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Annual dune plant communities in the Southwest coast
of Europe
J. C. Cost a
a
, C. Net o
b
b
, M. Mart ins & M. Lousã
a
a
Cent ro de Bot ânica Aplicada à Agricult ura (CBBA), Inst it ut o Superior de Agronomia,
Technical Universit y of Lisbon (TULisbon), Tapada da Aj uda, 1349-017, Lisboa, Port ugal
b
Cent ro de Bot ânica Aplicada à Agricult ura (CBBA), Inst it ut e of Geography and Spat ial
Planning (IGOT), Universit y of Lisbon, Edif ício da Faculdade de Let ras, Alameda da
Universidade, 1600-214, Lisboa, Port ugal
Available online: 22 Aug 2011
To cite this article: J. C. Cost a, C. Net o, M. Mart ins & M. Lousã (2011): Annual dune plant communit ies in t he Sout hwest
coast of Europe, Plant Biosyst ems - An Int ernat ional Journal Dealing wit h all Aspect s of Plant Biology, 145: sup1, 91-104
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Plant Biosystems, Vol. 145, Supplement, September 2011, pp. 91–104
Annual dune plant communities in the Southwest coast of Europe
J. C. COSTA1, C. NETO2, M. MARTINS2, & M. LOUSÃ1
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1
Centro de Botânica Aplicada à Agricultura (CBBA), Instituto Superior de Agronomia, Technical University of Lisbon
(TULisbon), Tapada da Ajuda, 1349-017 Lisboa, Portugal and 2Centro de Botânica Aplicada à Agricultura (CBBA),
Institute of Geography and Spatial Planning (IGOT), University of Lisbon, Edifı́cio da Faculdade de Letras, Alameda da
Universidade, 1600-214 Lisboa, Portugal
Abstract
This study presents an updated sintaxonomic review of the annual communities from coastal dunes in the Southwest coast of
Europe, specifically in the Iberian Peninsula, including both types: directly influenced by the sea salt spray and not affected
by this influence. The floristic segregation of the different plant associations was obtained by statistical agglomerative
processes (UPGMA, Unweighted Pair Group Method with Arithmetic Mean) plus principal coordinate analysis (PCoA) and
discussed based on the interpretation of phytosociological tables. Three new communities in central western Portugal were
found and are described: Pseudorlayo minusculae-Polycarpietum alsinifoli, Cerastio diffusae-Vulpietum fontqueranae and
Omphalodo kunzinskyanae-Evacietum ramosissimae.
Keywords: Biogeography, coastal dunes, chorology, sintaxonomy, Iberian Peninsula
Introduction
A general characteristic of annual plant communities
is the fact that they colonize extreme biotopes, within
the possible limits for vegetal life (Biondi 2007)
where physical conditions hinder the development of
most perennials. Globally, the diversity of therophitic
communities is enormous, because these communities easily change their floristic combination along
ecological gradients such as climatic, soil, topographic and human disturbance-related gradients
(Guarino et al. 2005). After the major review from
Dı́ez-Garretas et al. (2003), to extensive dune areas,
especially in the North of Lisbon (Dividing Portuguese Sector, Sadensean-Dividing Portuguese Subprovince, Lusitan–Andalusian Coastal Province), the
available information has been revealed to be scarce.
Thus, new complete studies on the annual dune
communities in the west of the Iberian Peninsula
were needed. Such communities belong to the
alliances Linarion pedunculatae (Cutandietalia maritimae), Anthyllido hamosae-Malcolmion lacerae (Malcolmietalia) and Thero-Airion (Helianthemetalia
guttati). This type of vegetation is threatened by the
invasion of alien plants (Campos et al. 2004; Stanisci
et al. 2010; Zedda et al. 2010). Their conservation
relevance is supported by the inclusion in the
European Union Network ‘‘Nature 2000’’, with the
habitat code ‘‘2230 Malcolmietalia dune grasslands’’
(Loidi 1999; Neto et al. 2007).
Materials and methods
The vegetation relevés have been made according to
the sigmatist landscape phytosociological approach
(Braun-Blanquet 1979; Weber et al. 2000; RivasMartinez 2005). Bioclimatic and biogeographic
typologic follow Costa et al. (1999) and RivasMartinez et al. (2002). A total of 235 relevés were
submitted to UPGMA and principal coordinate
analysis (PCoA), with Bray–Curtis coefficient using
SYNTAX 2000 software (Podani 2001). In addition
to these, some other relevés were taken from
literature (Rivas Goday 1958; Pinto Silva & Teles
1972; Dı́az González & Navarro 1978; RivasMartinez et al. 1980; Dı́ez-Garretas 1984; Izco
et al. 1988; Costa et al. 1994, 1997, 2000; Dı́ezGarretas & Asensi 2002; Neto 2002; Lomba 2004) to
Correspondence: José Carlos Costa, Centro de Botânica Aplicada à Agricultura (CBBA), Instituto Superior de Agronomia, Technical University of Lisbon
(TULisbon), Tapada da Ajuda, 1349-017 Lisboa, Portugal. Tel: 00 351 213653166. Fax: 00 351 213653195. Email: jccosta@isa.utl.pt
ISSN 1126-3504 print/ISSN 1724-5575 online ª 2011 Società Botanica Italiana
DOI: 10.1080/11263504.2011.602729
92
J. C. Costa et al.
compose the synthetic table and run the ordination
and cluster analysis.
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Results and discussion
The PCoA ordination (Figure 1) separated the
relevés of L. pedunculatae (relevés 1-123), Anthyllido
hamosae-Malcolmion lacerae (relevés 124–219) and
Thero-Airion (relevés 220–242). The performed
cluster analysis UPGMA showed a segregation of
the communities from L. pedunculatae (Figure 2) and
Anthyllido hamosae-Malcolmion lacerae (Figure 3).
Likewise, the new associations described show a fine
floristic individuality over the others of the same
alliance.
The order Cutandietalia maritimae was proposed
by Rivas-Martinez et al. (2002) to the Mediterranean
annual associations of semi-fixed dunes, including
the alliance L. pedunculatae, distributed by the west
and northwest of the Iberian Peninsula and North of
Africa. In the Iberian Peninsula and West Morocco,
the order Malcolmietalia comprises the interior sandy
soil communities, namely the Iberian alliance Anthyllido hamosae-Malcolmion lacerae which occurs in
the clearings of bushes and woodlands in the
themomediterranean dry belt.
In the central west Portuguese coast, the relevés
made in semi-fixed dunes do not show the endemics
Linaria ficalhoana Rouy or Herniaria algarvica
Chaudhri, which are present in Herniario algarvicaeLinarietum ficalhoanae in the south. Consequently,
the communities from the north of Lisbon demonstrate a clear floristic individuality that can be
explained by biogeography and climatic reasons.
Both communities are physically separated by an
extension superior to 100 km, of rocky cliffs, coastal
limestone and granite platforms, between Lisbon and
Carvoeiro Cap (Peniche). From a biogeographical
point of view, this lithological discontinuity is critical
in the distribution of the psammophilic plant
communities in the west of the Iberian Peninsula.
There are also important climatic differences, like
higher number of foggy days in spring and summer,
stronger influence of fronts, lower temperatures
through the year and higher levels of rainfall, in the
north. As a result, thermophilic plants have difficulty
in moving north.
When describing the association Violo henriquesiiSilenetum littoreae to the dunes of Galicia, Izco et al.
(1988) extended its distribution area to the Portuguese western coast. However, the relevés made in
this work have revealed that the dune community in
the center of Portugal is different from that occurring
in the northwest of the Iberian Peninsula (Table I,
the bold values mean the type of the association and
the values inside the box are the characteristics
and the different species of each association). Hence,
it is proposed as a new association, Pseudorlayo
minusculae-Polycarpietum alsinifolii (Table II, typus
relevé # 8), according to the following arguments:
Viola kitaibeliana Schultes var. henriquesii (Willk.)
Beck, Polycarpon diphyllum (L.) L., Omphalodes
littoralis Lehm. ssp. gallaecica Laı́nz, Pseudorlaya
pumila (L.) Grande ssp. microcarpa Loret and
Barrandon, Sedum arenarium Brot. and Silene scabriflora Brot. ssp. gallaecica Talavera do not occur in the
south of Aveiro; Pseudorlaya minuscula (Font Quer)
Laı́nz, Polycarpon alsinifolium (Biv.) DC., Hedypnois
Figure 1. PCoA Bray–Curtis coefficient: I L. pedunculatae II Anthyllido hamosae-Malcolmion lacerae III Thero-Airon.
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Annual dune communities in Southwest Europe
93
Figure 2. UPGMA with Bray–Curtis coefficient of the association of Iberian Peninsula of L. pedunculatae: A Ononido variegatae-Linarietum
pedunculatae B Herniario algarvicae-Linarietum ficalhoanae C Pseudorlayo minusculae-Polycarpietum alsinifoli D Violo henriquesii-Silenetum
littoreae.
Figure 3. UPGMA with Bray–Curtis coefficient of the association of Iberian Peninsula of Anthyllido hamosae-Malcolmion lacerae: A
Hymenostemmo pseudanthemidis-Arenarietum emarginatae B Malcolmio lacerae-Anthyllidetum hamosae C Corynephoro macrantheri-Arenarietum
algarbiensis D Omphalodo kunzinskyanae-Evacietum ramosissimae E Tolpido barbatae-Tuberarietum bupleurifoliae F Cerastio diffusi-Vulpietum
fontquerianae G Linario donyanae-Loeflingietum baeticae.
cretica (L.) Dumont-Courset, Silene nicaeensis All.,
Ononis dentata Sol ex Lowe and Pimpinella villosa
Schousb. appear as differentials of this new
community, when facing the Galician-Portuguese.
Malcolmia ramosissima (Desf.) Thell. is differential
to Herniario algarvice-Linarietum ficalhoanae. Violo
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94
Table I. Synthoptic table of dunes associations of Linarion pedunculatae, Anthyllido hamosi-Malcolmion lacerae and Thero-Airion dunes of the Iberian Peninsula and North Africa.
1
13
2
7
3
26
4
11
5
7
6
22
Characteristics and differentials of the Linarion pedunculatae associations
Viola kitaibeliana var. henriquesii
V
II
Pseudorlaya pumila ssp. microcarpa
IV
Polycarpon diphyllum
I
II
Sedum arenarium
II
Omphalodes littoralis ssp. gallaecica
II
Silene scabriflora ssp. gallaecica
I
Malcolmia ramosissima
þ
I
I
Silene littorea
V
III
V
V
IV
V
Polycarpon alsinifolium
V
V
V
V
Pseudorlaya minuscula
IV
III
Silene nicaeensis
II
II
II
Pimpinella villosa
II
II
Ononis dentata
I
Linaria ficalhoana
V
V
Herniaria algarvica
IV
Chaenorhinum serpyllifolium ssp. lusitanicum
Erodium laciniatum (Cav.) Willk.
Romulea gaditana G. Kunze
Hypecoum littorale Wulfen
Ononis variegata
V
Linaria pedunculata
Linaria munbyana ssp. pygmaea
7
11
8
4
9
7
10
15
11
15
12
4
13
3
14
10
15
5
16
13
17
10
18
1
20
28
21
6
22
16
I
V
V
III
4
4
2
IV
IV
V
III
V
IV
III
þ
4
V
3
3
1
II
IV
I
þ
I
1
4
2
4
Differentials of the Anthyllido hamosi-Malcolmion lacerae and Thero-Airion associations
Hymenostemma pseudanthemis
Arenaria emarginata
Scilla ramburei
Hippocrepis salzmannii
Trisetaria dufourei (Boiss.) Paunero
Anthoxanthum ovatum
Evax asterisciflora
Hymenocarpos hamosus
Malcolmia triloba
Loeflingia baetica var. baetica
Linaria tursica
Arenaria algarbiensis
Tuberaria bupleurifolia
Scilla odorata
Lotus castellanus
V
V
þ
I
þ
IV
V
V
V
3
4
I
23
13
24
10
III
III
þ
I
V
III
2
2
3
3
V
V
IV
III
II
III
III
V
V
þ
3
4
4
5
V
V
III
II
1
II
1
I
IV
II
V
IV
III
þ
(continued)
J. C. Costa et al.
Column number
Number of relevés
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Table I. (Continued).
Column number
Number of relevés
Corynephorus macrantherus
Malcolmia gracillima
Omphalodes kuzinskyanae
Jonopsidium acaule
Vulpia fontquerana
Lotus arenarius
Aira praecox
Cerastium diffusum
Asterolinon linum-stellatum
Petrorhagia prolifera (L.) P.W. Ball
Trifolium arvense L.
Koeleria albescens DC.
Ononis reclinata L.
Arenaria serpyllifolia L.
Phleum arenarium L.
Rumex bucephalophorus ssp. bucephalophorus
1
13
2
7
3
26
4
11
5
7
6
22
7
11
8
4
9
7
10
15
11
15
12
4
13
3
14
10
15
5
16
13
17
10
I
18
1
20
28
1
V
II
21
6
22
16
23
13
24
10
IV
II
II
I
IV
III
IV
IV
IV
I
II
I
III
I
III
V
II
IV
III
II
IV
III
II
III
III
þ
r
II
Characteristics of the Malcolmietalia and Anthyllido-Malcolmion lacerae
Erodium bipinnatum
IV
III
V
V
V
II
Rumex bucephalophorus ssp. hispanicus
þ
III
II
III
III
III
Ononis diffusa
þ
Evax ramosissima
III
Coronilla repanda
I
I
Vulpia membranacea.
II
þ
Andryala arenaria
þ
Ononis broterana
r
þ
Linaria spartea
Silene colorata
þ
Leucojum trichophyllum Schousb.
Ononis baetica Clemente
Corynephorus fasciculatus Boiss. & Reut.
Loeflingia baetica var. micrantha
Ornithopus isthmocarpus
Agrostis tenerrima
Silene scabriflora ssp. scabriflora
þ
Ornithopus sativus
III
V
III
3
2
V
4
IV
III
4
1
V
III
III
IV
V
IV
I
IV
II
V
V
III
III
II
IV
4
4
1
4
3
2
V
3
2
II
3
2
V
IV
4
4
2
I
2
II
3
1
2
3
V
III
1
1
IV
þ
I
II
III
þ
II
4
II
II
V
I
V
IV
I
II
III
III
V
V
IV
I
I
þ
IV
V
V
III
5
2
V
4
III
V
V
3
4
5
III
II
þ
I
II
II
IV
III
II
II
2
1
1
1
1
1
II
III
III
IV
1
1
V
V
þ
II
I
II
I
V
II
III
II
III
I
þ
IV
V
IV
I
V
III
IV
IV
I
II
III
I
IV
II
I
II
III
V
II
V
I
III
V
I
I
V
95
(continued)
Annual dune communities in Southwest Europe
Characteristics of the Cutandietalia maritimae
Medicago littoralis Rohde
þ
III
Cutandia maritima
Hedypnois cretica
Pseudorlaya pumila var. pumila
Hedypnois arenaria (Schousb.) DC.
r
II
III
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96
Column number
Number of relevés
1
13
2
7
3
26
4
11
Characteristics and differentials of the Helianthemetea guttatae
Mibora minima (L.) Dev.
II
Hypochoeris glabra L.
I
Leontodon longirostris
IV
II
IV
Ornithopus pinnatus
I
þ
þ
Tuberaria guttata
III
II
II
Trifolium scabrum
I
r
Crassula tillaea Lest.-Garl.
III
Teesdalia nudicaulis
I
Vulpia myuros
II
Logfia minima
I
Euphorbia exigua
r
Micropyrum tenellum
r
Trifolium campestre
I
Scorpiurus muricatus
Briza maxima L.
Silene portensis L.
Dipcadi serotinum
Vulpia bromoides
Ornithopus compressus L.
Teesdalia coronopifolia (J.P. Bergeret) Thell.
Logfia gallica (L.) Coss. & Germ.
Tolpis barbata
Pterocephalus diandrus
Petrorhagia nanteuilii
Lathyrus angulatus L,
Paronychia echinulata Chater
Plantago bellardii
Andryala integrifolia L.
Aira caryophyllea ssp. caryophyllea
Jasione montana
Trifolium angustifolium
Linum bienne
Medicago lupulina L.
Ornithopus perpusillus L.
5
7
6
22
7
11
8
4
9
7
10
15
r
þ
þ
I
2
4
11
15
12
4
13
3
14
10
15
5
16
13
17
10
18
1
20
28
III
4
þ
þ
1
II
V
IV
5
V
II
III
IV
V
1
1
I
V
4
III
21
6
III
IV
1
þ
I
I
3
1
II
III
V
I
II
I
I
II
II
II
II
II
III
4
2
5
3
3
V
II
II
1
22
16
þ
II
III
II
V
II
IV
V
II
III
II
II
II
1
1
V
I
II
II
II
II
III
I
V
II
I
II
III
I
II
24
10
II
II
II
III
III
III
þ
IV
23
13
III
IV
þ
þ
I
þ
II
r
II
1
1
1
II
II
III
II
I
II
V
V
þ
þ
II
III
I
V
I
II
(continued)
J. C. Costa et al.
Table I. (Continued).
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Table I. (Continued).
Column number
Number of relevés
I
þ
þ
þ
þ
I
þ
2
7
3
26
4
11
IV
II
I
III
II
I
IV
5
7
6
22
7
11
8
4
r
I
2
9
7
10
15
II
I
IV
11
15
12
4
13
3
14
10
15
5
16
13
2
2
III
1
I
4
II
II
V
17
10
18
1
20
28
21
6
þ
III
III
V
II
IV
V
2
II
II
IV
1
IV
þ
III
I
I
I
I
IV
II
III
þ
II
2
IV
4
23
13
24
10
III
V
III
þ
II
þ
III
IV
III
I
V
22
16
2
II
V
III
II
III
I
III
II
þ
II
III
IV
1
I
II
þ
II
II
II
II
II
II
I
5
II
IV
4
II
I
þ
II
V
II
I
I
II
IV
Note: Violo henriquesii-Silenetum littoreae: 1 Izco et al. 1988, 2 Lomba (2004); Pseudorlayae misnusculae-Polycarpietum alsinifoli: 3, 4 Costa et al. (2000) under the name Violo-Silenetum littoreae; Herniario
algarvicae-Linarietum ficalhoanae: 5 Diez Garretas (1984), 6 Neto (2002), 7 Costa et al. (1992); Herniario algarvicae-Linarietum ficalhoanae chaenerhitesosum lusitanicae: 8 Costa et al (1992); Ononido
variegatae-Linarietum pedunculatae: 9 Diez Garretas (1984), 10 Costa et al. (1996), 11 Diez Garretas & Ascensi (2002); Ononido variegatae-Linarietum pedunculatae linarietosum pygmaeae: 12 Diez
Garretas (1984), 13 Costa (1991), 14 Costa et al. (1996); Hymenostemmo pseudoanthemidis-Arenarietum emarginatum: 15 Rivas Goday (1958); Malcomio lacerae-Anthyllidetum hamosae: 16 Rivas Goday
(1958); Linario donyanae-Loeflingietum baeticae: 17 Rivas-Martı́nez et al. (1980); Tolpido barbatae-Tubarietum bupleurifoliae: 18 Costa et al. (1996); Corynephoro macrantheri-Arenarietum algarbiensis: 19
Pinto da Silva & Teles (1972), 20 Neto (2002); Omphalodo kunzinskyanae-Evacietum ramosissimae: 21; Cerastio diffusi-Vulpietum fontqueranae: 22; Petrorhagio-Trifolietum arvensis: 23 Dı́az González &
Navarro Andrés (1978); Asterolino linostellati-Rumicetum bucephalophori: 24 Dı́az González & Navarro Andrés (1978).
Annual dune communities in Southwest Europe
Principal companions
Vulpia alopecuros
Paronychia argentea
Lagurus ovatus
Anagallis arvensis
Senecio gallicus
Veronica arvensis L.
Catapodium marinum (L.) C.E. Hubb.
Centranthus calcitrapa
Medicago polymorpha L.
Poa annua L.
Sedum acre L.
Reichardia gaditana
Papaver setigerum DC.
Brassica oxyrrhina Cosson
Linaria viscosa (L.) Chaz.
Silene longicaulis Pourr.
Alkanna tinctoria (L.) DC.
Catapodium rigidum C.E. Hubb.
Chamaemelum mixtum (L.) All.
Lotus edulis L.
Galium murale
Blackstonia perfoliata (L.) Huds.
Lotus corniculatus L.
Aphanes arvensis L.
1
13
97
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98
Table II. Pseudorlayo minusculae-Polycarpietum alsinifoli.
1
6
15
Characteristics of association, alliance and order
Polycarpon alsinifolium
Silene littorea Brot.
2
Erodium bipinnatum
2
Medicago littoralis Rohd
1
Pseudorlaya minuscula
2
Cutandia maritima (L.) W. Barbey
Silene nicaeensis
Hedypnois cretica
Pimpinella villosa
Ononis dentata
Malcolmia ramosissima
Characteristics of Helianthemetea guttati
Leontodon longirostris (Finch & P.D. Sell) Talavera
Rumex bucephalophorus ssp. hispanicus
Vulpia myuros (L.) C.C. Gmel.
Tuberaria guttata (L.) Fourr.
Vulpia membranacea (L.) Dumort.
Logfia minima (Sm.) Dumort.
Coronilla repanda (Poir.) Guss.
Characteristics of Ammophiletea
Euphorbia portlandica L.
Lotus creticus L.
Crucianella maritima L.
Helichrysum picardii Boiss. & Reuter
Medicago marina L.
Seseli tortuosum L.
Armeria welwitschii Boiss.
Anagallis monelli var. microphylla (Ball) Vasc.
Calystegia soldanella (L.) R. Br.
Eryngium maritimum L.
Artemisia crithmifolia L.
Iberis procumbens Lange
Aetheorhiza bulbosa (L.) Cass.
Pancratium maritimum L.
Malcolmia littorea (L.) R.Br.
Cyperus capitatus Vandelli
Anchusa calcarea Boiss.
þ
2
4
18
3
6
21
4
4
16
5
4
12
6
6
20
7
4
20
8
4
11
9
4
14
10
4
15
11
6
21
12
6
22
13
6
21
14
4
13
15
6
17
16
6
22
17
6
22
18
6
19
19
4
18
20
4
15
21
2
18
22
4
20
23
6
25
24
4
19
25
4
14
26
4
23
þ
2
2
1
3
1
2
3
4
1
2
3
3
2
3
3
2
3
1
1
2
2
1
1
1
1
2
2
3
1
1
2
þ
1
3
3
1
þ
3
2
1
1
3
2
2
1
3
2
1
þ
þ
1
3
2
2
2
2
þ
þ
2
2
1
2
1
þ
þ
4
2
1
2
1
1
2
2
2
2
þ
1
1
2
2
3
1
1
þ
þ
1
4
2
2
1
3
2
2
2
þ
1
þ
3
2
3
2
þ
1
1
þ
2
þ
1
1
þ
2
þ
2
þ
2
3
þ
1
1
1
1
þ
þ
1
þ
2
1
þ
1
2
þ
1
þ
þ
1
þ
1
1
1
þ
þ
1
1
þ
1
2
þ
1
3
1
þ
1
þ
þ
1
þ
1
3
1
þ
1
2
1
2
2
1
þ
þ
þ
2
þ
þ
2
þ
1
þ
2
1
3
1
1
þ
2
2
þ
1
þ
þ
þ
þ
.
þ
þ
þ
1
þ
þ
þ
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þ
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þ
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þ
þ
þ
þ
þ
þ
þ
þ
þ
1
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
1
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
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þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
1
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
1
1
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
(continued)
J. C. Costa et al.
# relevé
Sampling area (m2)
Number of species
Downloaded by [Tech Univ of Lisbon Polo Ist] at 06:54 22 August 2011
Table II. (Continued).
# relevé
Sampling area (m2)
Number of species
2
4
18
3
6
21
1
1
þ
þ
þ
þ
þ
þ
1
4
4
16
5
4
12
þ
þ
þ
þ
1
þ
6
6
20
7
4
20
8
4
11
9
4
14
10
4
15
11
6
21
12
6
22
13
6
21
14
4
13
15
6
17
16
6
22
þ
1
þ
þ
1
1
1
þ
þ
1
þ
1
1
þ
þ
þ
2
þ
þ
þ
2
2
2
þ
þ
þ
þ
1
þ
18
6
19
19
4
18
2
þ
þ
þ
þ
1
þ
þ
þ
þ
þ
þ
þ
þ
þ
1
þ
20
4
15
21
2
18
22
4
20
23
6
25
1
þ
1
1
1
1
1
þ
þ
þ
þ
þ
þ
þ
þ
þ
1
1
þ
þ
þ
þ
þ
17
6
22
þ
þ
þ
24
4
19
þ
þ
26
4
23
þ
þ
þ
1
þ
þ
25
4
14
þ
þ
More: Characteristics of Helianthemetea: Lotus arenarius þ in 2, 21; Silene scabriflora ssp. scabriflora þ in 2, 1 in 23; Ornithopus pinnatus (Miller) Durc. þ in 21, 1 in 26; Andryala arenaria þ in 22, 24;
Euphorbia exigua L. þ in 3; Ononis broterana 1 in 11; Trifolium scabrum L. 1 in 14, Micropyrum tenellum (L.) Link 1 in 14; Urospermum picroides (L.) Scop. þ in 2, 24; Asteriscus maritimus (L.) Less. þ in 25;
Characteristics of Ammophiletea: Leontodon taraxacoides (Vill.) Merat þ in 6, 1 in 11; Linaria polygalifolia Hoffmanns. & Link þ in 13, 16; þEuphorbia paralias L. in 9; þMatthiola sinuata (L.) R.Br. in 10;
þOtanthus maritimus (L.) Hoffmanns & Link in 12; þHerniaria maritima Link; Companions: Catapodium marinum (L.) C.E. Hubb. þ in 1, 23, 24; Lobularia maritima (L.) Desv. þ in 23, 25, 26; Cakile
maritima Scop þ in 22, 23; Carex arenaria L. þ in 5; þBromus rigidus Roth in 22; Bromus diandrus Roth in 24.
Places: 1, 2, 3 South of Pedrogão Beach (Leiria), 4, 5 North of Pedrogão Beach (Leiria), 6, 8 Vieira de Leiria Beach (Marinha Grande), 7, 12 North of Nazaré, 9 S. Pedro de Muel (Marinha Grande),
10, 11 Pedra de Ouro Beach (Marinha Grande), 13 S. Martinho do Porto (Alcobaça), 14 Baleal Sul (Peniche), 15 Peniche, 16 Consolação Beach (Peniche), 17 Areia Branca (Lourinhã), 18 Palheiros
de Mira (Mira), 19 Quiaios Beach (Figueira da Foz), 20 Palheiros da Tocha (Cantanhede), 21 Cabo Raso (Cascais), 22 Oitavos (Cascais), 23 Raso Cap (Cascais), 24 Guincho Beach (Cascais), 25
Abano Beach (Cascais), 26 Magoito (Sintra).
Annual dune communities in Southwest Europe
Companions
Vulpia alopecuros (Schousb.) Dumort.
Senecio gallicus Vill.
Sedum sediforme (Jacq.) Pau
Reichardia gaditana (Willk.) Coutinho
Lagurus ovatus L.
Centranthus calcitrapa (L.) Drusf.
Plantago coronopus L.
Corynephorus canescens (L.) P. Beauv.
Anagallis arvensis L.
Parapholis filiformis (Roth) C.E. Hubb.
1
6
15
99
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100
J. C. Costa et al.
henriquesii-Silenetum littoreae have an exclusively
Eurosiberian distribution (Galician–Portuguese), in
thermotemperate (submediterranean) and upper
mesomediterranean bioclimate, occurring in the
clearings of Iberidetum procumbentis and extending
in Portugal until Aveiro. To the south of Aveiro until
Cascais (Coastal Portuguese and Olissiponesean
districts, Dividing Portuguese Sector), the former
association is replaced by Pseudorlayo minusculaePolycarpietum alsinifolii, occupying the clearings of
Loto cretici-Ammophiletum australis and Armerio welwitschii-Crucianelletum maritimae, in upper thermomediterranean to lower mesomediterranean, upper
dry to lower subhumid bioclimate.
The alliance L. pedunculatae (Table I) includes the
following four associations:
(1) Ononido variegatae-Linarietum pedunculatae (relevés 1–29 in Figure 2): a thermomediterranean
association of wide distribution in the southwest
of the Iberian Peninsula, between Algarve and
Almeria, also present in Morocco and Algeria
(Dı́ez-Garretas & Asensi 2002) (Figure 4), with
Linaria pedunculata (L.) Chaz. and Ononis
variegata L. as guideline species. On more
stabilized dunes occurs the subassociation
linarietosum pygmaeae, differentiated by Linaria
munbyana Boiss. and Reut. ssp. pygmaea
(Samp.) Rivas-Mart.;
(2) Herniario algarvicae-Linarietum ficalhoanae (30–
73): in Sadensean and Vincentine coastal
dunes, in the thermomediterranean dry belt.
For lithified calcareous dunes, the subassociation chaenorhinetosum lusitanicae characterized
by Chaenorhinum serpyllifolium (Lange) Lange
ssp. lusitanicum R. Fernandes is described;
(3) Violo henriquesii-Silenetum littoreae (111–123);
(4) Pseudorlayo minusculae-Polycarpietum alsinifolii
ass nova hoc loco (74–110).
Concerning the associations in the more interior
dunes, belonging to Anthyllido hamosae-Malcolmion
Figure 4. Distribution map of annual dune plant communities in Iberian Peninsula and west of North Africa.
101
Annual dune communities in Southwest Europe
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Table III. Cerastio diffusi-Vulpietum fontquereanae.
# relevé
Sampling area (m2)
Number of species
1
6
17
Characteristics of association, alliance
Silene scabriflora ssp. scabriflora
Linaria spartea
Andryala arenaria
Cerastium diffusum
Aira praecox
Vulpia fontquerana
Erodium bipinnatum
Lotus arenarius
Vulpia membranacea
Medicago littoralis
Coronilla repanda
Rumex bucephalophorus ssp. hispanicus
Ononis broteroana
Agrostis tenerrima
Ornithopus isthmocarpus Coss.
and order
4
3
3
1
þ
1
1
þ
3
3
8
24
3
1
1
1
1
þ
þ
þ
1
4
6
25
5
6
24
6
6
27
7
6
26
8
14
40
9
14
31
10
6
19
11
6
16
12
10
21
13
14
22
14
6
28
15
4
22
16
8
19
1
3
2
þ
þ
3
1
1
1
2
þ
1
1
3
2
2
þ
4
1
2
1
1
3
1
þ
þ
1
3
þ
þ
þ
1
2
1
2
þ
1
2
2
2
3
þ
þ
3
þ
1
þ
2
1
2
1
þ
3
þ
þ
3
1
1
1
1
2
þ
1
1
þ
þ
þ
þ
2
þ
2
þ
þ
þ
1
þ
1
1
þ
1
1
þ
1
3
1
þ
þ
1
þ
þ
2
þ
2
1
1
þ
1
3
1
þ
þ
þ
Characteristics of Helianthemetea guttati
Briza maxima L.
1
Tuberaria guttata
Asterolinon linum-stellatum (L.) Duby
Aira caryophellea L. ssp. caryophyllea
Leontodon longirostris
þ
Logfia minima
Tolpis barbata (L.) Gaertn.
Euphorbia exigua
þ
Vulpia myuros
Trifolium campestre L.
Trifolium scabrum
þ
Linum bienne
Plantago bellardii All.
Vulpia bromoides (L.) S.F. Gray
2
Jasione montana L.
Trifolium arvense L.
1
Teesdalia nudicaulis (L.) R.Br.
Ornithopus pinnatus
Companions
Lagurus ovatus
Corynephorus canescens var. maritimus
Aetheorhiza bulbosae
Vulpia alopecurus
Centranthus calcitrapa
Senecio gallicus
Avena barbata Pott ex Link ssp. barbata
Scilla monophyllus Link
Bromus diandrus
Sonchus oleraceus L.
Anarrhinum bellidifolium (L.) Wild.
2
6
24
1
þ
1
3
þ
þ
1
2
3
2
3
1
1
þ
þ
2
1
1
þ
þ
þ
þ
þ
þ
þ
þ
2
1
þ
þ
1
þ
þ
þ
þ
þ
þ
1
1
þ
þ
1
1
1
2
2
2
1
1
þ
þ
1
1
þ
1
þ
1
þ
2
þ
1
þ
þ
3
2
þ
2
þ
þ
þ
1
þ
þ
þ
þ
þ
2
1
1
þ
1
1
1
1
1
1
1
2
1
2
2
2
1
3
1
1
þ
2
1
1
þ
þ
þ
þ
1
1
þ
þ
1
2
þ
þ
þ
þ
1
þ
þ
2
þ
þ
1
þ
þ
þ
þ
1
1
þ
þ
þ
þ
þ
þ
þ
þ
2
þ
þ
þ
þ
þ
þ
þ
1
3
2
2
2
þ
þ
þ
þ
þ
1
2
þ
1
þ
þ
þ
1
þ
þ
þ
1
3
1
1
2
þ
2
þ
þ
þ
þ
þ
1
1
1
1
1
þ
þ
þ
þ
2
þ
þ
2
1
1
1
1
þ
þ
þ
þ
þ
þ
1
þ
1
þ
þ
1
þ
þ
þ
1
þ
þ
þ
þ
þ
þ
More: Characteristics of Helianthemetea: Polycarpon alsinifolium 1 in 1, þ in 8; Silene littorea þ in 1, 5; Scorpiurus muricatus L. þ in 2, 4;
Pterocephalus diandrus (Lag.) Lag. þ in 4, 14, Hedypnois cretica þ in 5, 14; Silene colorata Poir. þ in 6, 1 in 15; Trifolium angustifolium L. þ in 6,
9; Hypochoeris glabra L. þ in 6, 8; Petrorhagia nanteuilii (Lag.) Lag. þ in 7, 8; Mibora minima (L.) Dev. þ in 10; Companions: Herniaria
ciliolata subsp. robusta þ in 2, 9; Silene gallica L. þ in 3, 14; Juncus capitatus Weigel, þ in 3, 16; Lolium rigidum Gaudin, þ in 4, 9; Sedum
sediforme þ in 5, 9; Carex arenaria þ in 8, 12; Vulpia ciliata Dumort. þ in 8, 12; Dipcadi serotinum (L.) Medik. þ in 8, 13; Sesamoides
purpurascens (L.) G. López þ in 8, 16; Chamaemelum mixtum (L.) All. þ in 3; þSeseli tortuosum þ in 5; þMatthiola sinuata in 5; Rumex
acetosella L. ssp. angiocarpus (Murb.) Murb. þ in 8; þAnagallis monelli var. microphylla, in 8; þHelichrysum picardii (pl.) in 12; þSenecio
vulgaris L. in 13; þUrospermum picroides in 14; þOdontites tenuifolia (Pers.) G. Don f. in 16.
Places: 1 Between S.Pedro de Muel and Vieira de Leiria (Marinha Grande), 2 Vieira de Leiria Beach (Marinha Grande), 3 Between Marinha
Grande and Vieira de Leiria; 4, 6, 10, 13 Pataias (Alcobaça); 5, 9 Between Vieira de Leiria and Pedrogão Beach (Leiria), 7, 11 Marinha
Grande; 8 S. Pedro de Muel (Marinha Grande),12 Pilado (Marinha Grande), 13 Calhau (Nazaré).
102
J. C. Costa et al.
Table IV. Omphalodo kunzinskyanae-Evacietum ramosissimae.
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Numerical order
Sampling area (m2)
Altitude (m)
Number of species
1
4
50
18
Characteristics of association, alliance and order
Evax ramosissima
2
Pimpinella villosa
2
Erodium bipinnatum
þ
Omphalodes kuzinskyanae
4
Rumex bucephalophorus ssp. hispanicus
1
Ononis broteroana
1
Vulpia membranacea
Ononis dentata
2
Coronilla repanda
Silene scabriflora ssp. scabriflora
Jonopsidium acaule
Ornithopus isthmocarpus
Ononis diffusa Ten.
Ornithopus sativus Brot.
Linaria spartea
Characteristics of Helianthemetea guttati
Medicago littoralis
Euphorbia exigua
Tuberaria guttata
Ornithopus pinnatus
Scorpiurus muricatus
Companions
Anagallis arvensis
Centaurium erythraea Rafn
Galium murale (L.) All.
Centranthus calcitrapa
Paronychia argentea Lam.
Lagurus ovatus
Narcissus bulbocodium L.
Senecio vulgaris
Anagallis monelli var. microphylla
1
1
2
8
60
24
3
2
40
19
4
4
40
21
5
6
60
25
6
6
70
25
3
3
1
1
3
1
2
1
1
3
2
þ
1
4
1
2
2
2
2
1
3
2
1
1
2
2
1
1
1
þ
þ
1
þ
þ
3
2
þ
1
þ
þ
1
1
þ
1
1
1
þ
þ
1
1
1
2
þ
þ
2
1
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
1
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
þ
3
þ
þ
þ
þ
þ
1
1
þ
þ
þ
þ
More: Characteristics of Helianthemetea: Polycarpon alsinifolium þ in 1; 1 Trifolium arvense 1 in 3; Asterolinon linum-stellatum þ in 3;
Micropyrum tenellum þ in 3; Companions: Lobularia maritima þ in 1, 1 in 5; Armeria welwitschii þ in 1, 5; Chamaemelum nobile (L.) All. var.
discoideum (Willd.) P.Silva 1 in 3, 4; Dipcadi serotinum þ in 3, 4; Polycarpon tetraphyllum (L.) L. þ in 3, 6; Senecio gallicus þ in 4, 1 in 5;
Cerastium glomeratum þ Thuill. in 1; Bellis annua L. 1 in 2; Carlina corymbosa L. þ in 2; Calendula suffruticosa Vahl ssp. algarbiensis (Boiss.)
Nyman þ in 2; Polypogon maritimus Willd. þ in 4, Geranium molle L. þ in 4; Centaurea sphaerocephala L. þ in 5; Seseli tortuosum þ in 5;
Scabiosa atropurpurea (L.) Greuter et Burdet þ in 5; Lupinus angustifolius L. þ in 6.
Places: 1 Between Ursa and Adraga (Sintra), 2 Samarra (Sintra), 3, 4 Abano (Cascais), 5, 6 Between Adraga and Praia Grande (Sintra).
lacerae, the relevés from the Coastal Portuguese
district have revealed an original floristic combination, where elements from Malcolmietalia (like Silene
scabriflora ssp. scabriflora, Vulpia fontquerana Melderis and Stace, Erodium bipinnatum Willd., Linaria
spartea (L.) Chaz., Andryala arenaria (DC.) Boiss
and Reut, Lotus arenarius Brot., Ononis broterana
DC., Coronilla repanda (Poir.) Guss., Agrostis tenerrima Trin., and Rumex bucephalophorus L. ssp.
hispanicus (Steinh.) Rechb. f., etc.), the alliance
Thero-Airion (Cerastium diffusum Pers., Aira praecox
L.), and from the North of the Iberian Peninsula
(Cantabric-Atlantic) are mingled. Therefore, for the
community occurring on old dunes, in the clearings
of Stauracantho genistoidis-Corematetum albi, with
upper mesomediterranean and lower subhumid
bioclimate, the name Cerastio diffusae-Vulpietum
fontqueranae (Table III, typus relevé # 10) is
proposed, distinguishing it from other communities
of Anthyllido hamosae-Malcolmion lacerate, Petrorhagio-Trifolietum arvensis and Astrolino-Rumicietum bucephalophori (Table I) described for Asturias (Dı́az
González & Navarro 1978). A characteristic species
of this new association, Vulpia fontquerana, possesses
a disjoint distribution area, occurring again in the
southwest of the Iberian Peninsula, in Linario
donyanae-Loeflingietum baeticae.
To the Olissiponesean district, in the upper
thermomediterranean and upper dry belt, in dunes
on calcareous platforms, Omphalodo kuzinskyanae-
Annual dune communities in Southwest Europe
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Evacietum ramosissimae is proposed, which includes
Evax ramosissima Mariz, Omphalodes kuzinskyanae
Willk., Erodium bipinnatum, Pimpinella villosa, Ononis
dentata, Ononis broteroana and Jonopsidium acaule
(Desf.) Rchb., etc. (Table IV, typus relevé # 1).
Although near the sea, being positioned in the top of
cliffs in the clearings of Osyrio quadripartitae-Juniperetum turbinatae, these communities do not directly
suffer the sea spray influence. The statistic treatments
have positioned these communities in the alliance
Anthyllido hamosae-Malcolmion lacerae (Figure 1).
The alliance is composed as follows (Table I):
(1) Hymenostemmo
pseudanthemidis-Arenarietum
emarginatae (relevés 1–10 in Figure 3) occurs
on sandy soils, in the Gaditanean littoral
district, with the differentials Hymenostemma
pseudanthemis (G. Kunze) Willk., Arenaria
emarginata Brot., Scilla ramburei Boiss. and
Hippocrepis salzmannii Boiss and Reuter;
(2) Malcolmio lacerae-Anthyllidetum hamosae (24–
28) is an association from fossil dunes on deep
sandy soils of the Gaditan–Onubensean sector,
with Malcolmia triloba (L.) Spreng., Hymenocarpos hamosus (Desf.) Vis, Anthoxanthum ovatum Lag., Evax asterisciflora (Lam.) Pers., etc.;
(3) Linario donyanae-Loeflingietum baeticae (11–23)
on interior dunes of the National Park of
Doñana (Coastal Onubensean District), characterized by Linaria tursica Valdés and Cabezudo, Loeflingia baetica Lag. var. baetica, etc.;
(4) Tolpido barbatae-Tuberarietum bupleurifoliae (29–
38) on the littoral sands of the Algarvian
district, in the clearings of Cistetum bourgaeani
and Tuberario majoris-Stauracanthetum boivinii,
with Tuberaria bupleurifolia, Scilla odorata and
Lotus castellanus as differentials;
(5) Corynephoro macrantheri-Arenarietum algarbiensis
(39–67) on the old dunes of the Sadensean district,
characterized by Arenaria algarbiensis Willk., Corynephorus macrantherus Boiss. and Reut., Malcolmia
gracillima Samp., Loeflingia baetica var. micrantha
(Boiss. and Reut.) Samp., etc.;
(6) Cerastio diffusae-Vulpietum fontqueranae ass nova
hoc loco (74–89);
(7) Omphalodo kuzinskyanae-Evacietum ramosissimae Neto, J.C. Costa, J.P. Fonseca and Lousã
ass nova hoc loco (68–73).
Sintaxonomic scheme
HELIANTHEMETEA GUTTATI (Br.-Bl. in Br.-Bl.,
Roussine & Nègre 1952) Rivas Goday & Rivas-Martı́nez
1963 em. Rivas-Martı́nez 1978
*Helianthemetalia guttati Br.-Bl. in Br.-Bl., Molinier
& Wagner 1940 em. Rivas-Martı́nez 1978
103
**Thero-Airion Tüxen & Oberdorfer 1958
Asterolino lino-stellati-Rumicetum bucephalophori T.E.
Dı́az & F. Navarro 1978
Petrorhagio-Trifolietum arvensis T.E. Dı́az & F. Navarro
1978
*Malcolmietalia Rivas Goday 1958
**Anthyllido hamosae-Malcolmion lacerae Rivas
Goday 1958 em. Rivas-Martı́nez 1978
Hymenostemmo pseudanthemidis-Arenarietum emarginatae
Rivas Goday 1958
Malcolmio lacerae-Anthyllidetum hamosae Rivas Goday
1958
Linario donyanae-Loeflingietum baeticae Rivas-Martı́nez,
Costa, Castroviejo & E. Valdês 1980
Tolpido barbatae-Tuberarietum bupleurifoliae J.C. Costa in
J.C. Costa, Lousã & Espı́rito-Santo 1997
Corynephoro macrantheri-Arenarietum algarbiensis P. Silva
& Teles ex Rivas-Martı́nez & Izco 2002
Cerastio diffusi-Vulpietum fontquerianae J.C. Costa, Neto,
M. Martins & Lousã ass. nova hoc loco
Omphalodo kuzinskyanae-Evacietum ramosissimae Neto,
J.C. Costa, J.P. Fonseca & Lousã ass. nova hoc loco
*Cutandietalia maritimae Rivas-Martı́nez, Dı́ez Garretas & Asensi 2002
**L. pedunculatae Diez Garretas, Asensi & Esteve in
Diez Garretas 1984
Herniario algarvicae-Linarietum ficalhoanae Dı́ez Garretas 1984
linarietosum ficalhoanae
chaenorrhinetosum lusitanicae J.C. Costa, Espı́rito Santo
& Lousã 1994
Ononido variegatae-Linarietum pedunculatae Diéz Garretas ex Izco, P. & J. Guitián 1988
linarietosum pedunculatae
linarietosum pygmaeae Dı́ez Garretas 1984
Violo henriquesii-Silenetum littoreae Izco & Guitián
1988
Pseudorlayo minusculae-Polycarpietum alsinifoli J.C. Costa,
Neto, M. Martins & Lousã ass. nova hoc loco
All communities from L. pedunculatae and Anthyllido hamosae-Malcolmion lacerae have specific floristic
compositions and biogeographic areas, allowing the
characterization of districts. More rarely they correspond to unique thermotype and ombrotype. Considering the alliance Anthyllido hamosae-Malcolmion
lacerae as an endemic element characteristic from this
Province, its distribution to the entire Lusitan–
Andalusian Coastal Province is proposed.
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José Costa