136Vegetation Ecology and DiversityVED3033-1447Società Italiana di Scienza della Vegetazione (S.I.S.V.)10.3897/ved.142914142914Research ArticleAngiospermaeHabitats DirectivePhanerogamic and Cryptogamic Vegetation survey and classificationPlant Community Conservation and ManagementPlant Ecology and SynecologySyntaxonomy and NomenclatureVegetation classification and conservation aspects of Atlantic dune pine forests in the southwest of the Iberian PeninsulaCostaJosé Carloshttps://orcid.org/0000-0002-7619-840X1NetoCarloshttps://orcid.org/0000-0003-0912-02551Monteiro-HenriquesTiagohttps://orcid.org/0000-0002-4206-06992PinaAna Ritahttps://orcid.org/0000-0002-4928-25851AguiarCarloshttps://orcid.org/0000-0001-8643-71123MasciaFrancescohttps://orcid.org/0000-0002-8777-89284BonariGianmariagianmaria.bonari@unisi.ithttps://orcid.org/0000-0002-5574-606745Linking Landscape, Environment, Agriculture and Food (Leaf), Laboratory Associate TERRA, Instituto Superior de Agronomia, University of Lisbon, Lisbon, PortugalCentro de Estudos Geográficos, Laboratory Associate TERRA, Institute of Geography and Spatial Planning, University of Lisbon, Lisbon, PortugalGlobal Change and Conservation Lab (GCC), Faculty of Biological and Environmental Sciences, University of Helsinki, Helsinki, FinlandLinking Landscape, Environment, Agriculture and Food (Leaf), Laboratory Associate TERRA, Instituto Superior de Agronomia, University of Lisbon, Lisbon, PortugalCIMO-Centro de Investigação de Montanha, Instituto Politécnico de Bragança, Bragança, PortugalDepartment of Life Sciences, University of Siena, Siena, ItalyNBFC, National Biodiversity Future Center, Palermo, Italy
2025300620256211229D14CF0-8A4A-5592-A005-9FB1C0EAD4C7157890602711202403062025José Carlos Costa, Carlos Neto, Tiago Monteiro-Henriques, Ana Rita Pina, Carlos Aguiar, Francesco Mascia, Gianmaria BonariThis is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
This study refines the classification framework for Portuguese dune pine forests included in the alliance Coremato albi-Pinion pinastri at the association level after the syntaxonomic revision of Mediterranean pine forests of the class Pinetea halepensis. We collected 61 original vegetation plots between 2017 and 2022 in the southwestern Iberian Peninsula, analysing them through Principal Coordinate Analysis (PCoA) and UPGMA clustering. Numerical analysis identified four associations within the alliance Coremato albi-Pinion pinastri, specific to the sandy Atlantic coast of the Iberian Peninsula. Of these, three associations are newly described (Cytiso grandiflori-Pinetum pinastri, Aristolochio baeticae-Pinetum pineae, and Ulici australis-Pinetum pineae). Our study classified pine forests on sand in southwestern Iberia and identifies conservation-significant species in their understory. The floristic diversity in these psammophilous pine forests is enhanced by numerous endemics and relic species from the Tertiary period, shaped by past climatic refugia and multiple floristic migration routes, including Atlantic, Mediterranean, and North African pathways. This has resulted in a unique blend of thermophilic, oceanic, and xeric species with high conservation value. Our study contributes to the understanding of Atlantic dune pine forest ecology and inform habitat conservation efforts.
Florapsammophilous pine forestsphytosociologyPinetea halepensisPinus pinasterPinus pineavegetationsand dunesThis work was funded by the ‘Natura 2000 Habitat Mapping Project’ (CART-PG RN2000 - POSEUR-03-2215-FC-000005).Introduction
Pine forests are a widespread vegetation type across the Mediterranean Basin. Numerous studies have focused on these communities dominated by Pinus pinaster and/or Pinus pinea in various regions (Desole 1964; Arrigoni 1967; Agostini 1973; Brullo et al. 1977, 2002; Barbéro et al. 1998; Gianguzzi 1999; Biondi and Vagge 2015; Bonari et al. 2017, 2018, 2020; Calvia et al. 2022a, 2022b; Ciaramella et al. 2024).
During the early Holocene, forest communities dominated by Pinus pinaster thrived in sandy coastal areas of the southwestern Iberian Peninsula, with macrofossil evidence dated between 7,930 and 7,430 cal years BP (García-Amorena et al. 2007). In regions south of Lisbon, Pinus pinea is also well represented in the paleoecological record – particularly through macroremains and charcoal – dating back to approximately 6,300–6,400 ¹⁴C years BP (Carrión et al. 2000; Carrión Marco 2005). These data suggest that both species were key components of coastal vegetation during the early-mid Holocene, even after their decline inland, where Mediterranean taxa became more dominant (Figueiral 1995).
Paleoecological and historical evidence confirm that both Pinus pinaster and Pinus pinea are native to Portugal (Teixeira and Pais 1976; Mateus 1992; Van Leeuwaarden and Queiroz 2003; Aguiar and Capelo 2008). Its consistent presence in Portuguese coastal areas since the Holocene (Rego 2001; Reboredo and Pais 2014), along with its ecophysiological and phenological traits, supports its long-standing adaptation to dynamic dune ecosystems, particularly at elevations below 500 meters (Aguiar and Capelo 2008).
Ecological differences between the two species are notable, especially in their fire responses. Pinus pinaster is a pyrophytic species, typically associated with high-biomass, flammable communities dominated by ericaceous and other fire-adapted taxa on western coastal sands. In contrast, Pinus pinea forms less flammable communities, often with sparse understories and few ericaceous species. Its reproductive biology and resin production traits indicate a lower adaptation to frequent, short-interval fires (Aguiar and Capelo 2008). These differences support the hypothesis of an originally allopatric distribution in the western and southwestern Iberian Peninsula, with each species occupying distinct ecological niches. Currently, pine forest communities are (co-)dominated by these two species that define both the structure and physiognomy of the vegetation.
Along the coast, forests of Pinus pinaster subsp. pinaster and Pinus pinea are designated as priority habitats on dunes under the EU Habitats Directive 92/43/EEC (Costa et al. 2007). According to the Interpretation Manual of EU Habitats (European Commission 2013), these formations are classified as priority habitat 2270, defined as current or ancient dune systems supporting forests of Pinus pinaster and/or Pinus pinea. In the EUNIS habitat classification system, these forests fall under type T3A, “Mediterranean lowland to submontane Pinus forest”, and are found across southern European countries (Chytrý et al. 2020).
However, Mediterranean pine forests lacked a well-defined phytosociological framework. Formerly, Pinus pinaster dune forests of southwestern Iberian peninsula had been variably assigned to the classes Quercetea ilicis, Calluno-Ulicetea, and Cisto-Lavanduletea stoechadis. The study by Bonari et al. (2021) represents the first attempt to formally classify these dune pine forests within a revised phytosociological framework at the class and alliance levels. Within the recently defined phytosociological class Pinetea halepensis Bonari et Chytrý in Bonari et al. (2021), forests in the southwestern Atlantic region of the Iberian Peninsula – dominated by Pinus pinaster and/or Pinus pinea and primarily occurring on sandy coastal soils – have been classified under the alliance Coremato albi–Pinion pinastri J.C. Costa, Neto, Capelo, Aguiar, Monteiro Henriques et Bonari in Bonari et al. (2021). This alliance encompasses pine forests of Pinus pinaster and Pinus pinea with a psammophilous understory rich in species restricted to southwestern Iberia. It is distributed along the coast within the thermo- to meso-Mediterranean bioclimatic zones of the western and southwestern Iberian Peninsula (Lusitanian coast and West Andalusia province; Preislerová et al. 2022). Nevertheless, subordinate syntaxonomic levels (i.e. associations) remain undefined, a knowledge gap that the present study seeks to address.
This study presents a phytosociological analysis of dune pine forests classified in the alliance Coremato albi–Pinion pinastri within priority habitat 2270 along the Atlantic coast of the Iberian Peninsula. It aims to i) describe and characterise the distinct plant communities of this habitat in Portugal and ii) underscore the conservation value of these forest formations and related flora.
Material and methodsStudy area
The psammophilous pine forests studied here occupy a coastal region extending from the Ria de Aveiro in Portugal to the sands of the Costa del Sol in southwestern Spain. Influenced heavily by the Atlantic Ocean, this area is predominantly characterised by sandy and limestone substrates (Neto et al. 2007; Espírito-Santo et al. 2017). This coastal area faces west toward the Atlantic Ocean, where summer advection fogs are common (Daveau 1985).
Data collection and preparation
From 2017 to 2022, we conducted a phytosociological survey of pine forests on coastal sand dunes, collecting a total of 61 original vegetation plots (Suppl. material 1: table S1–S5) in habitat 2270, including both mature and newly stabilised dunes along the Iberian Atlantic coast, from Esmoriz to the Algarve. Fieldwork was concentrated in locations that were either previously documented in phytosociological literature or identified through a prior habitat mapping project.
The nomenclature for vascular plants in this study adheres to Flora Iberica (Castroviejo 1986–2012; Cubas 2000) and Flora-On (2023). Syntaxonomic nomenclature for associations outside of pine forests is based on Costa et al. (2012), while higher taxonomic ranks follow Mucina et al. (2016), apart from pine forests, for which we adhere to Bonari et al. (2021). Syntaxonomy aligns with the fourth edition of the International Code of Phytosociological Nomenclature (ICPN; Theurillat et al. 2021). All vegetation data are maintained in the CircumMed Forest Database (Bonari et al. 2019).
Bioclimatic classification for the description of the associations is based on Rivas-Martínez et al. (2011) and Monteiro-Henriques et al. (2016), while biogeographic units follow the system by Rivas-Martínez et al. (2011, 2017).
Analysis
To investigate differences in species composition among coastal sand dune pine forest types, we analysed the 61 original vegetation plots (relevés) collected in our survey. Transformation of cover-abundance values for numerical treatment follows van der Maarel (1979, 2007). The data were subjected to Principal Coordinate Analysis (PCoA) using the Bray-Curtis coefficient with CANOCO 5.5 software (ter Braak and Šmilauer 2012) and a UPGMA hierarchical clustering also based on Bray-Curtis coefficients, conducted with SYNTAX 2000 software (Podani 2001). We interpreted the UPGMA clusters by comparing species composition and analysing ecological gradients using literature-based references. Community definitions and synthetic table arrangements were developed using floristic-statistical group analysis following Müller-Dombois and Ellenberg (1974).
To emphasise the conservation significance of the vegetation and associated flora of pine forests (priority habitat 2270) in the Southwestern Iberian Peninsula, we prepared a list of taxa with conservation value based on their inclusion in the Habitats Directive annexes and their IUCN extinction risk category for mainland Portugal (IUCN conservation status; Carapeto et al. 2020), and level of endemism (Carapeto et al. 2020).
Results
The floristic composition analysis of the vegetation plots enabled us to distinguish four pine forest associations, of which three are newly described. The results of the Principal Coordinate Analysis (PCoA) are presented in Figure 1.
The PCoA shows plant community patterns on axis 1 according to latitude, reflecting the change in climatic conditions along the coast from north to south. Total rainfall values decrease from north to south and temperature values increase from north to south, in combination with summer fogs, which are very frequent north of Lisbon and gradually decrease southwards. These fogs are very important because they provide plants with water in the driest season. Thus, the associations found on axis 1 are organised according to this gradient, with the northernmost community (Cytiso grandiflori-Pinetum pinastri) occupying the highest values on axis 1, the Pinetum pinastro-pineae community occupying south-west Portugal and the other two associations (Aristolochio-Pinetum pineae and Ulici-Pinetum pineae) having a more southerly distribution, occupying the negative values on axis 1.
Our hierarchical clustering analysis highlights the separation among our pine forest communities (Figure 2). The first division can be interpreted as the distinction between the mesomediterranean community and the other thermomediterranean communities.
Figure 3 shows the distribution of the four pine forest communities. The abbreviated synthetic table (Table 1) presents species percentages across the four pine forest communities. The full table is reported in Suppl. material 1: table S6.
Principal Coordinate Analysis (PCoA) with Bray-Curtis coefficient, generated using CANOCO 5.5 software, based on 61 vegetation plots. Four distinct groups are evident, corresponding to the associations identified in this study. The spatial distribution of the associations is depicted in Figure 3.
Distribution of plant communities in the western and southwestern regions of the Iberian Peninsula.
https://binary.pensoft.net/fig/1362497
Shortened synthetic table of the four pine forest communities: 1 = Cytiso grandiflori-Pinetum pinastri; 2 = Pinetum pinastro-pineae; 3 = Aristolochio baeticae-Pinetum pineae; 4 = Ulici australis-Pinetum pineae. Cells in grey represent the diagnostic species of each plant community.
Pine forest community
1
2
3
4
N° of vegetation plots
13
22
16
9
Pine species
Pinus pinaster
100
100
88
11
Pinus pinea
.
100
100
100
Diagnostic species
Cytisus grandiflorus subsp. grandiflorus
92
.
.
.
Ulex latebracteatus
85
.
.
.
Myrica faya
69
.
.
.
Erica arborea
38
.
.
.
Ulex australis subsp. welwitschianus
.
52
.
.
Juniperus navicularis
.
26
.
.
Thymus capitellatus
.
30
.
.
Stauracanthus spectabilis
.
26
.
.
Santolina rosmarinifolia
22
.
.
Ulex jussiaei
.
13
.
.
Thymus camphoratus
.
9
.
.
Armeria rouyana
.
9
.
.
Chamaerops humilis
.
.
100
.
Aristolochia baetica
.
.
81
.
Stauracanthus boivinii
.
.
81
.
Asphodelus serotinus
.
.
81
.
Ulex argenteus subsp. subsericeus
.
.
75
.
Asparagus albus
.
.
63
.
Centaurea aspera subsp. stenophylla
.
.
38
.
Genista hirsuta subsp. algarbiensis
.
.
25
.
Cistus libanotis
.
.
19
.
Thymus lotocephalus
.
.
19
.
Tuberaria major
.
.
13
.
Juniperus macrocarpa
.
.
.
100
Ulex australis subsp. australis
.
.
.
56
Thymus mastichina subsp. donyaea
.
.
.
11
Other species
Juniperus turbinata
8
87
.
100
Corema album
100
61
.
33
Rosmarinus officinalis
.
17
.
100
Asparagus aphyllus
15
96
100
22
Pistacia lentiscus
46
91
94
89
Daphne gnidium
54
83
94
56
Helichrysum picardii
54
70
69
100
Rubia peregrina
69
61
63
67
Lavandula pedunculata subsp. lusitanica
38
57
75
44
Halimium calycinum
92
52
56
67
Stauracanthus genistoides
38
26
.
56
Asparagus acutifolius
8
.
31
100
Rhamnus oleoides
.
17
38
89
Osyris quadripartita
.
13
44
56
Myrtus communis
.
4
38
11
Halimium halimifolium subsp. multiflorum
62
48
.
.
Ruscus aculeatus
54
9
.
.
Scilla monophyllos
54
22
.
.
Cytisus grandiflorus subsp. cabezudoi
.
.
81
78
Halimium halimifolium subsp. halimifolium
.
.
56
100
Thymus albicans
.
.
56
11
Armeria macrophylla
.
.
31
11
Euphorbia boetica
.
.
31
11
Corynephorus canescens var. maritimus
38
57
6
11
Scrophularia frutescens
23
22
.
11
Artemisia crithmifolia
15
17
.
.
Pine forest communities
We describe the four pine forest communities (Figure 4) and provide information on the holotype for three newly described communities, diagnostic species, structure, ecology, dynamics, and chorology.
Photographs of the four pine forest communities. A)Cytiso grandiflori-Pinetum pinastri at Pinhal de Leiria (Portugal) – Pine forest of Pinus pinaster with an understorey of Myrica faya. B)Pinetum pinastro-pineae at Comporta (west coast, south of Lisbon) – Pine forest of Pinus pinaster with an understorey of Corema album, Juniperus turbinata, among others. C)Aristolochio baeticae-Pinetum pineae at Faro near Ria Formosa (southern coast, eastern Algarve) – Pine forest of Pinus pinea with an understorey of Ulex argenteus subsp. subsericeus, Osyris quadripartita, Daphne gnidium, Halimium halimifolium subsp. halimifolium, Stauracanthus boivinii, among others. D)Ulici australis-Pinetum pineae near Huelva (southwestern Spain) – Pine forest of Pinus pinea with an understorey of Juniperus macrocarpa, among others. Photo credits: A: J. C. Costa; B and C: C. Neto; D: A. R. Pina.
https://binary.pensoft.net/fig/1362498
Cytiso grandiflori-Pinetum pinastri J.C. Costa, Monteiro-Henriques et Neto ass. nova (typus relevé no. 12, Suppl. material 1: table S1)
Type relevé (holotypus): Pinus pinaster 4, Cytisus grandiflorus subsp. grandiflorus 2, Corema album 2, Halimium calycinum 2, Myrica faya 1, Phillyrea angustifolia 1, Rubia peregrina 1, Ulex latebracteatus +, Arbutus unedo +, Cistus salviifolius +, Pinus pinaster (juvenile) +, Other species: Calluna vulgaris. 1. Area: 200 m², Aspect: W; Locality: between Praia de Vieira and São Pedro de Muel, Leiria, Portugal.
Structure: A pine forest dominated by Pinus pinaster, ranging from 5 to 15 m in height, with a dense understorey of psammophilous and sclerophyllous shrubs, as well as some lianas and herbs. Typical species include Corema album, Cytisus grandiflorus, Phillyrea angustifolia, Halimium calycinum, Cistus salviifolius, Ulex latebracteatus, Myrica faya, Arbutus unedo, Rubia peregrina, Halimium halimifolium subsp. multiflorum, Ruscus aculeatus, Helichrysum picardii, Daphne gnidium, Stauracanthus genistoides, Lavandula pedunculata subsp. lusitanica, Erica arborea, Scilla monophyllos, Pistacia lentiscus, and Calluna vulgaris. In many areas, this association has been replaced by Acacia longifolia, an invasive species that has spread significantly in recent decades, particularly following fires.
Ecology: Found on recent and ancient Quaternary dunes at elevations of 5–60 m above sea level, within a lower sub-humid upper mesomediterranean bioclimate.
Dynamics: Generally situated as a vegetation strip adjacent to the dune interiors and Juniperus series (Osyrio quadripartitae-Juniperetum turbinatae, Rubio longifoliae-Coremetum albi). Further inland, Cytiso grandiflori-Pinetum pinastri may border Lauro nobilis-Quercetum rivasmartinezii or Myrico fayae-Arbutetum unedonis, which occupy thin dune sands, often overlaying Miocene-Pliocene formations in the innermost dune complex areas (Henriques and Neto 2002; Neto 2012). Subserial stages include the scrubland Ulici latebracteati-Cytisetum striati, the psammophilic thicket Stauracantho genistoidis-Corematetum albi, and the annual grasslands Cerastio diffusi-Vulpietum fontqueranae and Herniario robustae-Corynephoretum maritimi.
Chorology: Ranges from south of Espinho to Nazaré, covering Porto and Low Douro District (Galicia and North Portugal sector, Cantabrian-Atlantic subprovince, Atlantic European province, Eurosiberian region, and Littoral Beira district, Lusitanian coast, and West Andalusia province, Mediterranean region).
Pinetum pinastro-pineae J.C. Costa et Neto in Bonari et al. 2021 (Suppl. material 1: table S2)
Note: In the type relevé (holotypus) published in Bonari et al. (2021), Ulex parviflorus subsp. parviflorus is reported. However, this taxon is a complex of species including U. jussiaei, U. australis subsp. australis, U. australis subsp. welwitschianus, and U. airensis.
Structure: Mixed pine forest of Pinus pinea and Pinus pinaster, reaching heights of 5–20 m, characterised by a dense understorey of psammophilous, thermophilous, and sclerophyllous shrubs, vines, and herbs. Species include Asparagus aphyllus, Pistacia lentiscus, Cistus salviifolius, Juniperus turbinata, Daphne gnidium, Helichrysum picardii, Corema album, Rubia peregrina, Ulex australis subsp. welwitschianus, Lavandula pedunculata subsp. lusitanica, Halimium calycinum, Phillyrea angustifolia, Rhamnus alaternus, Halimium halimifolium subsp. multiflorum, Olea europaea var. sylvestris, Thymus capitellatus, Juniperus navicularis, Stauracanthus spectabilis, S. genistoides, Smilax aspera, Thapsia villosa var. platphyllos, Quercus rivasmartinezii, and other dune species such as Corynephorus canescens var. maritimus, Ononis ramosissima, and Euphorbia portlandica. The diagnostic species of this community include several endemic taxa, many of which are exclusive to Portugal and listed in Annexes II and IV of the Habitats Directive.
Ecology: Located on sandy soils of recent and ancient Quaternary dunes, at elevations of 5–60 m above sea level, within an upper dry to lower sub-humid thermomediterranean bioclimate.
Dynamics: The Pinetum pinastro-pineae represents the potential natural vegetation in most areas of recent dunes, with many species from Juniperus turbinata and Corema album shrub formations in the understorey, and primarily in ancient dunes where species of the Juniperus navicularis series are prevalent. Inland, it borders cork oak forests (Aro neglecti-Quercetum suberis). The subserial stages are primarily psammophilous plant communities, such as Thymo capitellati-Stauracanthetum genistoidis (Sado District), Thymo camphorati-Stauracanthetum spectabilis (São Vicente Coastal District), Erico umbellatae-Ulicetum welwitschiani (Sado and São Vicente Coasts), Euphorbio transtaganae-Celticetum giganteae (Sadense and Costeiro Vicentino), Lavandulo luisieri-Ulicetum jussiaei (Estremadura, Coimbra, and Lisbon Districts), Omphalodo kuzinskyanae-Evacetum ramosissimae (Lisbon), Corynephoro macrantheri-Arenarietum algarbiensis (Sado and São Vicente Coasts), Herniario maritimae-Corynephoretum maritimi (Lisbon and Sado), and Herniario unamunoanae-Corynephoretum maritimi (Sado and São Vicente Coasts).
Chorology: Extending from Nazaré to Cabo São Vicente, covering the Estremadura and Coimbra, Lisbon, Sado, and São Vicente Coastal Districts (including Ribatejo and Sado in the Portuguese sector, Cádiz and Sado subprovince, Lusitanian coast, and West Andalusia province, Mediterranean region).
Aristolochio baeticae-Pinetum pineae J.C. Costa, Neto et A.R. Pina ass. nova (typus relevé no. 42, Suppl. material 1: table S3)
Structure: Pinus pinea forest with Pinus pinaster, 10–20 m in height, supported by a dense understorey of psammophilous, thermophilous, and sclerophyllous shrubs, vines, and herbs, including Asparagus aphyllus, Chamaerops humilis, Pistacia lentiscus, Cistus salviifolius, Daphne gnidium, Olea europaea var. sylvestris, Thapsia villosa var. platyphyllos, Aristolochia baetica, Cytisus grandiflorus subsp. cabezudoi, Stauracanthus boivinii, Asphodelus serotinus, Ulex argenteus subsp. subsericeus, and more. This forest is rich in endemic and rare taxa, some listed in Annexes II and IV of the Habitats Directive.
Ecology: Sandy soils, sometimes with gravel beds near the coast, at 0–20 m a.s.l., within a dry lower thermomediterranean bioclimate.
Dynamics: Represents the potential natural vegetation of the Algarve’s sandy coastal soils. Substitution stages include: Cytisetum cabezudoi, Cistetum libanotidis, Tuberario majoris-Stauracanthetum boivinii, Armerio macrophyllae-Celticetum giganteae, and Tolpido barbatae-Tuberarietum bupleurifoliae. Inland, it transitions to cork oak forests (Aro neglecti-Quercetum suberis).
Chorology: Distributed in the Algarve district (Algarve and Monchique sector, Cádiz and Sado subprovince, Lusitanian coast, and West Andalusia province, Mediterranean region).
Ulici australis-Pinetum pineae J.C. Costa et A.R. Pina ass. nova (typus relevé no. 54, Suppl. material 1: table S4)
Structure: A Pinus pinea forest, 10–20 m in height, with a dense understorey of psammophilous, thermophilous, and sclerophyllous shrubs, vines, and herbs, such as Juniperus turbinata, Asparagus acutifolius, Halimium halimifolium subsp. halimifolium, Helichrysum picardii, Rosmarinus officinalis, Juniperus macrocarpa, Pistacia lentiscus, Rhamnus oleoides, and others.
Ecology: Within a dry lower thermomediterranean bioclimate at elevation between 5 and 30 m a.s.l.
Dynamics: On the coast, Ulici australis-Pinetum pineae contacts Juniperus turbinata communities (Osyrio quadripartitae-Juniperetum turbinatae) and other coastal communities, transitioning inland to cork oak forests (Aro neglecti-Quercetum suberis). Substitution stages include Halimio halimifolii-Stauracanthetum genistoidis, Asphodelo aestivi-Armerietum gaditanae, and Linario donyanae-Loeflingietum baeticae.
Chorology: Cadiz and Littoral Huelva sector (Cádiz and Sado subprovinces, Lusitanian coast, and West Andalusia province, Mediterranean region).
We found 19 taxa of conservation concern with reference to Habitats Directive annexes (17 Annex I species, of which 4 of priority importance, and 18 Annex IV species), IUCN conservation status (2 critically endangered, 1 endangered, 2 vulnerable, 4 near threatened, and 10 least concern) in mainland Portugal and endemism levels (17 endemic to mainland Portugal, 2 endemic to Iberian Peninsula) (Table 2).
Flora of conservation concern in the understory of Atlantic dune pine forests in the southwestern Iberian Peninsula (priority habitat 2270), with reference to Habitats Directive annexes, extinction risk category in mainland Portugal (IUCN conservation status; Carapeto et al. 2020), and level of endemism. Species are listed in alphabetical order. The asterisk indicates a species present in our vegetetation plots.
Species
Habitats Directive annex
Extinction risk in Portugal according to IUCN category
We identified distinct pine forest communities in southwestern Iberia, including new associations on ancient coastal dunes. Their rich, psammophilous flora is shaped by climate and biogeographic boundaries. This region hosts many endemic and relict species due to its role as a climatic refuge and intersection of Mediterranean, Atlantic, and North African migration routes.
Pine forests associations
A recent large-scale study examined pine forest communities across lowland and submontane areas in 20 Mediterranean countries, resulting in the classification of Mediterranean pine forests within the class Pinetea halepensis (Bonari et al. 2021). In the Iberian Peninsula, pine forests are a dominant forest type, ranging from mountainous to lower elevations. In southwestern Iberia, these forests are classified within the alliance Coremato albi-Pinion pinastri J.C. Costa, Neto, Capelo, Aguiar, Monteiro Henriques et Bonari in Bonari et al. 2021. Our analysis of vegetation plots of the alliance Coremato albi-Pinion pinastri identified four associations, of which three of them are new, within these ancient dunes, formed during the Pleistocene glaciations and often subject to podzolisation.
North of Nazaré to Porto, summer fog significantly reduces water stress for coastal plants, unlike the southern coastal regions. As a result, many thermophilic coastal psammophilous species with southern distributions are absent in this sector, leading to a separation between mesomediterranean and thermomediterranean communities. This separation reflects the biogeographic boundary between the Eurosiberian and Mediterranean regions, located between Aveiro and Lisbon, where flora and vegetation from both regions converge due to the lack of biogeographic barriers.
Flora of conservation concern
The floristic composition of the psammophilous understory in pine forests varies along a coastal-to-inland gradient, influenced by soil and air salinity, sandy substrate mobility, temperature, wind, and soil chemistry, as well as latitudinal changes in climate. An important ecological factor along the western Iberian coast is horizontal precipitation from coastal summer fog, which declines in frequency from north to south (Daveau 1977, 1985). This fog supplies water to plants during the crucial summer months, influencing the distribution of species and communities. Thus, topography, climate, and biogeographical history contribute to the understory composition, resulting in a diverse flora with numerous Portuguese and Iberian endemics along the sandy coast (Costa 1992; Costa et al. 1997; Neto 2002; Neto et al. 2004, 2005).
The coastal vegetation in southwestern Iberia is notably rich in endemic Paleomediterranean and Paleotropical lianoid and lauroid species with leathery leaves (Costa et al. 1998; Costa et al. 2000; Espírito-Santo et al. 2017; Fernández-Palacios et al. 2017; Neto et al. 2019). This region’s oceanic or hyperoceanic climate, with mild winters, has provided a refuge for numerous thermophilic and naked bud plants, which were relatively unaffected by Pleistocene glaciations and Holocene climatic shifts (Médail and Diadema 2009; Minkley et al. 2015; Gómez-Orellana et al. 2021). Such plants, typical of oceanic thermophilic forests, are absent in the more continental forests of holm and cork oaks, likely due to their inability to recolonise colder inland regions of Iberia during the Holocene.
This unique climatic and palaeoecological context also facilitated the persistence of palaeomediterranean Tertiary elements with affinities to the Macaronesian Subregion (e.g., Myrica faya, Convolvulus fernandesii, Cheilantes guanchica, Polypodium macaronesicum, Woodwardia radicans) and allowed the arrival of Mauritanian and Pontic-Indian flora (Costa et al. 1998; Vargas et al. 2018; Ramos-Capón et al. 2024). This area lies at the intersection of several floristic migration routes, primarily the Mediterranean coastal route and the Betic limestone range (from the Sierra Nevada to the Algarve Barrocal).
Broadleaved, soft-leaved Atlantic taxa of the class Carpino-Fagetea sylvaticae, including species of the genera Acer, deciduous Quercus, and Sorbus, also reached this territory from the north when the temperate macrobioclimate extended to lower latitudes (Alarcón et al. 2013; Vargas 2020), along with various Ericaceae species. The nanophanerophytic scrubland vegetation, represented by the alliance Asparago albi-Rhamnion oleoidis (order Pistacio lentisci-Rhamnetalia alaterni), contains a diversity of xeric paleotropical shrubs (Olea, Pistacia, Rhamnus, Myrtus, Asparagus), which endured the climatic shift from tropical to Mediterranean during the Miocene (Guzmán et al. 2009; Fiz-Palacios and Valcárcel 2013; López-Villalta 2014; Coello et al. 2021; Martín-Hernanz et al. 2023). These species now persist as either stable communities or as successional stages in low-rainfall, warm territories.
This region has also served as a significant area for speciation from ancient genotypes distinct from those in non-coastal western Iberia (e.g., Stauracanthus spp.). Various floristic migration routes have enriched the gene pool here, particularly two main coastal routes: an ascending route over mobile, halophilic dune substrates facilitating Mediterranean taxa migration, and a descending sublittoral route supporting Atlantic taxa displacement. Additionally, this region shows evidence of contributions from a North African migratory route, evidenced by the presence of Iberian-Mauritanian taxa (Suc 1984; Palamarev 1989; Costa et al. 1998; Verdú et al. 2003; Jiménez-Moreno et al. 2010; Suc et al. 2018).
Conclusions
We present an updated vegetation classification of psammophilous pine forests in western and southwestern Iberian Peninsula, identifying three newly described syntaxa dominated by sand dune-exclusive plants. Each of the four associations analysed in this study is distinct, with a unique floristic composition. Our findings indicate a clear distinction between the mesomediterranean community and the thermomediterranean community group, influenced by the biogeographic boundary between the Eurosiberian and Mediterranean regions, which lies between Aveiro and Lisbon. This diffuse boundary, without biogeographic barriers, allows the convergence of flora and vegetation from both regions. Consequently, the thermomediterranean plant communities contain a high number of taxa with predominantly Mediterranean distributions, particularly those native to southwestern Iberian sand habitats. The floristic assemblages, often hosting taxa previously considered diagnostic of other habitats – many of which are protected – highlights the conservation importance of these coastal Atlantic pine forests.
Syntaxonomic scheme
Class Pinetea halepensis Bonari et Chytrý in Bonari et al. 2021
Order Pinetalia halepensis Biondi, Blasi, Galdenzi, Pesaresi et Vagge 2014
Alliance Coremato albi-Pinion pinastri J.C. Costa, Neto, Capelo, Aguiar, Monteiro Henriques et Bonari in Bonari et al. 2021
Association Cytiso grandiflori-Pinetum pinastri J.C. Costa, Monteiro-Henriques et Neto ass. nova
Association Pinetum pinastro-pineae J.C. Costa et Neto in Bonari et al. 2021
Association Aristolochio baeticae-Pinetum pineae J.C. Costa, Neto et A.R. Pina ass. nova
Association Ulici australis-Pinetum pineae J.C. Costa et A.R. Pina ass. nova
Data availability
The data used in this article are accessible through the CircumMed Forest Database (GIVD Code: EU-00026).
Author contributions
JCC, CN, and TMH designed and conducted the survey; JCC and CN identified the plant specimens. CN, GB, and JCC wrote the manuscript. All authors contributed to the manuscript and approved the final submission.
Acknowledgements
This work was funded by the ‘Natura 2000 Habitat Mapping Project’ (CART-PG RN2000 - POSEUR-03-2215-FC-000005). GB was funded under the National Recovery and Resilience Plan (NRRP), Mission 4 Component 2 Investment 1.4 – Call for tender No. 3138 of 16 December 2021, rectified by Decree n.3175 of 18 December 2021 of Italian Ministry of University and Research funded by the European Union – NextGenerationEU; Award Number: Project code CN_00000033, Concession Decree No. 1034 of 17 June 2022 adopted by the Italian Ministry of University and Research, CUP B63C22000650007, Project title “National Biodiversity Future Center - NBFC”.
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Phytosociological tables of the Atlantic dune pine forests in the southwest of the Iberian Peninsula
docx
We report the phytosociological tables of the Atlantic dune pine forests in the Southwest of the Iberian Peninsula; table S1 = Cytiso grandiflori-Pinetum pinastri; table S2 = Pinetum pinastro-pineae; table S3 = Aristolochio baeticae-Pinetum pineae; table S4 = Ulici australis-Pinetum pineae; table S5 = Coordinates of the original vegetation plots; table S6 = Synthetic table of the four pine forest communities.
https://binary.pensoft.net/file/1362499This dataset is made available under the Open Database License (http://opendatacommons.org/licenses/odbl/1.0). The Open Database License (ODbL) is a license agreement intended to allow users to freely share, modify, and use this Dataset while maintaining this same freedom for others, provided that the original source and author(s) are credited.José Carlos Costa, Carlos Neto, Tiago Monteiro-Henriques, Ana Rita Pina, Carlos Aguiar, Francesco Mascia, Gianmaria Bonari