ARTICLES

Mediterranean Botany
ISSNe 2603-9109

https://doi.org/10.5209/mbot.78628

Synanthropisation of coastal vegetation in southern Spain

Álvaro Enríquez de Salamanca1

Received: 2 November 2021 / Accepted: 18 May 2022 / Published online: 16 January 2023

Abstract. Coastal dunes have a high ecological value but are often damaged or overexploited by beach tourism. The main
problems for the vegetation of this ecosystem are the physical destruction and the synanthropisation, the latter due to
colonisation by alien and apophyte species favoured by human action. This study determines the synanthropisation of the
vegetation of a stretch of coastline in southern Spain, based on the floristic analysis of six habitats: upper beach, foredune,
mid-dune, back-dune, creek and rocky area. The percentage of synanthropic species was 51%; 33% were apophytes, mainly
ruderal species and weeds, and 18% were alien species, mostly from gardens adjacent to the coastal zone. The degree of
synanthropisation varied for each habitat, with a marked increase from the beach inland: nil on the beach and up to 63% on the
back dune. Richness and diversity were higher in habitats with greater synanthropisation, but at the cost of less naturalness.
Land protection has ensured the physical conservation of the vegetation but is not preventing increased colonization by alien
species, some of them with high invasive potential. Measures must be taken to protect coastal vegetation from destruction
and synanthropisation.

Keywords: synanthropisation, alien plants, ruderal plants, native plants, coastal vegetation, vegetation protection.

How to cite: Enríquez de Salamanca, Á. 2022. Synanthropisation of coastal vegetation in southern Spain. Mediterr. Bot. 44,
e78628. https://doi.org/10.5209/mbot.78628.

Introduction sometimes invasive, and of apophytes, native species

not specific to the colonised habitats but favoured by
Coastal dunes are ecosystems of high ecological value human action and by his tendency to expand (Faliński,
but fragile and often overexploited by beach tourism 1975; Kornaś, 1982; Olaczek, 1982).
(Carboni et al., 2009; Tordoni et al., 2018; Kuroda Alien species are less functionally diverse than native
& Sawada, 2019; Sarmati et al., 2019). An indicator species. Among them, species that are invasive have a
of their value is the inclusion of most coastal habitats capacity to spread, reducing the physical space left for
in the European Union (EU) Directive 92/43/EEC, native species and driving the plant community towards
as EU interest habitats and in the European Red List functional homogenisation, favoured by changes
of Habitats, such as the Mediterranean coastal dune in climatology, such as rising temperatures already
grassland (B1.4b), endangered and present in southern occurring in the Mediterranean; as a consequence,
Spain (Janssen et al., 2016). synanthropic species proliferate, and there is a loss
The main causes of coastal vegetation disturbance of native species (Carboni et al., 2010; Tordoni et al.,
are the construction of houses, hotels, parking lots 2019; Del Vecchio et al., 2021).
and tracks, trampling by visitors and vehicular traffic The synathropisation of coastal vegetation is leading
(Hylgaard & Liddle, 1981; Andersen, 1995; Kutiel et to a progressive loss of habitats with good conservation
al., 2000; Lemauviel & Rozé, 2003; Martínez et al., status. For example, García-Mora et al. (2001) found
2004; Santoro et al., 2012; Malavasi et al., 2013; Šilc et that only 13% of dunes sampled in the southwest of the
al., 2017; Kuroda & Sawada, 2019; Giulio et al., 2020; Iberian Peninsula had a low vulnerability and Kutiel
Zhang et al., 2020). Significant consequences of these (2001) reported that 17% of coastal dunes had good or
threats are the physical destruction of vegetation and reasonable ecological status in Israel. Synathropisation
synanthropisation; the greater the accessibility of the of coastal habitats is a global problem, reported, for
area and the fragmentation of vegetation, the greater the example, in Australia (Batianoff & Franks, 2000),
degradation and synanthropisation (Muñoz et al., 2011; Europe (Giulio et al., 2021), Italy (e.g. Acosta et
Farris et al., 2013). al., 2007; Carboni et al., 2010; Ciccarelli, 2014; Del
Synathropisation is a process of alteration of the Vecchio et al., 2015), Spain (e.g. Campos et al., 2004;
vegetation as a consequence of direct or indirect human Asensi et al., 2014) or Ukraine (e.g. Dubyna et al., 2010;
impacts, which favors the colonisation of alien species, Kolomiichuk & Maltseva, 2014).

1
Biodiversity, Ecology and Evolution Department, Faculty of Biological Sciences, University Complutense of Madrid. E-28040 Madrid, Spain.
Email: alvenriq@ucm.es
Mediterranean Botany 44, e78628, 2023 1
2 Enríquez de Salamanca, Á. Mediterranean Botany 44, e78628, 2023

In Spain, vegetation protection against physical determine whether the current protection is enough for
destruction (such as urban planning constraints) has the conservation of coastal habitats.
improved greatly, although some threats remain, such
as the construction of access paths and beach bars or
the mechanical cleaning of beaches. However, there Materials and methods
is no control over synathropisation, which leads to
progressive degradation of natural habitats, and even the Study area
disappearance of some species; sometimes the degree
of synanthropisation is so high that coastal habitats are The study area is located in the municipality of Tarifa,
barely recognizable. As Sperandii et al. (2020) point out in the province of Cadiz (Andalusia), in southern
for Italy, nominal protection is not effective if it is not Spain. It is a stretch of the Atlantic coast, 23 km north
accompanied by effective management, conservation of Tarifa Cape, in the Strait of Gibraltar (Figure 1,
and monitoring measures. left). The site comprises an area covered by natural
In the study area, a stretch of coastline in southern coastal vegetation between the bare ground of the
Spain, despite anthropic pressures (especially urban beach and the existing buildings inland (Figure 1,
development and beach tourism), there is still a strip right). The elevation ranges between 3.5 and 10 m
of dune system preserved by urban constraints, where asl. The length of the site is 1,915 m, with an area
natural vegetation is apparently well preserved. However, of 214,817 m², resulting in an average width of 112 m.
the actual conservation status is unknown. The aim of The coordinates of the extremes (in ETRS89 datum)
this study is to analyze this strip of coastal vegetation by are north end 36°7’10.31’’N, 5°49’59.18’’W (UTM
assessing the degree of synathropisation of the different 245013, 4000924); and south end 36°6’13.97’’N,
habitats, from the upper beach to the back dune, to 5°49’27.73’’W (UTM 245749, 3999165).

Figure 1. Study area. Left: Location. Right: Delimitation on orthophoto.

The climate of the area is Mediterranean, with an been considered in this study, defined according to broadly
average annual temperature of 18.6°C, stable year-round established criteria, and also according to the possibility
(average maximum 21.6°C, average minimum 15.4°C) of their precise physical delimitation on the ground. The
and annual rainfall of 523 mm, with a marked period of habitats considered were:
summer drought, with very little or almost no rainfall
between June and August (8 mm on average in the three  Upper beach. Narrow strip on the upper beach with
months). widely scattered plants between the bare ground
of the beach and the foredune. This area should be
Habitats much larger under natural conditions, but tourist
pressure on the beach, both from visitors and
Coastal dunes can be divided into different zones according beach cleaning, has reduced it to a small, heavily
to their geomorphology and vegetation. Six habitats have endangered strip.
Enríquez de Salamanca, Á. Mediterranean Botany 44, e78628, 2023 3

 Foredune. Dune partially stabilized by vegetation, the introduction path, using in the latter case three
although with mobility on the front bordering the categories: (i) Gardens: ornamental species planted
upper beach. The substrate is loose beach sand. This in gardens close to the study area, which escape and
unit includes the embryonic dune and the mobile dune, become naturalised; (ii) Crops: species currently or
but it is not possible to delimit them precisely on the formerly cultivated, which have become naturalised in
ground. Some characteristic plants may occur in the the area; (iii) Indeterminate: alien species whose origin
embryonic dune but with irregular distribution; the in this area is uncertain, often accidentally introduced or
floristic composition is often indistinguishable from the expanded from distant or past introductions.
foredune. We consider alien plants to be naturalised in a
 Mid-dune or transition dune. Flat area between the conventional meaning, according to Richardson et
foredune and the back-dune, with transitional vegetation. al. (2000), i.e. when they reproduce and maintain
The substrate is more compact than the foredune, with populations without direct human intervention but do
incipient soil formation. not necessarily invade natural vegetation. Pyšek et al.,
 Back-dune or fixed dune. The presence of coastal (2004) propose a minimum of 10 years without direct
scrub and a lesser presence of psammophilous plants human intervention to consider a plant naturalised, but
mainly characterises it. The substrate is still beach this is not an objective criterion; for example, Pontaderia
sand but consolidated, with soil formation and little crassipes began to invade in Spain a few years after its
wind mobility of edaphic materials. introduction without reaching a decade. Invasive plants
 Small creeks cross the area, draining both natural runoff are naturalised plants that have the potential to spread
and drainage from gardens and built-up areas. They are over a large area (Pyšek et al., 2004).
visible on the back and mid-dune but disappear before
reaching the beach.
 Contact between rocky areas and foredune. At the Data collection
southern end of the study area, the mid and back dune
disappear, and the foredune comes into contact with Strip transects were used to sample the area (Burnham et
a rocky slope; it is not a cliff, but a steep slope with al., 1980; Melville & Welsh, 2001). For the main transects,
coastal scrub and rocky outcrops. a crossed design was used (Buckland et al., 2007), with
perpendicular strips running NW-SE direction (parallel
Plant classification to the sea) and NE-SW (from the sea inland). Sixteen
transects were conducted from the beach inland (or vice
Plant nomenclature has followed the International Plant versa), and seven transects parallel to the sea, one along
Names Index (IPNI, 2021). The main groups of Kornaś the upper beach, two along the foredune, two along the
(1990), widely accepted, were used to classify the plants mid-dune and two along the back-dune. A transect was
(Table 1); the lower categories, based on vague criteria also conducted along each of the four creeks detected.
according to Pyšek (1995) and difficult to delimit, were Finally, two more transects were carried out in the rocky
not applied. areas at the southern end. The objective was to achieve
the maximum sampling density, so no truncation in the
width of the strips was established; however, a detection
Table 1. Plant classification according to the origin and
human action. Source: Kornaś (1990).
width on each side of the transect axis (w) of 2 m for
herbaceous plants, 4 m for low shrubs and up to 10 m
Present in the natural coastal
vegetation without the for shrubs and trees was estimated. Consequently, the
Native concurrence of human action, estimated sampling density was 100% for tree and shrub
although occasionally may be species, and at least 50% for low bushes and 25% for
favoured by it. herbaceous vegetation.
Present in the natural coastal The boundaries between habitats were delimited
vegetation thanks to human during the walks, using orthophotos and GPS. This
Synanthropic
action, with intentional or
unintentional introduction. information was incorporated into a geographic
Native to the region but not in information system (GIS) to map the habitats and
the natural coastal vegetation, calculate the area, length and average width of each
Apophytes
where their presence is due to unit.
human action. All species observed along transects were
Not native to the region, intro- recorded, although detailed relevés were carried out
duced voluntarily or involun- at each habitat change in transects perpendicular to
Antropophytes
tarily, sometimes escaped from
(aliens)
cultivation and eventually natu- the sea and every 200 m on transects parallel to the
ralized. sea. Shorter transects in creeks and rocky areas were
Archaeophyta Introduced before 1500 A.D. treated as a single relevé. For each species detected,
the abundance and habitat in which it occurred were
Kenophyta Introduced after 1500 A.D. recorded. Plant abundance was established according
to the Braun-Blanquet (1932) scale. The overall
Within alien species, we have differentiated the abundance value for each species per habitat was
region of origin (continent) firstly, and secondly, calculated by averaging all observations.
4 Enríquez de Salamanca, Á. Mediterranean Botany 44, e78628, 2023

Data analysis Two scenarios were analysed, the current floristic

composition of the habitats (native and synanthropic
Once the floristic composition of the habitats was species, both apophyte and alien) and the composition
established, several analyses were carried out. excluding the synanthropic species (native flora only).
Firstly, a category was assigned to each of the species The results have been represented by dendrograms and
recorded: native (N), apophyte (Ap) or alien (Al). The scatter plots, and their adequacy has also been calculated
synanthropisation index (Jackowiak, 1990; Chmiel, using the cophenetic correlation coefficient and the
1993) is the ratio of apophyte and alien species to the Kruskal’s stress.
total number of established species: IS = (Ap + Al) / (N Finally, we calculated several indices of richness,
+ Ap + Al). It has been calculated for the whole flora, diversity and evenness, in the two previous scenarios,
and for each habitat. This index has been widely used, native and synanthropic species and only native
among others, in several studies on coastal vegetation species. We recorded the number of species (S) and
(e.g. Dubyna et al., 2010; Kolomiichuk & Maltseva, calculated the indices of Shannon and Weaver (1949)
2014; Valcheva et al., 2019). –applying neperian logarithms–, Pielou (1966) and
The proportion of alien species was also calculated, Simpson (1949). Table 3 shows the criteria used for the
named by some authors as anthropophytization index interpretation of these indices.
(Dąbkowska & Sygulska, 2012; Jaźwa & Stadnicka- All calculations were performed using PRIMER
Futoma, 2015; Ziaja & Wójcik, 2015): IAN = Al / 7 software (PRIMER-e, Quest Research Limited,
(N + Ap + Al). For alien species, we also analysed Auckland).
their origin, introduction path, life form (following
Raunkiaer, 1934), category (archaeophytes or Table 3. Interpretation of indices
kenophytes) and family. Index Interpretation
Habitats were subjected to a hierarchical clustering
0-2 Low diversity
(HC) and a non-metric Multidimensional Scaling Shannon-Weaver
2-3.5 Average diversity
(nMDS), using the Bray-Curtis (1957) index, according (neperian log)
>3.5 High diversity
to the proportion of native, apophyte and alien species.
0 Minimum evenness
The results were represented by a dendrogram in the HC Pielou’s evenness
1 Maximum evenness
and a scatter plot in the nMDS. To assess the adequacy
0 Maximum diversity
of the results, we calculated the cophenetic correlation Simpson
1 Minimum diversity
coefficient (Sokal & Rohlf, 1962) of the HC (the closer
to 100% the tighter the HC) and the stress for the nMDS,
using the formula number 1 of Kruskal (1964), whose
results were assessed according to the author’s proposal Results
(Table 2).
Habitat distribution and extent
Table 2. Evaluation of Kruskal’s stress.
Based on the results of the fieldwork, a GIS map of habitat
Stress Kruskal’s definition distribution has been produced (Figure 2). For each habitat
>20% Unlikely to be of interest the area and percentage of occupancy with respect to the
15-20% We must still be cautious total study area was calculated. Taking into account the
10-15% We wish it were better area of each habitat and its length (parallel to the sea except
5-10% Satisfactory in watercourses, where the current water line is measured),
its average width has been calculated (Table 4).
<5% Impressive
The most extensive habitat is the fixed dune,
followed by the foredune and then the transition zone
Changes in synanthropisation related to the
(Table 4). The rest of the habitats have a restricted
distance to the beach were analysed for the four
presence in the area.
habitats distributed continuously along the study area
parallel to the sea (upper beach, foredune, mid-dune
Table 4. Characteristics of the habitats
and back-dune).
Habitats were analysed according to their floristic Altitude Area Percentage Average
Habitat
(m asl) (ha) (%) width (m)
composition, also performing HCs and nMDSs. Two
different indices were used in this case, the Jaccard Upper beach 3.5-4.0 0.327 1.52 1.85
(1908) similarity index, which relates the number of Foredune 4.0-5.0 6.830 31.79 37.26
exclusive species to the number of shared species, Mid-dune 4.5-5.5 5.069 23.60 27.90
and the Bray-Curtis (1957) dissimilarity index, which Back-dune 5.0-8.0 8.713 40.56 58.40
measures the total difference in species abundance Creek 4.0-8.0 0.272 1.27 6.50
between two sites, divided by the total abundance at each
Rocky area 5.0-10.0 0.271 1.26 10.15
site; the former only takes into account the presence or
absence of species, and the latter also their abundance. Total 3.5-10.0 21.482 100.00 112.18
Enríquez de Salamanca, Á. Mediterranean Botany 44, e78628, 2023 5

Figure 2. Distribution map of the habitats.

Floristic composition of the habitats Africa (41%), mainly Southern Africa and the Americas
(38%), both South and Central-North America, and to a
As a result of the sampling, 149 plants have been recorded lesser extent Asia (14%) and Oceania (7%).
in the study area. The species have been compiled in a Most of the alien species (59%) came from nearby
table, including their name, category (N, Ap, Al), habitat gardens, 33% were accidentally introduced or planted in
and abundance (Table S1). the past and naturalised, and 2 species (8%) came from
former cultivation, Ficus carica, traditionally planted
Synanthropisation of the flora for its figs, and Opuntia maxima, introduced in the past
for cochineal production. Among the 27 alien species
Native species account for 49% of the total (Figure 3). found, there are 19 whose presence in the vicinity of
These plants are highly adapted to coastal environments the study area is limited to the gardens bordering the
(e.g. Achillea maritima, Cakile maritima, Calamagrostis coastal dune, while there are another eight species that
arenaria, Calystegia soldanella, Crithmum maritimum, are not associated with these gardens. Considering that
Crucianella maritima, Eryngium maritimum, Lotus the gardens were planted from 2000 onwards, 70% of
creticus, Marcus-kochia littorea, Ononis variegata, the current alien plants have entered the area in the last
Pancratium matimum or Sporobolus pungens), with two decades.
limited or no presence outside these habitats. Some Kenophytes predominate. Only Ficus carica is
native species are favoured by synanthropisation, such clearly an archaeophyte, perhaps the first plant to be
as Echium gaditanum. domesticated in the Neolithic (Kislev et al., 2006);
Synanthropic plants accounted for 51% of the total. Arundo donax could also be a kenophyte, as the first
Most of them are apophytes (33%), ruderal weeds references in the western Mediterranean date back to
introduced from nearby crops and as a consequence the 16th century (Sanz et al., 2004). The latter species
of disturbance caused by urban development and the is also the only one that is clearly invasive and also
opening of pedestrian paths. Although the study area a transformer (Richardson et al., 2000). Although
currently borders on built-up areas, these were developed naturalised and common, Agave americana and
mainly from the late 1990s onwards on former arable Opuntia maxima are not invasive in the study area;
land. Acacia saligna is locally abundant, but only in wet
There were 27 alien plants, and they accounted for areas; Lantana×strigocamara and Yucca aloifolia
18%, all of them naturalised, although frequently only are recent colonisers, which appear to be expanding;
with few (or just one) individuals and limited expansion. two species (Aizoon pubescens and Nicotiana glauca)
Alien species were distributed among 19 families, with are still sparse, but have great invasive potential;
a maximum of 3 species per family (Asteraceae and Carpobrotus acinaciformis is currently very sporadic,
Poaceae). Within life forms (classified according to but eradication programmes have been carried out in
Raunkiaer, 1934), phanerophytes dominated (55.6%), this coastal stretch (Andreu et al., 2010).
followed by hemicryptophytes and chamaephytes
(14.8% each), therophytes (11.1%) and geophytes Synanthropisation of the habitats
(3.7%). Some 26% of the species were succulent.
Asensi et al. (2016) identified 26 alien species on the The composition of vegetation in each habitat was analysed
southern coast of Spain (including the study area); 12 of to determine the degree of synanthropisation (Table 6).
them have also been recorded in our study, but 15 other The upper beach was free of synanthropic species. It
species located by us had not been cited in that work. was followed by the foredune and rocky areas, although
The most frequent areas of origin of alien species were colonisation by some apophytes and alien species was
6 Enríquez de Salamanca, Á. Mediterranean Botany 44, e78628, 2023

detected. In the mid-dune native species were reduced invasive alien species, and Avena sterilis, an apophyte.
by about half and in the creeks, even more. Finally, in the Among the frequent species (3 o greater in Braun-Blanquet
back-dune native species account for just over a third, the scale), 40% were native (Echium gaditanum, Equisteum
rest being apophytes and alien species. ramosisimum, Festuca alopecuros, Marcus-kochia littorea,
The vegetated area on the upper beach occupies a narrow Olea europaea subsp. europaea, Pycnocomon rutifolium,
strip at the foot of the foredune, with very low vegetation Scirpoides holoschoenus and Verbascum giganteum
cover. The only frequent species were Ononis variegata and subsp. martinezii) and 60% apophytes (Bromus rigidus,
Silene nicaeensis, the rest being sporadic. No synanthropic Cynara cardunculus, Daucus carota, Dittrichia viscosa,
species were detected. The vegetation in this area is affected Festuca ambigua, Lavatera cretica, Plantago bellardii,
by trampling, beach cleaning and the construction of beach P. coronopus, Scolymus hispanicus, Silybum marianum,
bars and terraces. A characteristic species of this habitat, Sisymbrium officinale and Sonchus tenerrimus). The less
Cakile maritima, frequent in the area a few decades ago frequent species were almost equally divided between
according to our observations, is now very rare. native and synanthropic, while in sporadic species, 32%

Table 6. Synanthropisation of habitats.

Synanthropic
Native Total Total
Habitat Apophytes Alien
synanthropic
Nº % Nº % Nº % Nº % Nº
Upper beach 11 100.0 - - - - - - 11
Foredune 27 79.4 5 14.7 2 5.9 7 20.6 34
Mid-dune 36 54.5 24 36.4 6 9.1 30 45.5 66
Back-dune 41 37.6 44 40.4 24 22.0 68 62.4 109
Creek 7 41.2 7 41.2 3 17.6 10 58.8 17
Rocky areas 21 72.4 3 10.4 5 17.2 8 27.6 29

In the foredune 34 species were found, mostly native were native, 37% apophytes and 31% alien. The abundance
(79.4%). The dominant species were Calamagrostis arenaria, of alien plants, although scattered, showed that this habitat
Lotus creticus, Orobanche densiflora and Pancratium was the main introductory path from nearby gardens.
matimum, followed by Achillea maritima, Crucianella The dominant species in the creeks was Arundo
maritima, Eryngium maritimum, Marcus-kochia littorea and donax, an invasive alien. It was followed in importance
Silene nicaeensis. Less frequent were Calystegia soldanella, by two native species Rubus ulmifolius and Scirpoides
Cyperus capitatus, Echium gaditanum, Medicago marina, holoschoenus. Scarcer were some native (Tamarix africana,
Ononis variegata, Pycnocomon rutifolium and Sporobolus Typha domingensis), alien (Acacia saligna, Xanthium
pungens. All dominant species were native, with synanthropic strumarium) and apophyte species (Rumex crispus).
species occurring sporadically. Among the sporadic species, synanthropic dominated
In the mid-dune native species were reduced (54.5%), (67%) compared to native (33%).
with synanthropic species gaining weight (45.5%), especially The rocky areas were dominated by bushes of Pistacia
apophytes. The vegetation was dominated by 3 native species lentiscus and Rhamnus alaternus, and the contact with
(although favoured by human action), Echium gaditanum, the dune by grasslands with Festuca alopecuros and
Pycnocomon rutifolium and Verbascum giganteum subsp. Pycnocomon rutifolium, all of them native species.
martinezii, and two synanthropic species, Plantago bellardii Among the less frequent and sporadic species, the native
and Scolymus hispanicus; all of them dominated mainly dominated (68%), especially those typical of foredunes and
around the paths and trampled areas. A second group of bushes, compared to the synanthropic species (32%); there
abundant species was dominated by foredune plants such was a significant percentage of alien species with sporadic
as Calamagrostis arenaria, Festuca alopecuros, Lotus presence.
creticus, Marcus-kochia littorea, Orobanche densiflora, Considering the percentages of native, apophyte and
Pancratium matimum and Silene nicaeensis as well as other alien plants, a HC and a nMDS were performed, using the
elements such as Lagurus ovatus and Paronychia argentea. Bray-Curtis index (Figure 3). The cophenetic correlation
Three synanthropic species, Plantago coronopus, Dittrichia coefficient for the HC was 79.28%, quite consistent, and
viscosa and Festuca ambigua, and one alien plant, Arundo the value of Kruskal’s stress for the nMDS was 1%, very
donax, were frequent. Among the sporadic species, the satisfactory. Two clearly differentiated groups can be
synanthropic dominated (58%). observed, one covering the upper beach, foredune and
On the back-dune, the proportion of native species rocky areas, with lower synanthropisation, and the other
decreased to 37.6% compared to 62.4% of synanthropic covering the mid and back dune and the creeks, with
species, with a significant weight of alien plants (22%). greater synanthropisation.
The dominant species were Pistacia lentiscus, native In the four habitats occurring continuously along
and main coastal scrub-forming plant; Arundo donax, an the study area, distributed parallel to the sea (upper
Enríquez de Salamanca, Á. Mediterranean Botany 44, e78628, 2023 7

beach, foredune, mid-dune and back-dune), an inverse beach inland the number of native species decreased,
distribution of native versus synanthropic species was while the number of synanthropic species increased
observed from the beach inland: moving away from the (Figure 4).

Figure 3. Similarity according to plant categories (native, apophyte and alien) using Bray-Curtis index.
Left: Dendrogram. Right: Scatter plot graph. Ccc - Cophenetic correlation coefficient; Ks - Kruskal’s stress.

Figure 4. Synanthropisation of the main habitats. Left: Percentages of species per category and habitat.
Right: Changes of plant categories related to beach distance.

In order to determine the floristic similarity of the The upper beach is closely related to the foredune
habitats, HCs and nMDSs were performed using the Jaccard when all species were considered, but separated when
and Bray-Curtis indices in two scenarios, the current floristic only native species were considered. The mid and
composition and the composition excluding synanthropic back-dune were closely related when all species were
species (Figure 5). The Kruskal stress value in all cases was considered; when only native plants were analysed this
1%, very satisfactory. The cophenetic correlation coefficients situation persisted with the Bray-Curtis index, but not
for the HCs were in all cases consistent, although higher with Jaccard index, where the mid dune was closer to
when only native flora was analysed: 85.57% for Jaccard the foredune. The rocky areas occupied an independent
index and the entire flora; 91.75% for Jaccard index and position, close to the dune system.
native flora; 89.75% for Bray-Curtis index and the whole Richness, diversity and evenness indices were
flora; and 91.08% for Bray-Curtis index and native flora. calculated for each habitat, considering all species
The results showed differences in the distance currently present and only native species (Table 7). The
between habitats, derived from the consideration or not back-dune presented high species richness, diversity and
of species abundance and the consideration of the whole evenness, as many species had scarce presence. It was
flora or only native species. Creeks appeared separated followed in richness and diversity by mid-dune. The
from the rest of the habitats in all cases; they have foredune and rocky areas had somewhat lower diversity
moisture-dependent azonal vegetation unrelated to the and evenness. The upper beach and creeks had the
rest of the other plant communities. lowest diversity and evenness.

Table 7. Diversity, richness and evenness indices.

Habitat Species Species Pielou Shannon Simpson
Upper beach Native + synanthropic 11 0.79 1.88 0.23
Native 11 0.79 1.88 0.23
Foredune Native + synanthropic 34 0.84 3.05 0.06
Native 27 0.88 2.91 0.07
8 Enríquez de Salamanca, Á. Mediterranean Botany 44, e78628, 2023

Habitat Species Species Pielou Shannon Simpson

Mid-dune Native + synanthropic 66 0.90 3.76 0.03
Native 36 0.90 3.24 0.05
Back-dune Native + synanthropic 109 0.90 4.23 0.02
Native 41 0.90 3.29 0.05
Creek Native + synanthropic 17 0.83 2.35 0.12
Native 7 0.79 1.54 0.26
Rocky areas Native + synanthropic 29 0.87 2.95 0.07
Native 21 0.86 2.62 0.09

(a) Jaccard index. Native + synanthropic species.

(b) Jaccard index. Native species.

(c) Bray-Curtis index. Native + synanthropic species.

Enríquez de Salamanca, Á. Mediterranean Botany 44, e78628, 2023 9

(d) Bray-Curtis index. Native species.

Figure 5. Floristic similarity with and without synanthropic species. Left: Dendrograms.
Right: Scatter plot graphs. Ccc - Cophenetic correlation coefficient; Ks - Kruskal’s stress.

Considering only native species, richness was coastal areas of northern Spain, 56% of the species come
reduced, especially in the most synanthropic habitats: from America and only 7% from Africa (Campos et al.,
in the back dune, diversity was reduced by almost two- 2004), and on the Italian coast, American species also
thirds and in the mid-dune by nearly half. In the rocky dominate (Acosta et al., 2007; Del Vecchio et al., 2015).
areas, diversity was somewhat reduced by the presence However, the study area, only 30 km from the African
of a significant number of alien species. In the creeks, coast, has a drier and warmer climate than northern
richness and diversity were drastically reduced. In Spain or most Italian coasts. This could justify the high
the foredune, the change was slight, and on the upper percentage of African species recorded here (41%).
beach, the situation did not change because there was no Giulio et al. (2021) indicate that alien plants from Africa
synanthropisation. might have a slightly higher invasion success in coastal
dunes in Europe, although species from North America
and the Mediterranean-Turanian region dominate at the
Discussion European level (Giulio et al., 2020).
The main paths of the introduction of alien plants
Our results showed a synanthropisation rate in the area in Spain are gardening (48%), agriculture (18%),
of 51%, which varies between habitats, with a null unintentional introduction and weeds (31%) and forestry
value on the upper beach and 63% on the back dune. In (3%) (Sanz et al., 2004). In northern Spain, these
coastal areas of Ukraine, Dubyna et al. (2010) obtained percentages are quite similar (Campos & Herrera, 2009),
a synanthropisation rate of 37.1% (20.9% apophytes, with a higher weight of unintentional introduction (44%)
16.2% alien), and Kolomiichuk & Maltseva (2014) 44% and a lower rate of garden plants (32%). In our study
(24.5% apophytes, 19.5% alien). Valcheva et al. (2019) area the weight of gardening is much higher (59%), and
recorded 27.1% of synanthropic species (17.4% weeds, that of agriculture lower (8%); this is explained because
9.7% alien) on the Bulgarian coast. the area borders on landscaped areas, which makes it
The percentage of alien plants reached 18%. Sobrino easier for species to escape from the gardens and become
et al. (2002) recorded 20% of alien species in coastal feral. Batianoff & Franks (2000) point to the dumping of
zones of NE Spain, a very similar value. Giulio et al. garden waste as a source of alien plant introduction in
(2020) recorded an average of 7% of kenophytes in coastal dunes, a situation that may be occurring in this
coastal dunes in Europe and Tordoni et al. (2018) a area. The vast majority of species were kenophytes, a
8.6% on the Italian Adriatic coast, while in Australia result consistent with that obtained by Del Vecchio et al.
naturalised alien species reach up to 59% of coastal (2015) in Italy.
dune flora (Batianoff & Franks, 2000). Szwed & Sýkora In the studied area, only Arundo donax competes
(1996) detected a majority of apophytes on roadside with the natural vegetation. However, there are some
verges in coastal dunes in the Netherlands. Tordoni et aggressive invasive species, which are not currently
al. (2021) highlight those anthropogenic factors, such as common, but whose populations could skyrocket if left
gross domestic product, are positively associated with unmanaged. There are also apophyte species that compete
alien species richness in coastal areas. The area studied with the natural vegetation, although not aggressively
is a popular summer resort (like most Spanish coasts), (e.g. Dittrichia viscosa, Plantago coronopus, Plantago
which implies heavy investment in urbanisation and bellardii or Scolymus hispanicus). Acosta et al. (2006)
landscaping and high tourist pressure, which explains analysed the flora of coastal dunes in Italy according
the high percentage of alien species. to functional groups, finding that invasive alien
In Spain, the dominant origin of alien plants is in species were mainly annuals, which mature quickly, or
North and South America (Sanz et al., 2004). In the perennials often with stolons or rhizomes that allow for
10 Enríquez de Salamanca, Á. Mediterranean Botany 44, e78628, 2023

rapid spread, two strategies that enable them to occupy pressure and abiotic and biotic factors is favourable for
coastal habitats. the establishment of the invader.
Asensi et al. (2016) identified 26 alien species on the In the study area, a clear differentiation of habitats
southern coast of Spain (including the study area); about with distance from the coastline was detected,
half of them have also been detected in this study, but consistent with the results obtained by Bazzichetto et al.
in addition there are 15 new species now detected. This (2016) in Italy, as well as an increase in diversity and
could be partly because they have gone unnoticed, as synanthropisation inland. The influence of wind and soil
this study is more local but more comprehensive, but properties on coastal vegetation varies between areas,
also probably (at least in some cases) because of their with the former or the latter being more important (Fenu
recent colonisation. et al., 2013; Ciccarelli, 2014). The study area is among
Anthropopressure favours synanthropisation, the windiest in the Mediterranean region, constantly
making it difficult to identify the plant communities subjected to westerly or easterly winds, which justifies
of coastal dunes (Sarmati et al., 2019). In the study the dominant influence of wind on vegetation, and the
area, it is still possible to differentiate the main natural strong inland gradient.
habitats, although the presence or even dominance of The synanthropisation of the area has led to an
some apophyte species makes the interpretation of some increase in species richness, which has doubled;
vegetation relevés difficult. approximately half of the species recorded are native and
On the upper beach and on the foredune, trampling half are synanthropic. This result is consistent with those
causes a loss of vegetation cover and richness, which recorded in other regions, such as Denmark (Brunbjerg
is even greater on the beach as a result of mechanical et al., 2015) or Italy (Prisco et al., 2016). Diversity
cleaning (Nordstrom et al., 2000) and the construction is an indicator that should be handled with caution,
of bars and terraces. However, there is no significant considering at the same time the synanthropisation of
synanthropisation. The harsh ecological conditions the vegetation. In the study area, the mid and back-dune
limit the colonisation of ruderal plants (Malavasi et were very diverse, but due to the presence of numerous
al., 2016); in the study area this is demonstrated by the species not native to these habitats, favoured by human
absence of synanthropic species on the upper beach, action.
and their scarcity on the foredune, where they are also Anthropopressure produces a synanthropisation
concentrated in the area closest to the mid-dune. In of coastal vegetation which, in this area, results in an
Italy, Acosta et al. (2009) note that the upper beach has increase in richness but also in a loss of naturalness and
a lower richness, as in this case, but at the same time, a risk of biological invasions. The fragility, richness and
the highest rarity values and the highest proportion of ecological value of coastal ecosystems, together with
endangered species. their reduced surface representation, make it necessary
Pedestrian traffic is concentrated on the main paths, to adopt measures for their conservation, which must
where wooden walkways have been constructed. take into account the problem of synanthropisation.
Outside these paths, there is no appreciable loss of Firstly, it is important to protect the entire dune
vegetation cover, partly because it is already moderate system, regardless of its species richness, as there are
on the foredune and mid-dune. Some studies indicate areas of low richness and high uniqueness, such as the
that the effects of trampling on Mediterranean dunes are upper part of the beach (Acosta et al., 2009; Kuroda &
temporary, with good potential for recovery, although Sawada, 2019). The declaration of vegetation micro-
not those of vehicle traffic (Kutiel et al., 2000; Farris et reserves – a protection figure that already exists in several
al., 2013). Rickard et al. (1994) report a high recovery regions of Spain – in well-preserved coastal areas, such
potential for pioneer dune communities in South Africa as the one studied, would be highly recommended.
but a low recovery potential for shrubs, which is also In the management of coastal dune systems, it is
applicable to this area; analysis of historical aerial important to establish vegetation recovery periods
photographs of the area showed that there has been (Lemauviel & Rozé, 2003). Fencing of dune systems
virtually no recovery of shrub cover for decades. reduces trampling, a passive recovery system that
In mid-dune, tracks were built for beach service increases vegetation cover and the richness of protected
vehicles (especially quads), taking advantage of the areas (Acosta et al., 2013). Symbolic fencing is
gentle relief, the greater compactness of the terrain and the preferable for controlling pedestrian access because it is
absence of scrub. This has increased synanthropisation, less visually intrusive and does not alter the natural wind
and the proliferation of species like Echium gaditanum, transport regime (Nordstrom et al., 2000; Grafals-Soto
Scolymus hispanicus and Arundo donax. The back & Nordstrom, 2009). Rerouting pedestrian traffic is also
dune is the area with the greatest synanthropisation and interesting (Nordstrom et al., 2000), a measure that has
presence of alien species. already been adopted in the study area by constructing
Carboni et al. (2010) indicated for Italy that the wooden pedestrian walkways. However, uncontrolled
high levels of invasion in the transition dune could be access to the beach still exists and should be regulated.
partly explained by the higher propagule pressure in Vehicle traffic should be limited to the intertidal zone to
this area. However, Bazzichetto et al. (2018) applied a avoid the destruction of vegetation and dunes (García-
complex assessment model, finding that invasion does Mora et al., 2001).
not occur homogeneously across the coastal landscape A major problem is beach cleaning and the placement
but occurs where the combined action of propagule of bars and terraces. Garcia-Mora et al. (1999) indicate
Enríquez de Salamanca, Á. Mediterranean Botany 44, e78628, 2023 11

that beach management in Spanish tourist resorts thirds on the back-dune, with a marked increase from
is oriented towards sand sieving and gardening of the beach inland. Although only one invasive species,
embryo dunes, affecting native species and favouring Arundo donax, was detected, there are several others
kenophytes. Nordstrom et al. (2000) point out the need to with high invasive potential, whose populations could
modify beach cleaning procedures. Mechanical cleaning grow rapidly if left unmanaged.
is destructive for the vegetation and should therefore Comparing the orthophotos from 2001 and 2020, it
be limited to the bare beach, with manual cleaning of can be observed that there has been no appreciable loss
the upper area. In addition, beach bars and terraces are of vegetation in those 20 years. However, the degree of
often located precisely in the upper part of the beach, synanthropisation is high, especially in the back dune,
destroying its vegetation and the embryo dunes; it would and several alien species that only grow in gardens
be desirable to relocate them inland, outside the dune bordering the coast have now also been detected in
system, or at least away from the upper part of the beach, coastal habitats. Therefore, although protection against
with lighter mobile installations. urban development has been effective in physically
The use of alien species in areas adjacent to coastal maintaining the vegetation, it is not preventing the
dunes should be restricted (Nordstrom et al., 2000), progressive entry of alien species, some of them with
especially those with proven or potential invasiveness. high invasive potential.
There are several experiences of eradication of Measures must be taken to protect coastal vegetation
invasive plants in coastal dunes, especially Carpobrotus: from destruction and synanthropisation, especially from
Buisson et al. (2021) showed successful eradication the spread of invasive species. Including these areas
results; Andreu et al. (2010) analysed their manual in a legal protection scheme, and applying specific
removal in Andalusia (including the study area), management, conservation and monitoring measures
noting that although the density was greatly reduced, would be desirable, as mere nominal protection is not
repeated uprooting treatments were required; Lazzaro effective in guaranteeing their conservation (Sperandii
et al. (2020) noted the efficacy of using glyphosate, a et al., 2020). Other desirable measures are: regulating
controversial herbicide (it is authorised in the EU until 15 access to avoid trampling; avoiding vehicle traffic;
December 2022, and an extension is under evaluation). symbolic fencing; respecting regeneration periods;
However, eradication of alien plants may also encourage avoiding mechanical cleaning of the beach and the
synanthropisation. Kim et al. (2019) noted for Korea placement of bars and terraces on the upper part of the
that invasive species removal was costly and, if not done beach; or the eradication of invasive species, carefully
very carefully, could even increase the initial invasion. assessing their advantages and disadvantages to avoid
Novoa et al. (2013) found that dunes regenerated after unwanted synanthropisation as a result of these actions.
Carpobrotus removal were occupied by opportunistic
ruderal species, which competed with native dune plants.
Eradication decisions should be based on the importance Conflict of interest
of the invasion, weighing the benefits and risks.
None.

Conclusions
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