae
(Sic
ily,
Cap
oB
ian
co;
Bru
llo
etal
.1
98
5,
Tab
.1).
8-
Lim
onio
cata
nzaro
i-S
als
olet
um
oppos
itifol
iae
(Sic
ily,
Rib
era,
Tab
.2A
).
9-
Lim
onio
calc
ara
e-S
uaed
etum
ver
ae
(CS
icily,
S.
An
gel
oM
uxar
o,
Tab
.2B
).
10
-S
als
olet
um
agr
igen
tinae
(CS
icily;
Bru
llo
etal
.1
98
5,
Tab
.2).
11
-S
als
olet
um
agr
igen
tinae
(ES
icily,
Ad
ran
o;
Bru
llo
&S
irac
usa
20
00
,T
ab.1
8).
12
-S
als
olet
um
agr
igen
tinae
(Sic
ily,
Tab
.2C
).
13
-S
als
olo
oppos
itifol
iae-
Suaed
etum
pel
agi
cae
(Lam
ped
usa
;B
arto
loet
al.
19
90
,T
ab.1
1).
14
-S
uaed
over
ae-
Lim
onia
stre
tum
mon
opet
ali
(Lam
ped
usa
;B
arto
loet
al.
19
90
,T
ab.1
2).
15
-S
uaed
over
ae-
Lim
onia
stre
tum
mon
opet
ali
(Sic
ily,
Tab
.3A
).
16
-H
alim
iono
por
tula
coid
is-S
als
olet
um
oppos
itifol
iae
(SS
icily,
Man
fria
;B
rullo
etal
.2
00
0T
ab.9
).
17
-H
alim
iono
por
tula
coid
is-S
als
olet
um
oppos
itifol
iae
(Sic
ily,
Rib
era,
Tab
.4C
).
18
-C
am
phor
osm
om
onsp
elia
cae-
Atr
iplice
tum
halim
i(B
asilic
ata;
Bio
nd
iet
al.
19
92
,T
ab.2
).
19
-A
trip
lici
halim
i-P
olyg
onet
um
tenor
eani
(Bas
ilic
ata;
Bio
nd
iet
al.
19
92
,T
ab.3
).
20
-A
trip
lici
halim
i-H
alim
ionie
tum
por
tula
coid
is(S
icily,
Cat
ania
;T
ab.3
B).
21
-S
uaed
over
ae-
Atr
iplice
tum
halim
i(A
pu
lia,
Tre
mit
iIs
lan
ds;
Bio
nd
i1
98
8,
Tab
.4).
22
-S
uaed
over
ae-
Atr
iplice
tum
halim
i(A
bru
zzo
;P
iro
ne
19
95
,T
ab.2
1,
ril.
1-3
).
23
-A
spara
gost
ipula
ris-
Sals
olet
um
ver
mic
ula
tae
typ
icu
m(S
ard
inia
,C
apo
S.
Elia;
Bio
nd
i&
Mo
ssa
19
92
,T
ab.1
5,
rel.
3-8
,su
bS
als
olo
ver
mic
ula
tae-
Atr
iplice
tum
halim
i).
24
-A
spara
gost
ipula
ris-
Sals
olet
um
ver
mic
ula
tae
typic
um
(Sar
din
ia,
Cal
aM
osc
a,T
ab.5
A,
rel.
1-4
).
25
-A
spara
gost
ipula
ris-
Sals
olet
um
ver
mic
ula
tae
art
emis
ieto
sum
arb
ores
centi
s(S
ard
inia
,C
apo
S.
Elia;
Bio
nd
i&
Mo
ssa
19
92
,T
ab.1
5,
rel.
9-1
3,
sub
Sals
olo
ver
mic
ula
tae-
Atr
iplice
tum
halim
i).
26
-A
spara
gost
ipula
ris-
Sals
olet
um
ver
mic
ula
tae
art
emis
ieto
sum
arb
ores
centi
s(S
ard
inia
,C
ala
Mo
sca,
Tab
.5A
,re
l.5
-10
).
27
-A
spara
gost
ipula
ris-
Sals
olet
um
ver
mic
ula
tae
art
emis
ieto
sum
arb
ores
centi
s(S
ard
inia
,C
agliar
i;B
ion
di
etal
.1
99
4b
,T
ab.2
,su
bA
trip
lici
halim
i-A
rtem
isie
tum
arb
ores
centi
s).
28
-L
ycio
euro
paei
-Art
emis
ietu
marb
ores
centi
s(S
icily,
Tab
.7B
).
29
-L
ycio
euro
paei
-Art
emis
ietu
marb
ores
centi
s(S
icily,
Agir
a;C
ost
anzo
etal
.,2
00
5,
Tab
.15
).
30
-L
imon
ioop
tim
ae-
Sals
olet
um
oppos
itifol
iae
(Sic
ily,
Po
nte
Cin
qu
eA
rch
i,T
ab.7
A).
31
-C
oron
illo
vale
nti
nae-
Art
emis
ietu
marb
ores
centi
s(S
icilia
,C
alta
vult
uro
,T
ab.7
C).
32
-L
ycio
intr
icati
-Sals
olet
um
oppos
itifol
iae
(Sic
ily,
Man
fria
;B
rullo
etal
.2
00
0,
Tab
.8su
bS
uaed
over
ae-
Sals
olet
um
oppos
itifol
iae)
.
33
-L
ycio
intr
icati
-Sals
olet
um
oppos
itifol
iae
(Sic
ily,
Tab
.5B
).
34
-A
trip
lici
halim
i-A
rtem
isie
tum
arb
ores
centi
ssa
lsol
etum
oppos
itifol
iae
(Sic
ily,
Sic
ulian
a,T
ab.6
A).
35
-A
trip
lici
halim
i-A
rtem
isie
tum
arb
ores
centi
ssa
lsol
etum
oppos
itifol
iae
(Sic
ily,
Agir
a;C
ost
anzo
etal
.2
00
5,
Tab
.14
).
36
-A
trip
lici
halim
i-A
rtem
isie
tum
arb
ores
centi
shalim
ionet
usu
mpor
tula
coid
is(S
icily,
Fo
ceS
imet
o;
Sar
din
ia;
Tab
.6B
).
37
-A
trip
lici
halim
i-A
rtem
isie
tum
arb
ores
centi
sty
pic
um
(Ap
ulia,
Gar
gan
o&
Tre
mit
iIs
lan
ds;
Bio
nd
i1
98
8,
Tab
.1).
38
-A
trip
lici
halim
i-A
rtem
isie
tum
arb
ores
centi
sty
pic
um
(NS
ard
inia
;B
ion
di
etal
.1
98
8,
Tab
.1).
39
-A
trip
lici
halim
i-A
rtem
isie
tum
arb
ores
centi
sty
pcu
m(N
Cal
abri
a;B
ion
di
etal
.1
99
4a,
Tab
.8).
40
-M
edic
agi
ni
arb
orea
e-S
als
olet
um
oppos
itifol
iae
(Sic
ily,
Lic
ata,
Tab
.5C
).
Pegano-Salsoletea in Italy 5
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steep, heavily eroded badlands formed by alternating
clayey and sandy layers. Due to the fast weathering of
the dwelled sites, this vegetation never achieves high
cover values. The structure is given by clumps of
Polygonum tenoreanum, mixed with more scattered
bushes of Atriplex halimus.
Distribution: Basilicata, in the valley of Basento River
(Figure 3 online Appendix).
Limonio opulenti-Salsoletum oppositifoliae
Brullo, Grillo & Scalia 1980, nom. inver.
propos. – Table I (6).
Syn.: Salsolo oppositifoliae-Limonietum opulenti Brullo,
Grillo & Scalia 1980
Holotypus: rel.14, tab.1, Brullo et al. (1980).
Character species: Limonium opulentum, Herniaria
empedocleana and Suaeda kocheri, all endemic to S-
Sicily, plus Reaumuria vermiculata, a South-Medi-
terranean and Saharo-Syndic element, in Italy found
only in few places along the southern coast of Sicily.
Structure and ecology: Chamaephytic and nanopha-
nerophytic vegetation colonizing coastal badlands
and steep heavily eroded marly outcrops, within the
lower thermo-Mediterranean lower dry bioclimate
belt. The most abundant species are Salsola opposi-
tifolia, Atriplex halimus, and Suaeda vera. On less
sloping sites, this vegetation comes in contact with
the Euphorbietum dendroidis Guinochet in Guinochet
and Drounieau 1944. According to the Art 42 of the
ICPN (Weber et al. 2000), it is proposed here to
modify the name of the association, since Salsola
oppositifolia is the species physiognomically
dominant.
Distribution: Sicily, near Porto Empedocle and
Agrigento (Figure 5 online Appendix). Very loca-
lized but, until a recent past, relatively common in
this area, both inland and along the coast (Brullo
et al. 1980). At present times, a deregulated urban
sprawl, stone-quarries and landfills are seriously
threatening this plant community.
Salsoletum agrigentinae Brullo, Guglielmo &
Pavone 1985 – Table I (10–12); Table II C
Holotypus: rel.4, tab.2, Brullo et al. (1985).
Figure 1. Dendrogram of the releves of Table S1. Algorithm: simple average, Euclidean distance.
6 S. Brullo et al.
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Tab
leII
.L
imon
ioca
tanzaro
i-S
als
olet
um
oppos
itifol
iae
(A),
Lim
onio
calc
ara
e-S
uaed
etum
ver
ae
(B),
Sals
olet
um
agr
igen
tinae
(C)
(con
tinued
)
Pegano-Salsoletea in Italy 7
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Character species: Salsola agrigentina, endemic to
central and southern Sicily, belonging to the cycle
of Salsola vermiculata (Botschantzev 1975).
Structure and ecology: Nanophanerophytic vegetation
colonizing steep heavily eroded clayish badlands
within the thermo-Mediterranean dry bioclimatic
belt. It tolerates high concentrations of chlorides, as
testified by the occurrence of salt crystals on the
ground surface during the summer months. This
vegetation is relatively dense with Salsola agrigentina,
the dominant species, growing together with Salsola
oppositifolia, Suaeda vera and Atriplex halimus. On less
sloping sites, this vegetation comes into contact with
the perennial dry grasslands of Lavatero agrigentinae-
Lygeetum sparti Brullo 1985.
Distribution: Central and southern Sicily, both inland
and near the coast (Figure 4 online Appendix).
Limonio catanzaroi-Salsoletum oppositifoliae
Brullo, Guglielmo & Pavone 1985, nom. inver.
propos. – Table I (7–8); Table IIA
Syn.: Salsolo oppositifoliae-Limonietum catanzaroi
Brullo, Guglielmo & Pavone 1985.
Holotypus: rel.8, tab.1, Brullo et al. (1985).
Character species: Limonium catanzaroi, endemic to
few scattered sites of western Sicily, and Hel-
minthotheca aculeata, having a Sicilian-Maghrebid
distribution.
Structure and ecology: The physiognomy of this
association is given by the dominance of Salsola
oppositifolia and Limonium catanzaroi, growing on
salty lithosoils deriving from the erosion of marls or
marly clays interposed to sandstone layers. It occurs
within the thermo-Mediterranean dry bioclimatic
belt. On more mature soils, as the salinity decreases,
this vegetation is replaced by the maquis of the
Pistacio-Chamaeropetum humilis Brullo e Marceno
1985 or by perennial dry grasslands of the Astragalo
huetii-Ampelodesmetum mauritanici Minissale 1995.
According to the Art 42 of the ICPN (Weber et al.
2000), it is here proposed to modify the name of the
association, since Salsola oppositifolia is the species
physiognomically dominant.
Distribution: South-western Sicily, from Ribera to
Capo Bianco (Figure 5 online Appendix).
Salsolo oppositifoliae-Suaedetum pelagicae
Bartolo, Brullo, Minissale & Spampinato
1990 – Table I (13)
Holotypus: rel.4, tab.11, Bartolo et al. (1990)
Character species: Suaeda pelagica, endemic to Lam-
pedusa and belonging to the cycle of S. palestina Eig
& Zohary (Bartolo et al. 1988).
Structure and ecology: Nano-phanaerophytic vegeta-
tion colonizing steep south-facing marly outcrops
and the detritus accumulating at their base, within
the infra-Mediterranean dry bioclimatic belt. TheTab
leII
.(C
onti
nued
).
Rel
.1
-2,
Cap
oB
ian
co(S
icu
lian
a),
04
.06
.20
09
;3
-4,
Mo
nta
gn
aM
ago
ne
(Rib
era)
,0
5.0
6.2
00
9;
5-9
,C
.da
An
cid
ra(R
iber
a),
05.0
6.2
00
9;
Rel
.1
0,
San
t’A
ngel
oM
uxar
o(A
gri
gen
to),
13
.05
.20
09
;1
1-1
5,
ibid
.,0
5.0
6.2
00
9;
16
-17
,C
.da
Val
angh
e(A
dra
no
),2
5.0
5.2
00
9;
18-1
9,
ibid
.,2
6.0
3.2
00
9;
Rel
.20
-23
,M
acca
lub
ed
iA
rago
na,
13
.05
.20
09.
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Tab
leII
I.S
uaed
over
ae-
Lim
onia
stre
tum
mon
opet
ali
(A),
Atr
iplici
halim
i-H
alim
ionie
tum
por
tula
coid
is(B
).
(con
tinued
)
Pegano-Salsoletea in Italy 9
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climatic stress is further amplified by the intense
direct sunlight, by the frequent coastal winds and salt
spray. Near the sea, this association gets in contact
with the Limonietum lopadusani Bartolo et al. 1990,
while on vertical cliffs, it is replaced by the
Chiliadenetum lopadusani Bartolo et al. 1990.
Distribution: Island of Lampedusa, along the south-
western coast (Figure 2 online Appendix).
Suaedo verae-Limoniastretum monopetali
Bartolo, Brullo, Minissale & Spampinato
1990 – Table I (14–15); Table IIIA
Holotypus: rel.2, tab.12, Bartolo et al. (1990)
Character species: Limoniastrum monopetalum, a
South-Mediterranean species normally found in
coastal salt marshes, as a member of the vegetation
ascribed to the class Sarcocornietea fruticosae. In the
context of Pegano-Salsoletea, it can be regarded as a
differential species.
Structure and ecology: Association dominated by the
big prostrate shrubs of Limoniastrum monopetalum
mixed with succulent Chenopodiaceae like Suaeda
vera, Salsola Oppositifolia, and Atriplex halimus. It
colonizes loamy sediments resulting from the weath-
ering of marly clays, marls, and carbonatic sand-
stones. Even if linked to thermo- and infra-
Mediterranean dry bioclimatic belt, it benefits from
a slight edaphic humidity provided by the shallow
water-table. It is typically found on flat or gently
sloping coastal sites, exposed to southern winds and
to the sea-spray. In more sheltered and elevated
places, it is replaced by the maquis of Oleo-Ceratonion
and Periplocion angustifoliae Rivas-Martinez 1975.
Distribution: This association was first described for
the Island of Lampedusa by Bartolo et al. (1990).
Recently, it has been recorded, as well, in some
localities along the southern and western Sicilian
coast (Figure 3 online Appendix).
Halimiono portulacoidis-Salsoletum oppositi-
foliae Brullo, Guarino & Ronsisvalle 2000 –
Table I (16–17); Table IVC
Holotypus: rel.5, tab.9, Brullo et al. (2000).
Character species: Halimione portulacoides.
Structure and ecology: Association colonizing hol-
lowed sites and basins created by the weathering on
horizontal layers of limestone and carbonatic sand-
stones, where the combined action of sea-storms and
coastal winds accumulates loamy sediments and
organic matter. This explains the abundance of the
Halimione portulacoides, a species normally found in
coastal salt marshes belonging to the class Sarcocor-
nietea fruticosae Br.-Bl. & R. Tx. ex A. & O. Bolos
1950, and particularly in the Halimiono portulacoidis-
Suaedetum verae. In this context, instead, Halimione
portulacoides grows together with many characteristics
of the class Pegano-Salsoletea, such as Suaeda vera,Tab
leII
I.(C
onti
nued
).
Rel
.1-3
,P
ort
oE
mp
edo
cle,
nea
rV
illa
Ro
man
a,0
4.0
6.2
00
9;4
-8,S
cala
dei
Tu
rch
i,R
ealm
on
te,0
4.0
6.2
00
9;R
el.9
,L
ido
Ro
ssel
lo,R
ealm
on
te,0
4.0
6.2
00
9;1
0-1
1,P
un
taB
racc
etto
(Rag
usa
),(B
arto
loet
al.
19
82
,ta
b.2
2,
rel.
14
-15
);R
el.
12
-17
,R
ock
yco
ast
of
Cat
ania
,2
5.0
4.1
99
5;
18
-19
,ib
id.,
29
.03
.199
4.
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Salsola Oppositifolia, and Lycium intricatum, within
the thermo-Mediterranean dry bioclimatic belt.
Distribution: Southern Sicily (Figure 5 online Appen-
dix), near Gela (Brullo et al. 2000; Guarino et al.
2008), and Ribera (Agrigento).
Atriplici halimi-Halimionietum portulacoidis
ass. nova – Table I (20); Table IIIB
Holotypus: rel.13, tab.3, hoc loco.
Character species: Halimione portulacoides.
Structure and ecology: Nano-phanerophytic vegetation
colonizing flat or gently sloping volcanic outcrops
along the sea, influenced by the sea-spray and by the
organic waste accumulated during the sea-storms. It
is linked to thermo-Mediterranean subhumid biocli-
matic belt, in catenal contact with the Crithmo-
Limonietea vegetation. Like the Halimiono portulacoi-
dis-Salsoletum oppositifoliae, this association is char-
acterized by the dominance of Halimione
portulacoides, but it clearly differs from the latter for
the habitat conditions and for the absence of Salsola
oppositifolia.
Distribution: North-eastern Sicily, along the volcanic
coast of Catania (Figure 5 online Appendix).
Table IV. Asparago albi-Salsoletum oppositifoliae (A), Capparido siculae-Salsoletum oppositifoliae (B), Halimiono portulacoidis-Salsoletum
oppositilofilae (C)
Rel. 1-2, Sicily, Porto Empedocle, 04.06.2009; 3-4, Sicily, Porto Empedocle near Villa Romana, 04.06.2009; Rel. 5, Sicily, C.da Ancidra
(Ribera), 05.06.2009; 6, Calabria, rocky coast near Capo dell’Armi, 04.05.1987 (Brullo et al. 2001, tab.48, rel.2); Rel. 7, Calabria, Saline
Ioniche, 06.06.1996 (Brullo et al. 2001, tab.48, rel.3); 8-9, Sicily, Porto Empedocle, 04.06.2009; Rel. 10-11, Sicily, Porto Empedocle near
Villa Romana, 04.06.2009; 12, C.da Ancidra (Ribera), 05.06.2009.
Pegano-Salsoletea in Italy 11
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Tab
leV
.A
spara
gost
ipula
ris-
Sals
olet
um
ver
mic
ula
tae
(A),
Lyc
ioin
tric
ati
-Sals
olet
um
oppos
itifol
iae
(B),
Med
icagi
ni
arb
orea
e-S
als
olet
um
oppos
itifol
iae
(C).
(con
tinued
)
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Capparido siculae-Salsoletum oppositifoliae
ass. nova – Table I (1–3); Table IVB
Syn.: Suaedo verae-Salsoletum oppositifoliae auct. sic.
non Rivas Goday & Rigual 1958
Holotypus: rel.10, tab.5, hoc loco.
Character species: Capparis sicula.
Structure and ecology: Nano-phanerophytic vegetation
colonizing heavily eroded clayey badlands, both
inland and in coastal sites. It can tolerate extreme
edaphic conditions, where the concentration of
chlorides is so high that often, during the summer
drought, they become visible as salt crystals on the
ground surface. This association is a permanent
edaphophilous community linked to the lower
thermo-Mediterranean lower dry bioclimatic belt,
within the climatic plant communities of the Oleo-
Ceratonion.
Distribution: Western flank of Mt. Etna, near Adrano,
and southern Sicily, from Porto Empedocle (Agri-
gento) to Capo Passero (Fig.4 online Appendix). By
previous authors (Bartolo et al. 1982; Brullo &
Siracusa 2000), this vegetation was ascribed to the
Suaedo verae-Salsoletum oppositifoliae (O. Bolos 1957)
Rivas Goday and Rigual 1958, association described
for the Province of Alicante, in southern Spain.
Despite the relatively similar physiognomy, chiefly
given by the dominance of Salsola oppositifolia and
Suaeda vera, the Spanish vegetation is well char-
acterized by a pool of W-Mediterranean species
missing from Sicily. Therefore, a new association is
proposed here.
Limonio calcarae-Suaedetum verae ass. nova –
Table I (9); Table IIB
Holotypus: rel.10, tab. 2, hoc loco.
Character species. Limonium calcarae, endemic to
central Sicily.
Structure and ecology: Halo-nitrophilous association
colonizing clayish badlands belonging to the Sicilian
evaporitic series. It dwells heavily eroded gullies and
steep slopes within the thermo-Mediterranean dry
bioclimatic belt. The structure is given by scattered
tufts of Limonium calcarae, by far the most abundant
species in this context, mixed with sparse individuals
of Suaeda vera and Moricandia arvensis.
Distribution: Central Sicily (Figure 5 online Appen-
dix), far from the sea (Caltanissetta and Agrigento
provinces).
Thapsio pelagicae-Salsoletum oppositifoliae
ass. nova – Table I(5)
Syn.: Suaedo verae-Salsoletum oppositifoliae Bartolo
et al. 1990 non Rivas Goday & Rigual 1958
Holotypus: rel.5, tab.10, Bartolo et al. (1990), hoc
loco.
Character species: Thapsia pelagica, endemic to Lam-
pedusa (formerly attributed to T. garganica byTab
leV
.(C
onti
nued
).
Rel
1-4
,6
-8C
agliar
i,C
ala
Mo
sca,
26
.4.1
989
;5
,9
,1
0,
ibid
.,2
4.0
6.2
00
9;
11
-12
,S
cala
dei
Tu
rch
i,R
ealm
on
te,
04
.06.2
00
9;
13
,ro
cky
coas
tn
ear
Mt.
Polisc
ia(L
icat
a),
28
.05
.200
9;
Rel
.1
4-1
9,
Mt.
Lu
ngo
(Gel
a),
16.0
8.2
00
5;
20
-22
,C
.da
Mo
llar
ella
(Lic
ata)
,0
5.0
5.2
00
7;
23
,ib
id.,
28
.05
.20
09
;2
4,
rock
yco
ast
nea
rM
t.P
olisc
ia(L
icat
a),
28
.05
.20
09
.
Pegano-Salsoletea in Italy 13
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Bartolo et al. 1990), and Reichardia tingitana, a
South-Mediterranean species ranging up to the
Pelagic Islands.
Structure and ecology: Nano-phanerophytic vegetation
linked to marly substrata dunged by the seabirds,
within the infra-Mediterranean semiarid bioclimatic
belt. Its physiognomic characterization is given by a
dense layer of Salsola oppositifolia, Suaeda vera, and
Atriplex halimus that, towards the inland, are pro-
gressively replaced by the climatophilous vegetation
of the Periplocion angustifoliae.
Distribution: Island of Lampedusa (Figure 4 online
Appendix).
Note: This vegetation was previously ascribed by
Bartolo et al. (1990) to the Suaedo verae-Salsoletum
oppositifoliae (O. Bolos 1957) Rivas Goday and
Rigual 1958, association described for the Province
of Alicante, in southern Spain. Despite the relatively
similar physiognomy given by the dominance of
Salsola oppositifolia and Suaeda vera, the Spanish
vegetation is well characterized by a pool of W-
Mediterranean species, lacking from Lampedusa.
The Lampedusa populations ascribed by Bartolo
et al. (1990) to Thapsia garganica have been recently
described by Brullo et al. (2009) as new species,
namely T. pelagica.
Asparago albi-Salsoletum oppositifoliae ass.
nova – Table I (4), Table IVA
Holotypus: rel.4, tab.4, hoc loco.
Table VI. Atriplici halimi-Artemisietum arborescentis subass. salsoletosum oppositifoliae (A), subass. halimionetosum portulacoidis (B).
Rel. 1-5, Siculiana Marina, 15.05.1993; 6-8, Capo Bianco (Siculiana), 04.06.2009; Rel. 9-11, Foce del Simeto (Brullo et al. 1988, tab.19);
12-13, Cagliari, Stagno di Santa Gilla, 25.06.2009.
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Tab
leV
II.
Lim
onio
opti
mae-
Sals
olet
um
oppos
itifol
iae
(A),
Lyc
ioeu
ropaei
-Art
emis
ietu
marb
ores
centi
s(B
),C
oron
illo
vale
nti
nae-
Art
emis
ietu
marb
ores
centi
s(C
)
(con
tinued
)
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Character species: Asparagus albus.
Structure and ecology: Nano-phanaerophytic vegeta-
tion chiefly colonizing coastal stands, or rarely inner
stand, characterized marly soils. It usually dwells
very steep slopes, mainly represented by cliffs or
rocky outcrops. It is floristically differentiated by
Salsola oppositifolia, usually reaching high cover
values, and by the occurrence of Asparagus albus.
This association is a permanent edaphophilous
community linked to the lower thermo-Mediterra-
nean lower dry bioclimatic belt, within the climatic
plant communities of the Oleo-Ceratonion.
Distribution: Southern Sicily, near Porto Empedocle
and Ribera (Agrigento), and southern Calabria, near
Reggio Calabria, along the rocky coast between Capo
dell’Armi and Saline Ioniche (Figure 4 online
Appendix).
Asparago stipularis-Salsoletum vermiculatae
ass. nova – Table I (23–27); Table VA
Syn.: Salsolo vermiculatae-Atriplicetum halimi Biondi
& Mossa 1992 non Rivas-Martınez et al. 1991.
Holotypus: rel.8, tab.15, Biondi & Mossa (1992), hoc
loco.
Character species: Salsola vermiculata, a W-Mediterra-
nean species ranging up to southern Sardinia, and
Limonium dubium, endemic to Sicily, Sardinia and
Corsica.
Structure and ecology: Vegetation colonizing relatively
steep marly slopes exposed to the salt-spray and
dunged by the seabirds, within the thermo-Mediter-
ranean dry bioclimatic belt. Its ecological niche
stretches between the vegetation of Crithmo-Limonie-
tea Br.-Bl. in Br.-Bl. et al. 1952 and the thermo-
xerophilous maquis ascribed to the Euphorbio den-
droidis-Anagyretum foetidae Biondi and Mossa 1992
(Oleo-Ceratonion Br.-Bl. ex Guinochet and Droui-
neau 1944). The structure is given by Salsola
vermiculata, that together with Atriplex halimus,
Asparagus stipularis, and Suaeda vera, forms a tangled
scrubland, densely covering the ground.
We agree with Biondi and Mossa (1992) which,
besides to the typical aspect without Artemisia
arborescens, describe the subass. artemisietosum arbor-
escentis (Biondi & Mossa 1992) comb. nov. which is
characterized by the occurrence of A. arborescens,
often with high cover values. From the ecological
viewpoint, the subass. typicum is exclusively linked to
primary stands, as testified by the lacking of A.
arborescens, while the subass. artemisietosum arbores-
centis grows on disturbed habitats, with an higher
concentration of nitrates.
Distribution: Southern Sardinia, on promontories and
coastal capes eastward of Cagliari (Figure 3 online
Appendix).
Note: This vegetation was already sampled by Biondi
and Mossa (1992) and ascribed to the SalsoloTab
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16 S. Brullo et al.
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vermiculatae-Atriplicetum halimi (Br.-Bl. & Bolos
1958) Rivas-Martınez et al. 1991. Despite the similar
physiognomy, the Sardininan stands are quite
different from those of the Iberian Peninsula for
ecological and floristic reasons: the Spanish associa-
tion grows inland, within the meso-Mediterranean
dry bioclimatic belt and belongs to the Salsolo-
Peganion harmalae, an alliance that is not represented
in Italy.
Artemision arborescentis
Atriplici halimi-Artemisietum arborescentis
Biondi 1988 – Table I (34–39); Table VI
Holotypus: rel.4, tab.1, Biondi (1988).
Character species: Atriplex halimus and Artemisia
arborescens.
Structure and ecology: Synanthropic shrublands domi-
nated by the two nominal species of the associations,
sometimes growing together with Suaeda vera. This
very poor floristic settlement is often found in coastal
sites, on marly and conglomeratic substrata, some-
times used as dumping places for urban waste and
rubbles (Biondi 1988). It can be considered as a
permanent edaphic community whose evolution is
contrasted by the human disturbance, combined to
the action of natural ‘‘hecklers’’ like coastal winds,
seabirds, salt-spray, etc. It is found within the
thermo-Mediterranean dry to subhumid bioclimatic
belt, often in contact with the Oleo-Ceratonion-
vegetation. In addition to the typical aspect and to
the var. with Suaeda vera, already described by
Biondi (1988), the following two sub-associations
can be recognized: (a) subass. salsoletosum oppositi-
foliae Costanzo et al. 2005 (holotypus: rel.3, tab.14,
Costanzo et al. 2005), dwelling steep marly cliffs
exposed to the sea; (b) subass. halimionetosum
portulacoidis subass. nova (holotypus: rel.9, tab.6,
hoc loco), which is found on loamy sediments
surrounding coastal salt marshes.
Distribution: Southern Italy (Tremiti Islands, Garga-
no, and northern Calabria), Sardinia and Sicily
(Figure 3 online Appendix).
Limonio optimae-Salsoletum oppositifoliae
ass. nova – Table I (30); Table VIIA
Holotypus: rel.5, tab.7, hoc loco.
Character species: Limonium optimae, endemic to
central Sicily (Raimondo 1993, Falci 2000).
Structure and ecology: Association colonizing marly
limestone and warps along the Salso river and its
tributaries, known since ancient times for the salinity
of their water, resulting from the leaching of
evaporitic layers. This vegetation is found between
300 and 400 m a.s.l., within the thermo-Mediterra-
nean subhumid bioclimatic belt. Typically, it has
quite an open structure, given by sparse shrubs of
Salsola oppositifolia and Artemisia arborescens, mixing
together with denser clumps of Limonium optimae.
Distribution: Central Sicily, near Caltanissetta (Figure
5 online Appendix).
Coronillo valentinae-Artemisietum arbores-
centis ass. nova – Table I (31); Table VIIC
Holotypus: rel.21, tab.7, hoc loco.
Character species: Coronilla valentina, Ruta chalepensis.
Structure and ecology: Association colonizing steep,
heavily eroded, marly slopes, within the thermo-
Mediterranean subhumid bioclimatic belt, between
400 and 500 m a.s.l. The structure is given by sparse
shrubs of Artemisia arborescens, Salsola oppositifolia
and Anagyris foetida, mingled with the smaller bushes
of Coronilla valentina and Ruta chalapensis. This
vegetation is ecologically quite specialized, linked to
relatively fine-grained debris frequently disturbed by
the deposition of new eroded material. In more
stable contexts, it tends to be replaced by the Oleo-
Ceratonion vegetation.
Distribution: Central and northern Sicily (Figure 3
online Appendix).
Lycio europaei-Artemisietum arborescentis
ass. nova –Table I (28–29); Table VIIB
Holotypus: rel.15, tab.7, hoc loco.
Character species: Lycium europaeum.
Structure and ecology: Halo-nitrophilous association
colonizing clayey substrata next to urban and rural
settlements, within the meso-Mediterranean dry to
subhumid bioclimatic belt, at 500–700 m a.s.l. It has
some more mesophilous requirements in comparison
with the above-mentioned associations and often
achieves high cover values. The structure is given by
the big bushes of Artemisia arborescens and Atriplex
halimus, tangled with Lycium europaeum: a synan-
thropic species frequently found in secondary habi-
tats like road-sides, pathways, and embankments.
Distribution: Central and northern Sicily (Figure 2
online Appendix).
Notes: This plant community was formerly described
by Costanzo et al. (2005) as ‘‘aggr. Lyceum euro-
paeum’’ from central Sicily.
Lycio intricati-Salsoletum oppositifoliae ass.
nova – Table I (32–33); Table VB
Syn.: Suaedo verae-Salsoletum oppositifoliae Brullo
et al. 2000 non Rivas Goday & Rigual 1958
Holotypus: rel.14, tab.5, hoc loco.
Character species: Lycium intricatum, a SW-Mediter-
ranean thermo-xerophilous species.
Structure and ecology: Halo-nitrophilous association
growing on steep marly to clayish badlands, often
with southern exposures. Its optimum is within the
thermo-Mediterranean dry bioclimatic belt, near the
coast. The structure is rather dense and intricate,
Pegano-Salsoletea in Italy 17
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with big bushes of Salsola oppositifolia, Suaeda vera,
Asparagus stipularis and Lycium intricatum, growing
together with Artemisia arborescens, which tends to
prevail in the most disturbed sites. On gently sloping
sites, this vegetation is progressively replaced by dry
grasslands with Lygeum spartum.
Distribution: Coasts of southern Sicily, from Realm-
onte (Agrigento) to Gela (Fig.2 online Appendix).
Notes: In a previous contribution (Brullo et al. 2000),
this vegetation was ascribed to the Suaedo verae-
Salsoletum oppositifoliae (O. Bolos 1957) Rivas Goday
and Rigual 1958, association described for the
Province of Alicante, in southern Spain. Depite the
relatively similar physiognomy, due to substantial
floristic and ecologic differences, it is here preferred
to describe a new association.
Medicagini arboreae-Salsoletum oppositifoliae
ass. nova – Table I (40); Table VC
Holotypus: rel.19, tab.5, hoc loco.
Character species: Medicago arborea, a NW-Mediter-
ranean species which is very rare in Sicily.
Structure and ecology: This association is found on
limestone and carbonatic sandstones disturbed by
the sea-spray and dunged by the seabirds. It can be
considered an edaphic vicariant of the Lycio intricati-
Salsoletum oppositifoliae. Also in this case, the
structure is very dense, with Medicago arborea as
dominant species, and Salsola oppositifolia, Artemisia
arborescens, Suaeda vera and Atriplex halimus as
subordinates.
Distribution: Southern coast of Sicily, near Licata
(Figure 2 online Appendix).
Discussion
The phytosociological surveys carried out on this
very peculiar vegetation allowed to examine not only
the ecological differences of the investigated plant
communities, but also to evaluate the anthropogenic
effects on their structure and floristic composition.
Furthermore, thanks to synecologic approach, it was
possible to analyse the transformation of the plant
landscape in Sicily (Biondi 2011; Pott 2011; Blasi &
Frondoni 2011). All the associations occurring in
Italy, and examined in this study, are structurally,
floristically, and ecologically well differentiated, as
also confirmed by the numerical analysis. Figure 1
shows two clusters clearly separated, corresponding
to the Salsolo-Suedion mollis (cluster A) and to the
Artemision arborescentis (cluster B). Cluster B is quite
homogenous, since it includes plant communities
floristically poor, and chiefly playing a secondary
role, while three subclusters can be recognized within
cluster A. In particular, subcluster A1 groups the
associations characterized by the occurrence and
dominance of Salsola oppositifolia, thus suggesting
that these phytocoenoses are strictly linked to
particularly dry environments. The associations fall-
ing in subcluster A1 are found on stands of central-
southern Sicily, Lampedusa, as well as the south-
ernmost point of Calabria (i.e., Capo dell’Armi);
from the bioclimatic viewpoint, the thermotypes of
these growing sites are ranging from the infra- to the
thermo-Mediterranean dry. Subcluster A2 gathers
the plant communities dominated by Suaeda vera,
plant having its optimal growth conditions on soils
with a higher water capacity. The ecological beha-
viour of S. vera is also testified by the fact that it is the
key-species of the Suaedion verae (Sarcocornietea
fruticosae), alliance including subhalo-nitrophilous
phytocoenoses colonizing the edges of salt-marshes.
The associations of this subcluster are known only
for Sicily and Lampedusa. Finally, subcluster A3
groups the associations exclusively occurring in
Sardinia and central-southern Italy. Such plant
communities are characterized by the dominance of
Atriplex halimus, usually growing together with
Suaeda vera, while Salsola oppositifolia is definitely
missing. This is probably due to the bioclimatic
features of the dwelled sites, which are usually less
arid than the previous ones, since they fall into meso-
Mediterranean thermotype, or more rarely into the
thermo-Mediterranean one.
Conclusions
The class Pegano-Salsoletea is represented in Italy by
several well-differentiated phytosociological units,
that deserve more attention than what is normally
attributed to such vegetation. Apart from the
botanical interest, it is worth to remind that all the
surveyed associations are to be ascribed to the habitat
‘‘1430 – Halo-nitrophilous shrubs (Pegano-Salsole-
tea)’’, mentioned in the Supplement 1 of the
European Directive 43/92.
Moreover, most of the sites colonized by this
vegetation have an outstanding stratigraphic and geo-
morphological interest. Just to make an example, the
Global Stratotype Section and Point (GSSP) iden-
tifies in Mt. San Nicola, near Gela (S Sicily), the
stratigraphic reference for the Quaternary/Pleisto-
cene lower boundary. Similarly, the badlands of
Vrica, in southern Calabria, identify the early stage of
Pleistocene, known as ‘‘Calabrian Stage’’ (Gibbard
et al. 2010).
It can be noted, as well, that most of the species
characterizing the Pegano-Salsoletea vegetation have
an outstanding bio-geographical interest, to track
back the Plio-Pleistocenic phases of the Mediterra-
nean Basin. In fact, they originated from Irano-
Turanian and Saharo-Arabian elements and spread
around the Mediterranean basin during the Messi-
nian Age, of which they can be considered as ‘‘living
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traces’’ (Guarino et al. 2006). Quite relevant is the
occurrence within the surveyed plant communities of
several rare endemic species, such as Thapsia
pelagica, Suaeda kocheri, S. pelagica, Herniaria empe-
docleana, Limonium opulentum, L. dubium, L. catan-
zaroi, L. calcarae, L. optimae, Salsola agrigentina, and
Polygonum tenoreanum. The dwelled sites are chiefly
represented by ecologically very specialized habitats,
whose geographical and overall ecological isolation
strongly favoured the speciation processes of these
halophytes.
Unfortunately, many of the stands colonized by
the Pegano-Salsoletea vegetation are prone to the
recent demographic changes that invested the
Mediterranean people in last decades. The so-called
‘‘coastalization’’ (Cori 1999; Ivanov et al. 2008)
associated to a deregulated urban development has
determined quite an high human density in the close
proximity of most of the Pegano-Salsoletea stands in
Italy.
The proximity of these sites to areas of rapid
population growth and urban development is a
source of increasing concern, also because the
naturalistic and scientific value of such vegetation is
ignored by the most, the public access to sensitive
habitats/biotopes is totally uncontrolled. The resi-
dential development appropriated (often embezzled)
a great deal of the areas where the Pegano-Salsoletea
vegetation was growing relatively undisturbed until
recent years. In this sense, the case of the Limonio
opulenti-Salsoletum oppositifoliae is emblematic: it is
estimated that in the last three decades, 80% of the
habitat went lost, even if some floristic reminiscences
of the former vegetation can be sadly found in
dumpsites, parking places, garaging, and towing
services, along the walls of the power station of
Porto Empedocle. The same holds true for the many
‘‘badlands’’ that, due to the waterproof properties of
the clay, have been transformed in landfills for the
urban solid waste. This is the case, for instance, of a
relevant percentage of the habitat formerly occupied
by the Capparido siculae-Salsoletum oppositifoliae, near
the town of Adrano.
Impacts from human activities are resulting in a
severe habitat loss and declining environmental
quality for the Pegano-Salsoletea vegetation. Roads,
dust, pollution, industrial activities, and uncon-
trolled off-road access are limiting factors in the
population dynamics of the most sensitive species.
Moreover, the overall disturbance will probably
make introduced feral species more competitive,
opening the way to multiple problems, such as the
potential to out-compete native species, habitat
alteration, and alteration of the trophic levels. Up
to now, the only non-native species having a
significant presence in the Pegano-Salsoletea vegeta-
tion of Italy are as follows: Boerhavia repens, Ricinus
communis, Solanum sodomaeum, and Nicotiana glauca,
but if the human disturbance increases, it cannot be
excluded that, in the next future, new exotic species
become stronger competitors of the native ones.
An integrated management plan is urgently
needed. It should include a zoning framework to
protect key habitats and species from detrimental
human activities as well as to allow (and suggest)
appropriate sustainable uses. It is to be hoped that, in
the next future, men and resources will be found in
order to make more acceptable the conflict between
human activities and conservation needs for the still
surviving patches of Pegano-Salsoletea vegetation.
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Appendix 1. Syntaxonomical scheme of the
class Pegano-Salsoletea in the Mediterranean
Basin (up to alliance level)
PEGANO HARMALAE-SALSOLETEA VERMI-
CULATAE Br.-Bl & O.Bolos 1958
Syn.: Ruderali-Manihotetea utilissimae Leonard in
Taton 1949 corr. O.Bolos 1988
SALSOLO VERMICULATAE-PEGANETALIA HARMALAE Br.-
Bl. & O.Bolos 1954
Syn.: Onopordo-Salsoletalia Rivas Goday & Rivas-
Martınez 1963
Atriplicetalia glaucae Rivas Goday & Rivas-Martınez
1963
Ipomoeetalia purpureae Oberd. ex O.Bolos 1988
Salsolo vermiculatae-Peganion harmalae Br.-Bl. &
O.Bolos 1954
Syn.: Salsolo-Fagonion creticae Rivas Goday &
Rigual Magallon 1958
Salsolo-Carthamion arborescentis Rivas Goday &
Rivas-Martınez 1963
Salsolo-Artemision Folch 1981 n.n.
Salsolo oppositifoliae-Suaedion mollis Rigual 1972
Syn.: Suaedenion fruticoso-verae Peinado, Martınez
Parrasa & Bartolome 1986
Suaedenion verae Peinado, Martınez Parrasa &
Bartolome 1986 corr. Peinado et al. 1989
Artemision arborescentis Gehu & Biondi in Gehu
et al. 1986
Hammado articulatae-Atriplicion glaucae Rivas
Goday & Rivas-Martınez ex Rigual 1972 corr.
Alcaraz et al. 1991
Syn.: Haloxilo-Atriplicion Rivas Goday & Rivas-
Martınez 1963 n.n.
Haloxilo-Atriplicion Rivas Goday & Rivas-Martınez
ex Rigual 1972
Atriplicion glaucae Folch 1981 n.n.
Atriplicenion glaucae Peinado, Martınez Parrasa &
Bartolome 1986
Medicagini citrinae-Lavaterion arboreae O.Bolos,
Folch & Vigo in O. Bolos & Vigo 1984
Lyceo europaei-Ipomoeion purpureae O.Bolos 1988
HAMMADO SCOPARIAE-ANABASIETALIA ARTICULATAE
Brullo & Furnari 1996
Limonion tubiflori Brullo & Furnari 1996
Thymelaeion hirsutae Tadros & Atta 1958
CHENOLEOIDETALIA TOMENTOSAE Sunding 1972
Chenoleoideion tomentosae Sunding 1972
HELICHRYSO-SANTOLINETALIA SQUARROSAE Peinado &
Martinez Parras 1984
Artemisio glutinosae-Santolinion rosmarinifoliae
Costa 1975
Santolinion pectinato-canescentis Peinado & Marti-
nez Parras 1984
FORSSKAOLEO ANGUSTIFOLIAE-RUMICETALIA LUNARIAE
Rivas-Martınez et al. 1993
Launaeo arborescentis-Schizogynion sericeae Rivas-
Martınez et al. 1993
Artemision thusculae-Rumicion lunariae Rivas-Mar-
tınez et al. 1993
Argyranthemo succulenti-Calendulion maderensis
Capelo et al. 2000
NICOTIANO GLAUCAE-RICINETALIA COMMUNIS Rivas-
Martınez, Fernandez-Gonzalez & Loidi 1999
Nicotiano glaucae-Ricinion communis Rivas-Martı-
nez, Fernandez-Gonzalez & Loidi 1999
Appendix 2. Syntaxonomical scheme of the
surveyed associations
PEGANO-SALSOLETEA Br.-Bl & O.Bolos 1958
Salsolo vermiculatae-Peganetalia harmalae Br.-Bl. &
O. Bolos 1954
Salsolo oppositifoliae-Suaedion MOLLIS Rigual
1972
Asparago albi-Salsoletum oppositifoliae ass. nova
Atriplici halimi-Halimionietum portulacoidis ass.
nova
Atriplici halimi-Polygonetum tenoreani Biondi, Bal-
lelli & Taffetani 1992
Camphorosmo monspeliacae-Atriplicetum halimi
Biondi, Ballelli & Taffetani 1992
Capparido siculae-Salsoletum oppositifoliae ass.
nova
Halimiono portulacoidis-Salsoletum oppositifoliae
Brullo, Guarino & Ronsisvalle 2000
Limonio calcarae-Suaedetum verae ass. nova
Salsoletum agrigentinae Brullo, Guglielmo & Pavone
1985
Limonio catanzaroi Salsoletum oppositifoliae Brullo,
Guglielmo & Pavone 1985 nom. invers. propos.
Limonio opulenti-Salsoletum oppositifoliae Brullo,
Grillo & Scalia 1980 nom. invers. propos.
Salsolo oppositifoliae-Suaedetum pelagicae Bartolo
et al. 1990
Suaedo verae-Atriplicetum halimi Biondi 1988
Suaedo verae-Limoniastretum monopetali Bartolo
et al. 1990
Pegano-Salsoletea in Italy 21
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Thapsio pelagicae-Salsoletum oppositifoliae ass. nova
Asparago stipularis-Salsoletum vermiculatae ass. nova
a) typicum
b) artemisietosum arborescentis (Biondi & Mossa
1992) comb. nova
ARTEMISION ARBORESCENTIS Gehu & Biondi in Gehu
et al. 1986
Atriplici halimi-Artemisietum arborescentis
Biondi 1988
a) salsoletosum oppositifoliae subass. nova
b) halimionetosum portulacoidis subass. nova
Coronillo valentinae-Artemisietum arborescentis ass.
nova
Limonio optimae-Salsoletum oppositifoliae ass. nova
Lycio europaei-Artemisietum arborescentis ass. nova
Lycio intricati-Salsoletum oppositifoliae ass. nova
Medicagini arboreae-Salsoletum oppositifoliae ass.
nova
Figure 2. Distribution map of some Pegano-Salsoletea associations occurring in Italy, Sardinia, and Sicily.
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Figure 3. Distribution map of some Pegano-Salsoletea associations occurring in Italy, and Sicily.
Pegano-Salsoletea in Italy 23
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Figure 5. Distribution map of some Pegano-Salsoletea associations occurring in Sicily.
Figure 4. Distribution map of some Pegano-Salsoletea associations occurring in Calabria and Sicily.
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Tab
leS
1.
Asp
arag
oal
bi-
Sal
sole
tum
op
po
siti
foliae
(rel
.1-3
),C
app
arid
osi
cula
e-S
also
letu
mo
pp
osi
tifo
liae
(rel
.4-6
),H
alim
ion
op
ort
ula
coid
is-S
also
letu
mo
pp
osi
tifo
liae
(rel
.7-9
),L
imo
nio
cata
nza
roi-
Sal
sole
tum
op
po
siti
foliae
,L
imo
nio
calc
arae
-Su
aed
etu
mve
rae
(rel
.13
-15),
Sal
sole
tum
agri
gen
tin
ae(r
el.1
6-1
8),
Su
aed
ove
rae-
Lim
on
iast
retu
mm
on
op
etal
i(r
el.1
9-2
1),
Atr
iplici
hal
imi-
Hal
imio
nie
tum
po
rtu
laco
idis
(rel
.22
-24
),C
amp
ho
rosm
om
on
spel
iaca
e-A
trip
lice
tum
hal
imi
(rel
.25
-27
),A
trip
lici
hal
imi-
Po
lygo
net
um
ten
ore
ani
(rel
.28
-30
),T
hap
sio
pel
agic
ae-S
also
letu
mo
pp
osi
tifo
liae
(rel
.31
-33
),
Sal
solo
op
posi
tifo
liae
-Su
aed
etu
mp
elag
icae
(rel
.34
-36
),S
uae
do
vera
e-A
trip
lice
tum
hal
imi
(rel
.37
-39
),L
imo
nio
op
ule
nti
-Sal
sole
tum
op
po
siti
foliae
(rel
.40
-42),
Atr
iplici
hal
imi-
Art
emis
ietu
m
arb
ore
scen
tis
typ
us
(rel
.43
-45
),A
trip
lici
hal
imi-
Art
emis
ietu
mar
bo
resc
enti
ssa
lso
leto
sum
op
po
siti
foliae
(rel
.46
-48
),A
trip
lici
hal
imi-
Art
emis
ietu
mar
bo
resc
enti
sh
alim
ion
eto
sum
po
rtu
laco
idis
(rel
.49
-
51
),L
imo
nio
op
tim
ae-S
also
letu
mo
pp
osi
tifo
liae
(rel
.52
-54
),L
ycio
euro
pae
i-A
rtem
isie
tum
arb
ore
scen
tis
(rel
.55
-57
),C
oro
nillo
vale
nti
nae
-Art
emis
ietu
mar
bo
resc
enti
s(r
el.5
8-6
0),
Asp
arag
ost
ipu
lari
s-
Sal
sole
tum
verm
icu
lata
ety
pic
um
(rel
.61
-63),
Asp
arag
ost
ipu
lari
s-S
also
letu
mve
rmic
ula
tae
sued
ieto
sum
(rel
.64
-66
),L
ycio
intr
icat
i-S
also
letu
mo
pp
osi
tifo
liae
(rel
.67
-69
),M
edic
agin
iar
bo
reae
-
Sal
sole
tum
op
po
siti
foliae
(rel
.7
0-7
2).
(con
tinued
)
Pegano-Salsoletea in Italy 25
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Tab
leS
1.
(Con
tinued
).
26 S. Brullo et al.
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