Albufera

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Spatial and temporal variation of community
composition and species cover following dune
restoration in the Devesa de Albufera (Valencia,
Spain).
P. Zuccarini
a
, A. Aguilella
b
& G. Bedini
a
a
Department of Biology, University of Pisa, via Luca Ghini 13, I-56126Pisa, Italy
b
Botanic Garden, University of Valencia, C/Quart 80, E-46008Valencia, Spain
Accepted author version posted online: 28 Jan 2015.Published online: 24 Feb 2015.
To cite this article: P. Zuccarini, A. Aguilella & G. Bedini (2015): Spatial and temporal variation of community composition
and species cover following dune restoration in the Devesa de Albufera (Valencia, Spain)., Plant Biosystems - An
International Journal Dealing with all Aspects of Plant Biology: Official Journal of the Societa Botanica Italiana, DOI:
10.1080/11263504.2015.1012134
To link to this article: http://dx.doi.org/10.1080/11263504.2015.1012134
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ORIGINAL ARTICLE
Spatial and temporal variation of community composition and species
cover following dune restoration in the Devesa de Albufera (Valencia,
Spain).
P. ZUCCARINI1,
*, A. AGUILELLA2,
**, & G. BEDINI1
1
Department of Biology, University of Pisa, via Luca Ghini 13, I-56126 Pisa, Italy and 2
Botanic Garden, University of
Valencia, C/Quart 80, E-46008 Valencia, Spain
Abstract
Plant populations were reintroduced to the coastal dune bar of the Devesa de Albufera from 1988 to 2004; different coastline
sections received different species composition and cover. With the aim to detect spatial and temporal variation of floristic
diversity, we compared current species composition and cover across the length of the Devesa and across the dune bar with
those imposed at the time of restoration. Non-metric multidimensional scaling (NMDS) detected significant differences both
across the dune faces and across the coast sections. Differences across the dune faces reflect the sea-inland ecological gradient
and resulted from a spatial rearrangement of plant populations: Calystegia soldanella, Achillea maritima and Polygonum
maritimum prefer the windward face; Malcolmia littorea and Lagurus ovatus the leeward. Differences across coast sections are
related to those at restoration time, with a slow trend towards the homogenization of plant communities. At the current level
of anthropic pressure, the plant cover is likely to evolve following the trends pointed out in this research.
Keywords: Coastal sand dunes, dune restoration, Mediterranean sea, spatial dynamics, temporal dynamics
Introduction
The Natural Park of Albufera (Valencia, Spain) is an
important wetland area, home to more than 250
species of birds and 800 species of plants, protected
under the Ramsar Convention, the “Birds Directive”
EU 79/409, and the “Habitat Directive” EU 92/43.
The Devesa is the sand bar that separates the
Albufera lagoon from the Mediterranean Sea.
During the 1960s and 1970s of past century, the
dune bar was destroyed and replaced by a paved
promenade (Go´mez-Pina et al. 2002). After strong
protests against the local Administration, in 1986 the
Oficina Tecnica Devesa de l’Albufera (OTDA) drew
up a plan for the restoration of the dune bar.
The plan involved the removal of the promenade,
the erection of a new dune bar with sand extracted
from the backdune area or from the Valencia
harbour, and the restoration of plant communities
on the new dune bar.
After a series of pilot-experiments (Davidson-
Arnott & Law 1996; Curr et al. 2000), dune
restoration took place between 1988 and 2004 as
separate projects on different stretches of coastline
(Figure 1; Table SI), with native plants propagated by
ODTA. Plant communities were restored by planting
individuals of selected species as whole plants,
cuttings, bulbs, root balls or seedling (Benavent
Olmos et al. 2004). Because of the stepwise progress
of the restoration, of the availability of suitable plant
material, of variations in the ODTA staff and of
progressive refinement of protocols, different sections
of the Devesa received different species composition
and cover at the time of restoration. Escaray et al.
(2010) evaluated the restoration as successful based
on the measurement of ecological indices of restored
and undisturbed communities.
We started this work to answer the following
questions:
(1) Do current differences – if any at all – in species
composition and cover across the length of the
Devesa match those determined by the different
restoration protocols?
(2) Do current spatial patterns – if any at all – in
species composition and cover across the width
q 2015 Societa` Botanica Italiana
Correspondence: G. Bedini, Department of Biology, University of Pisa, via Luca Ghini 13, I-56126 Pisa, Italy. Email: gianni.bedini@unipi.it
Plant Biosystems, 2015
http://dx.doi.org/10.1080/11263504.2015.1012134
Downloadedby[Plant&FoodResearch]at19:2601March2015

of the dune bar (i.e. from windward to leeward
face) match those imposed at the time of
restoration?
The answers will shed light on the spatial and
temporal variation of community composition and
species cover following dune restoration in the
Mediterranean coast of the Valencia Region, allow
an ex post assessment of the effectiveness of
restoration protocols and enable to make predictions
on the evolution of the plant cover.
Materials and methods
Study area
The Devesa de la Albufera, about 20 km south of
Valencia, is part of a barrier beach delimited by the
Tu´ria River and Cape Cullera, and separates the
Albufera lagoon from the sea (Figure 1).
Its climate is of Mediterranean subarid type; soil
type is mostly calcareous arenosol (Escaray et al.
2010).
We considered ﬁve beaches along the Devesa de
Albufera, from North to South: El Saler (SAL), Els
Ferros-Garrofera (GAR), La Brava (BRA), La
Malladeta (MAL) and El Canyar (CAN) (Figure 1),
for which restoration data were available either for
the whole dune or for the distinct dune faces (WW,
windward face; CS, crest; LW, leeward face),
(Benavent Olmos et al., 2004; Table SI).
Species composition and cover
Repopulation data. For CAN, GAR and MAL, the
number of propagation units is given for each species
in terms of plants, cuttings, bulbs, root balls or seed
launches (Benavent Olmos et al. 2004), irrespective
of dune faces. For BRA and SAL, the number of
individuals per area unit is given for each species and
for each dune face (WW, CS, LW). To obtain a
coherent dataset, we recalculated the data for CAN,
GAR and MAL as number of individuals per area
unit with the following procedure:
(1) based on Vizcaino Matarredona (OTDA, per-
sonal communication), on Benavent Olmos
et al. (2004) and on Escaray et al. (2010), we
assumed that, at the moment of repopulation,
plant propagules of any given species were
homogeneously distributed across the beach
and, consequently, that their density was the
same in all dune faces;
(2) for each species, we calculated the number of
propagules per plot in each beach as pSB ¼ PSB x
(Q/AB), where PSB is the total number of
propagules of the species S in beach B, Q is the
plot size in m2
and AB is the area in m2
of beach
B; given the assumption in 1), we assumed pSB
to be constant in the whole beach; plot size has
been deﬁned based on repopulation modules
(25 m2;
Benavent Olmos et al. 2004);
(3) we assumed that the plant propagules used
(plants, cuttings, bulbs, root balls) were all vital
and capable of developing into a full-grown
individual, whereas for “seed launches” (golpe de
semilla in Benavent Olmos et al. (2004)), we
assumed that each launch included 50 seeds of
which only 10 would reach adult stage; the
number of individuals is summarized in Table SI
along with original data for BRA and SAL.
Figure 1. Map of the Devesa showing the position of the investigated beaches. SAL, El Saler; GAR, Els Ferros – Garrofera; BRA, La Brava;
MAL, La Malladeta; CAN, El Canyar.
2 P. Zuccarini et al.

Then, with homogeneous data for all beaches,
we calculated plant cover for each species as
follows: (1) we assigned a standard cover area DS
(m2
) to each species’ individuals based on
estimates of plant sizes (Table SI, first item of
conversion factor); (2) we calculated the
percentage cover for each species S in each
beach B and in each dune face F as CSBF ¼
pSBF £ DS/Q £ 100; given the assumptions in
(2), (3) above, CSBF was constant in the whole
beach (B ¼ GAR, MAL, CAN) or in the
separate dune faces (B ¼ SAL, BRA) (Tables Ia
and Ib). Finally, we constructed a plot-species
matrix (T0 matrix), with species in columns and
cover data in rows, (10 releve´s for each dune
face £ 3 dune faces £ 5 beaches ¼ 150 rows).
Current data. To record current species composition
and cover, we set 10 transects in each beach, spanning
the first dune perpendicularly to its axis, equidistant
from one another; the same equidistance separated
the outer transects from the side ends of the beach.
Along each transect, we set three rectangular plots (7
£ 4 m): one at the windward face of the dune (plot
WW), one at its crest (plot CS), and one at its leeward
face (plot LW), tallying 30 plots in each beach (10
WW, 10 CS and 10 LW) and 150 plots in total (50
WW, 50 CS and 50 LW). In each plot, we assessed
floristic composition and estimated visually each
species’ cover, yielding 150 releve´s. Then, we
constructed a plot-species matrix (T1 matrix) with
censused species in columns and cover data in 150
rows (one for each releve´).
Both matrices (T0 and T1) are available from the
corresponding author. Nomenclature follows
Euro þ Med (2006). Authorities are specified only
at the first mention of a species and omitted
thereafter.
Analysis of spatial variability
We subjected matrices T0 and T1 to non-metric
multidimensional scaling (NMDS) ordination, with
Bray-Curtis distance, to detect differences in species
composition and cover across beaches and across
dune faces. The environmental factors “beach”
(SAL, GAR, BRA, MAL, CAN) and “dune faces”
(WW, CS, LW) were then fitted to the ordination; the
goodness of fit was calculated as R2
¼ 1 ¼ ssf/sst,
where ssf is the sum of squares within each factor, and
sst is the total sum of squares (R software, package
vegan version 1.11-4). If the analysis showed
differences across dune faces and beaches, we
analysed separately each species’ releve´s (i.e. single
columns of matrices T0 and T1) to establish what
species were responsible for the observed difference:
because most plot data failed both the normality
(Shapiro-Wilk) and the homoscedasticity (Bartlett)
tests, we analysed the releve´s by a Friedman test (R
software, standard package). We rejected the null
hypothesis for p-value , 0.05 and submitted signifi-
cant data to a post hoc Schaich and Hamerle test
(code obtained from http://www.r-statistics.com/
Table Ia. Species composition and cover by beach at repopulation time. Figures are the number of populated plots and associated cover
percentage (in brackets).
SAL GAR BRA MAL CAN
Achillea maritima 10 [3.0] a
30 [3.6] b
10 [3.0] a
30 [6.6] b
30 [1.8] a
Ammophila arenaria subsp. arundinacea 10 [18.4] a
30 [17.6] b
10 [18.4] a
30 [6.4] ab
30 [2.4] a
Cakile maritima subsp. maritima 10 [1.0] a
0b
10 [1.0] a
0b
0b
Calystegia soldanella 20 [2.2] a
30 [2.4] b
20 [2.2] a
30 [0.6] g
30 [1.0] a
Crucianella maritima 10 [3.2] a
0b
10 [3.2] a
30 [0.1] ag
30 [2.0] g
Cyperus capitatus 20 [1.6] a
0b
20 [1.6] a
0b
0b
Echinophora spinosa 10 [3.2] ab
0a
10 [3.2] ab
30 [0.1] b
30 [6.4] g
Elytrigia juncea 10 [18.0] a
30 [9.0] a
10 [18.0] a
30 [7.2] a
30 [14.4] b
Eryngium maritimum 10 [2.4] a
30 [3.0] b
10 [2.4] a
30 [0.2] a
30 [18.6] b
Euphorbia paralias 10 [1.6] a
0b
10 [1.6] a
0b
0b
Lotus creticus 20 [4.4] a
30 [12.0] b
20 [4.4] a
30 [2.0] a
30 [9.6] b
Malcolmia littorea 10 [3.2] a
30 [3.2] b
10 [3.2] a
30 [0.4] a
30 [9.2] g
Medicago marina 20 [1.3] a
30 [3.5] b
20 [1.3] a
30 [0.5] a
30 [5.1] b
Ononis natrix 10 [3.2] a
30 [2.6] b
10 [3.2] a
30 [0.3] ab
30 [7.4] g
Pancratium maritimum 10 [1.9] a
30 [0.5] a
10 [1.9] a
30 [1.2] b
30 [0.5] a
Polygonum maritimum 10 [1.0] a
0b
10 [1.0] a
30 [0.2] g
0b
Sporobolus pungens 10 [15.0] a
0b
10 [15.0] a
0b
0b
Teucrium dunense 0a
0a
0a
0a
30 [2.2] b
Total number of species 17 10 17 13 13
Notes: SAL, El Saler; GAR, Els Ferros-Garrofera; BRA, La Brava; MAL, La Malladeta; CAN, El Canyar. All species significantly contribute
to differences between beaches (p , 0.001). Superscript Greek letters indicate differences between beaches (same letters ¼ no statistical
difference; different letters ¼ statistical differences). Differences assessed by a Friedman’s test (see material and methods for details).
Spatial and temporal variation of community composition 3

2010/02/post-hoc-analysis-for-friedmans-test-r-
code). Differences between plots were considered
significant for p-value,0.05.
For a graphical representation of selected species
distribution across the WW, CS and LW plots, we
recalculated each cover data as the percentage of the
cumulated value, as
%coveri ¼ coveri £ 100/(coverWW þ coverCS þ
coverLW), for i ¼ WW, CS, LW and coverWW þ
coverCS þ coverLW . 0) and plotted them in a
ternary plot (Trinity software)
In some calculations of cover, we have used the
average value of populated plots, i.e. those with a
cover . 0 for a single species, rather than the average
value of all plots, to allow for a better representation
of the plants’ density.
Comparison between current and repopulation data
We used the data from repopulation and current
releve´s to build two plot-species matrices of identical
size: first, we assembled an inclusive list of species,
taking into account those present in either group of
releve´s, and assigned a column of both matrix to each
species; then we added to both matrices 150 rows
and filled them with the current cover (matrix with
current data) or the repopulation cover of each
species (matrix with repopulation data). We assessed
correlation between the two datasets by a Mantel test
(Bray-Curtis distance, Pearson moment); the con-
tribution of each species to differences between the
two datasets was assessed by a pairwise Wilcoxon
test. Both tests were run with R software (“ape” and
“stats” packages, respectively).
Results
Repopulation data. At repopulation time, 18 species in
total were used in the five beaches (SAL, BRA ¼ 17;
GAR ¼ 10; MAL, CAN ¼ 13; see Table SI). Ten
species were used in all beaches, with % cover
varying between 0.20 and 18.60 (Table SI).
Current data. As regards current releve´s, we identified
23 species in total (Table IIa). The most frequent
species were Lotus creticus (130 plots), Elytrigia juncea
(123), Malcolmia littoralis (97), Calystegia soldanella
(79), Sporobolus pungens (52) and Achillea maritima
(49). Nine species were observed in less than 15
(10%) plots. The remaining species were recorded in
16–38 plots, according to an irregular distribution
pattern as shown in Figure S1(a). The number of
species per plot varied from 2 to 10 with a left-skewed
distribution (Figure S1(b)), mainly caused by the
pattern recorded at SAL, GAR, CAN (Figures S1
(c)–S1(g)).
Eight species found in the current releve´s had not
been part of the repopulation planting (Table IIa).
Repopulation data. The NMDS analysis of the
repopulation plots (Figure S2) shows that GAR,
MAL and CAN are represented by a single dot (due
Table Ib. Species composition and cover by dune face at repopulation time. Figures are the number of populated plots and associated cover
percentage (average ^ SE, in brackets).
WW CS LW
Achillea maritima*** 50 [3.60 ^ 0.23] a
30 [4.00 ^ 0.37] b
30 [4.00 ^ 0.37] b
Ammophila arenaria subsp. arundinacea*** 30 [8.80 ^ 1.19] a
50 [12.64 ^ 0.98] b
30 [8.80 ^ 1.19] a
Cakile maritima subsp. maritima*** 20 [1.00 ^ 0.00] a
0 [0.00 ^ 0.00] b
0 [0.00 ^ 0.00] b
Calystegia soldanella*** 50 [1.68 ^ 0.10] a
50 [1.68 ^ 0.10] a
30 [1.33 ^ 0.14] b
Crucianella maritima*** 20 [1.04 ^ 0.22] a
20 [1.04 ^ 0.22] a
40 [2.12 ^ 0.20] b
Cyperus capitatus*** 0 [0.00 ^ 0.00] a
20 [1.60 ^ 0.00] b
20 [1.60 ^ 0.00] b
Echinophora spinosa*** 20 [3.24 ^ 0.72] a
20 [3.24 ^ 0.72] a
40 [3.22 ^ 0.36] b
Elytrigia juncea*** 50 [13.32 ^ 0.64] a
30 [10.20 ^ 0.57] b
30 [10.20 ^ 0.57] b
Eryngium maritimum*** 30 [7.26 ^ 1.50] a
30 [7.26 ^ 1.50] a
50 [5.32 ^ 0.96] b
Euphorbia paralias*** 20 [1.60 ^ 0.00] a
0 [0.00 ^ 0.00] b
0 [0.00 ^ 0.00] b
Lotus creticus*** 50 [6.48 ^ 0.53] a
50 [6.48 ^ 0.53] a
30 [7.87 ^ 0.79] b
Malcolmia littorea*** 30 [4.27 ^ 0.68] a
30 [4.27 ^ 0.68] a
50 [3.84 ^ 0.41] b
Medicago marina*** 50 [2.34 ^ 0.25] a
50 [2.34 ^ 0.25] a
30 [3.04 ^ 0.36] b
Ononis natrix*** 30 [3.41 ^ 0.55] a
30 [3.41 ^ 0.55] a
50 [3.33 ^ 0.33] b
Pancratium maritimum*** 30 [0.72 ^ 0.06] a
30 [0.72 ^ 0.06] a
50 [1.20 ^ 0.09] b
Polygonum maritimum*** 30 [0.69 ^ 0.07] a
10 [0.16 ^ 0.00] b
10 [0.16 ^ 0.00] b
Sporobolus pungens*** 0 [0.00 ^ 0.00] a
0 [0.00 ^ 0.00] a
20 [15.00 ^ 0.00] b
Teucrium dunense 10 [2.24 ^ 0.00] 10 [2.24 ^ 0.00] 10 [2.24 ^ 0.00]
WW, windward face; CS, crest; LW, leeward face. Species in bold significantly contribute to differences between dune faces
(*** ¼ p , 0.001). Superscript Greek letters indicate differences between dune faces (same letters ¼ no statistical difference; different
letters ¼ statistical differences). Differences assessed by a Friedman’s test (see material and methods for details).

TableIIa.Currentspeciescompositionandcoverbybeach.
SALGARBRAMALCANALLBEACHES
Achilleamaritima*5[10.6067.39]a
<14[43.4367.50]b
<8[23.7566.18]ab
"10[37.5064.73]ab
<12[33.7565.61]ab
<49[33.2964.76]<
Ammophilaarenariasubsp.arundinacea1[40.00^0.00]<0#0#2[7.50^2.50]#1[50.00^0.00]#4[26.25^4.70]#
Cakilemaritimasubsp.maritima**2[7.5062.50]a
<5[2.3061.10]a
!5[10.1063.47]a
<14[9.1062.06]b
!5[8.4062.25]a
!32[7.9862.60]"
Calystegiasoldanella16[9.13^2.35]"18[3.86^1.25]<11[7.05^1.29]<17[5.29^1.03]<17[5.88^0.73]"79[6.11^2.42]"
Carpobrotusacinaciformis**5[15.6063.98]ab
!9[6.6761.67]b
!1[35.0060.00]a
!0a
<1[30.0060.00]a
!16[12.6963.25]!
Centaureaseridis**5[13.0063.00]a
!0b
<1[2.0060.00]ab
!0b
<0b
<6[11.1762.74]!
Crucianellamaritima*0a
#0a
<0a
#3[6.6761.67]b
#0a
#3[6.6761.70]#
Cyperuscapitatus**7[5.7161.92]ab
<8[1.8860.48]ab
!9[15.6165.32]ab
<11[17.1865.42]a
!1[2.0060.00]b
!36[10.7463.76]"
Echinophoraspinosa***2[1.2560.75]a
#0a
<3[3.5061.50]ab
#12[9.2163.16]b
<11[4.3261.37]b
#28[6.1162.84]#
Echiumsabulicola***8[7.3861.88]a
!0b
<0b
<0b
<1[5.0060.00]b
!9[7.1162.24]!
Elytrigiajuncea**27[43.1563.97]a
"26[38.0864.39]ag
"20[26.7563.13]bgd
"23[22.0463.22]bgd
"27[30.1963.93]ad
"123[32.6264.51]"
Eryngiummaritimum12[5.83^1.66]<8[5.38^3.59]#7[3.14^1.40]<6[6.00^2.31]<5[2.60^1.02]#38[4.84^2.49]#
Euphorbiaterracina10[5.45^1.70]!2[2.75^2.25]!6[14.33^5.75]!6[7.58^2.08]!7[9.36^4.35]!31[8.29^3.04]!
Lagurusovatussubsp.ovatus***12[6.0061.85]a
!0b
<0b
<0b
<1[0.5060.00]b
!13[5.5862.51]!
Lotuscreticus***26[27.1262.47]a
"30[24.1762.08]a
"18[19.7263.60]b
"28[33.1163.39]a
"28[30.7163.56]a
"130[27.4863.99]"
Malcolmialittorea***21[8.0561.51]a
"13[13.2362.92]a
<25[32.2065.47]b
"20[20.5064.21]ab
"18[10.2262.08]a
#97[17.9464.43]"
Medicagomarina**3[9.0063.79]ab
#7[11.4361.80]a
<2[2.7562.25]ab
#0b
#2[10.2569.75]ab
#14[9.5062.54]#
Pancratiummaritimum*11[1.3260.25]a
<9[1.0660.29]ab
#2[0.5060.00]bg
#4[0.6360.13]ag
#7[0.8660.21]ag
#33[1.0260.86]#
Paronychiaargentea1[15.00^0.00]!0<mrk[RELSP]><0<mrk[RELSP]><0<mrk[RELSP]><0<mrk[RELSP]><1[15.00^0.00]!
Polygonummaritimum0#0<mrk[RELSP]><4[15.50^4.94]<2[5.00^0.00]#2[22.50^7.50]!8[14.63^3.18]#
Salsolakali7[5.07^1.41]!4[0.50^0.00]!6[3.08^1.56]!5[6.00^1.00]!4[3.13^1.13]!26[3.79^1.82]!
Sedumsediforme*6[3.8361.33]a
!0b
<0b
<2[5.0060.00]ab
!2[17.75617.25]ab
!10[6.8563.20]!
Sporoboluspungens***17[27.7665.25]a
"9[9.4463.35]b
!12[10.7562.53]ab
<3[5.0060.00]b
!11[13.1862.55]ab
!52[16.2764.02]"
Totalnumberofspecies211417172023
SAL,ElSaler;GAR,ElsFerros-Garrofera;BRA,LaBrava;MAL,LaMalladeta;CAN,ElCanyar.Figuresarethenumberofpopulatedplotsandassociatedcoverpercentage(average^SE,in
brackets).Inbold,speciessigniﬁcantlycontributingtodifferences(*¼p,0.05;**¼p,0.01;***¼p,0.001).SuperscriptGreeklettersindicatedifferencesbetweenbeaches(sameletters¼no
statisticaldifference;differentletters¼statisticaldifferences).DifferencesassessedbyaFriedman’stest(seematerialandmethodsfordetails).Upwardarrows¼increasedcoverfromrepopulationto
current;downwardarrows¼decreasedcoverfromrepopulationtocurrent;circasign¼nostatisticaldifferencefromrepopulationtocurrent;exclamationmark¼speciesnotintroducedat
repopulation.DifferencesassessedbyWilcoxonpairedtest.

to their artificially imposed homogeneity) and are
well separated in the ordination space; SAL and BRA
are represented by three widely spaced dots,
matching the different floristic composition of the
windward, crest and leeward dune faces imposed at
repopulation time; as the two beaches were
repopulated with the same modules, the two sets of
points are coincident. Both environmental factors –
“beach” and “dune face” – affected releve´s in a
significant way (r 2
¼ 0.2109, p , 0.001 for factor
“beach”; r 2
¼ 0.3156, p , 0.001 for factor “dune
face”). The relevance of species in determining the
observed differences is shown in Table Ia (Greek
superscript letters).
Current data. As regards the current releve´s, the
number of species varied across beaches from 14 at
GAR to 21 at SAL (Figure S1(h)). Tables IIa and IIb
show species composition and cover for each beach
and dune face, respectively.
TheNMDSresultsareplottedinFigures2–3.Both
environmental factors – “beach” and “dune face” –
affected releve´s in a significant way (r 2
¼ 0.29,
p , 0.001 for factor “dune face”; r 2
¼ 0.17,
p , 0.001 for factor “beach”), but pairwise compari-
sons showed that the releve´s of beaches GAR and
CAN are not significantly different (data not shown).
The relevance of species in determining the
observed differences is shown in Tables IIa and IIb.
Some species observed in #10 plots are found
mainly in SAL: Lagurus ovatus subsp. ovatus, Echium
sabulicola, Centaurea seridis, Sedum sediforme; Crucia-
nella maritima was observed only in MAL; the
invasive Carpobrotus acinaciformis is found mainly in
SAL and GAR; the native Echinophora spinosa is
found in MAL and CAN.
The releve´s of WW plots are well separated by
those of LW plots, whereas those of CS plots sit in
between (Figure 3). This pattern is confirmed by
the post hoc tests, as shown in Table IIb: Achillea
maritima, Calystegia soldanella, Salsola kali, Cakile
maritima subsp. maritima exhibit a preference for
the windward face and the crest; on the other hand,
Malcolmia littorea, Cyperus capitatus, Echinophora
spinosa, Euphorbia terracina, Pancratium maritimum,
Echium sabulicola, Lagurus ovatus subsp. ovatus and
to a lesser extent Sporobolus pungens prefer the
crest and the leeward face; among the most
represented species, Elytrigia juncea and Lotus
creticus span the three faces without preferences.
A graphical representation of these patterns is given
in Figure 4.
Comparison between current and repopulation data
The number of species increased by 4–7 units in all
beaches except BRA from repopulation to current
Table IIb. Current species composition and cover by dune face.
WW CS LW
Achillea maritima*** 41 [34.83 6 4.75] a
" 8 [25.38 6 4.78] b
< 0b
#
Ammophila arenaria subsp. arundinacea 1 [10.00 ^ 0.00] # 3 [31.67 ^ 4.86] # 0 #
Cakile maritima subsp. maritima* 14 [5.29 6 2.12] a
< 13 [10.42 6 2.35] a
! 4 [11.50 6 3.59] b
!
Calystegia soldanella*** 44 [7.63 6 2.65] a
" 30 [4.15 6 1.78] b
< 5 [4.60 6 1.87] g
#
Carpobrotus acinaciformis 4 [20.00 ^ 3.29] ! 6 [10.50 ^ 3.50] ! 6 [10.00 ^ 2.78] !
Centaurea seridis 1 [2.00 ^ 0.00] ! 2 [12.50 ^ 3.26] ! 3 [13.33 ^ 2.40] !
Crucianella maritima 0 # 0 # 3 [6.67 ^ 1.70] #
Cyperus capitatus*** 2 [0.75 6 0.59] a
! 16 [12.31 6 3.90] bg
< 18 [10.44 6 3.73] g
<
Echinophora spinosa*** 2 [0.50 6 0.00] a
# 9 [6.72 6 2.78] a
< 17 [6.44 6 2.95] b
#
Echium sabulicola* 0a
! 5 [7.40 6 2.26] b
! 4 [6.75 6 2.38] b
!
Elymus farctus 45 [27.60 ^ 4.51] " 43 [36.98 ^ 4.34] " 35 [33.71 ^ 4.62] "
Eryngium maritimum** 6 [5.75 6 3.45] a
# 13 [4.65 6 2.31] ab
# 18 [4.92 6 2.12] b
#
Euphorbia terracina*** 3 [0.67 6 0.54] a
! 11 [6.32 6 2.58] bg
! 17 [10.91 6 3.25] g
!
Lagurus ovatus subsp. ovatus* 1 [1.00 6 0.00] a
! 5 [5.60 6 2.87] ab
! 7 [6.21 6 2.36] b
!
Lotus creticus 48 [25.25 ^ 3.75] " 45 [30.22 ^ 4.20] " 37 [27.03 ^ 3.98] "
Malcolmia littorea*** 11 [5.55 6 1.58] a
< 39 [20.72 6 4.82] b
" 47 [18.53 6 4.20] b
"
Medicago marina 3 [11.67 ^ 1.70] # 5 [11.40 ^ 2.89] # 6 [6.83 ^ 2.42] #
Pancratium maritimum*** 6 [0.75 6 0.78] a
# 7 [1.43 6 1.05] a
# 19 [0.97 6 0.77] b
#
Paronychia argentea 0 <mrk[RELSP]> < 0 <mrk[RELSP]> < 1 [15.00 ^ 0.00] !
Polygonum maritimum* 6 [13.67 6 2.96] a
< 0b
# 2 [17.50 6 4.20] ab
<
Salsola kali** 12 [3.42 6 1.71] a
! 12 [4.71 6 1.91] a
! 2 [0.50 6 0.00] b
!
Sedum sediforme 1 [5.00 ^ 0.00] ! 3 [13.00 ^ 4.36] ! 6 [4.08 ^ 1.85] !
Sporobolus pungens* 13 [6.54 6 2.14] a
! 18 [18.94 6 4.37] ab
! 21 [20.00 6 4.03] b
<
Figures are the number of populated plots and associated cover percentage (average ^ SE, in brackets). In bold, species significantly
contributing to differences (* ¼ p , 0.05; ** ¼ p , 0.01; *** ¼ p , 0.001). Superscript Greek letters indicate differences between dune
faces (same letters ¼ no statistical difference; different letters ¼ statistical differences). Differences assessed by a Friedman’s test (see
material and methods for details). Upward arrows ¼ increased cover from repopulation to current; downward arrows ¼ decreased cover
from repopulation to current; circa sign ¼ no statistical difference from repopulation to current; exclamation mark ¼ species not introduced
at repopulation. Differences assessed by Wilcoxon paired test.

releve´s, while being stable in BRA, although with a
different floristic composition (Tables Ia, Ib and IIa).
A Mantel test (Mantel statistic r: 0.2269, significance:
0.001) revealed a significant correlation of the floristic
composition and cover at the time of repopulation
with the current one. As indicated by Wilcoxon test
(Table IIa, upward arrows), some species significantly
increased their average cover, whereas others
decreased (Table IIa, downward arrows). Moreover,
we could not record at all three species that had been
included in the repopulation project (Euphorbia
paralias, Ononis natrix, Teucrium dunense), while
eight species originally not included were found in at
least one releve´ (Table IIa, exclamation mark).
In general, the “retreaters” have decreased to low or
null cover values in all beaches, irrespective of the
Figure 2. NMDS of the current relevees by beach. SAL, El Saler; GAR, Els Ferros – Garrofera; BRA, La Brava; MAL, La Malladeta; CAN,
El Canyar.

initial differences; newly observed species have a low,
uniform cover across beaches; species that have
increased their cover may show either a uniform cover
or a preferential distribution across beaches.
Discussion
The number of species recorded in our survey is in
line with those reported for the same area by Escaray
et al. (2010) and, in general, with the floristic
diversity measured in dunal communities, subjected
to highly selective environmental factors (Costa &
Mansanet 1981; Vagge & Biondi 1999; Acosta et al.
2007; Bertacchi & Lombardi 2014).
Furthermore, the number of species increased in all
beaches but one: this finding points to a positive
outcome of the restoration projects in quantitative
terms.
We recorded mostly native species and only one
alien species, Carpobrotus acinaciformis, a well-known
invader in several Mediterranean ecosystems (de
Montmollin & Strahm 2005; Sheppard et al. 2006;
Dal Cin D’Agata et al. 2009; Landi et al. 2012).
In our investigation, this alien species was found in
16/130 plots, with a low average cover compared with
documented cases of invasion where this species
poses serious threats to native communities (Fraga
et al. 2006; Traveset et al. 2008). The restored
communities seem therefore to possess some ability
to contain this invasive species.
The most frequent species are well represented in
Mediterranean coasts, extending either towards the
Atlantic or warmer areas. In Spain, they participate
in the well-defined phytosociological associations:
Agropyretum mediterraneum Br.-Bl. 1933; Medicago
marinae–Ammophiletum arundinaceae Br.-Bl. (1931)
1933 subass. lotetosum cretici Rivas Goday & Rivas-
Martinez 1958; Crucianelletum maritimae Br.-Bl.
(1931) 1933; (Costa & Mansanet 1981). Caespitose
species such as Achillea maritima and Elytrigia juncea
have a high cover, in agreement with this parameter
in other dunal communities (Vagge & Biondi 1999);
also Lotus creticus, Malcolmia littorea and Sporobolus
pungens have a good cover value. Surprisingly, Ononis
natrix, which had been implanted in all beaches, and
Euphorbia paralias, used only in two beaches, are
Figure 3. NMDS of the current relevees by dune face. WW ¼ windward face; CS, crest; LW, leeward face.

missing from our current releve´s; Ammophila arenaria
subsp. arundinacea, originally implanted in all
beaches, and Crucianella maritima (in four beaches)
are missing from most releve´s, despite their
important ecological role in the stabilization of
white dunes (De Lillis et al. 2004) and their
participation in psammophilous communities along
European coasts (Doody 1991). Our data do not
allow advancement of meaningful hypothesis to
interpret such findings. Euphorbia paralias was used
only in SAL and BRA with a low amount of
propagules, and we may speculate that they were
outcompeted by other, more expansive species.
However, Ononis natrix has disappeared everywhere
and Ammophila arenaria subsp. arundinacea was
observed in only 4/150 plots. Apparently, other
species have been favoured and were able to increase
their cover.
At the time of repopulation, all species contributed
significantly to the differences in species composition
and cover, whereas; in current releve´s some species
lost this feature, leading to a progressive homogen-
ization of the beach communities. However, new
species settled in, some contributing to significant
differences, albeit with low cover values. The species
that were used in repopulation mostly followed a
similar trend in all beaches over time, some
increasing, some decreasing, some remaining stable,
suggesting a common evolutionary pattern of the
dunal plant community, independently from how
remote the interventions of repopulation were. This
is in accordance with Benavent Olmos et al. (2004)
and Escaray et al. (2010), who noticed how it is
possible to reach, already 2–3 years after repopula-
tion, a satisfactory level of equilibrium of the
vegetation and a good fixing of the substrate,
provided that the dune bar has been left undisturbed.
The common trend shown by most species across
beaches explains the correlation between repopula-
tion and current matrices evidenced by Mantel test.
In this respect, the current differences observedacross
the beaches seem to be linked to past interventions.
In addition, the floristic assemblage in the
beaches that underwent the oldest interventions
(GAR, CAN) shows the same patterns but with more
extreme trends. This suggests us that their obser-
vation might provide hints about the future
evolutions of the plant community in the other
beaches (see also Escaray et al. 2010). SAL and GAR
are visited by a high number of tourists and are
served by public infrastructures, while the southern
beaches (BRA, MAL, CAN), devoid of infrastruc-
tures, experience a much lower level of disturbance.
However, our statistical analysis failed to detect a
significant effect of disturbance (data not shown).
The different volume of repopulation was another
factor of differentiation among the beaches. In MAL,
Figure 4. Ternary plots of selected species’distribution across dune faces. Species: a, Lotus creticus; b, Achillea maritima; c, Euphorbia terracina;
d, Elymus farctus; e, Calystegia soldanella; f, Malcolmia littorea. Plot vertices: WW, windward releve´s; CS, crest releve´s; LW, leeward releve´s.
At each vertex, value is 100% of the matching relevees and 0% of the others; ticks are set at 10% intervals. Each species’ data point is
represented by the % of distribution of their measured cover along the three releve´s (WW, CS, LW) of a single transect, see text for more
details. N, number of transects where the species was observed in at least one of the three releve´s WW, CS, LW. Transects where the species
was absent from all three releve´s were excluded. Dot size is proportional to the number of transects where the same distribution along the
WW, CS, LW releve´s was measured.

where the number of implanted specimens was lower,
the increase of coverage was on average more
dramatic. This can suggest how the implanted sand-
dune vegetation can compensate for a lower initial
density (as long as above a critical threshold) with a
subsequent more marked development (Perrow &
Davy 2002).
SAL and BRA, on the other hand, underwent the
most massive repopulations. SAL also scored a
significant increase of the presence of the introduced
plant species, but this increase concerned more the
spatial distribution of the species rather than their
total coverage, suggesting that once an optimal level
of coverage has been reached, the tendency of sand
plant communities is to reach an ecological balance
(Anwar Maun 2009).
Species composition and cover were significantly
linked to the dune face, matching the sea-inland
gradient of ecological factors occurring across the
whole dune system (Acosta et al. 2003; Martıńez &
Psuty 2004; Acosta et al. 2007; Biondi 2007). Again,
this result is in agreement with the vegetation series
occurring in Mediterranean sandy coasts. Sy´kora
et al. (2003) pointed out how in the coasts of Greece
and of other Aegean areas the strandline vegetation,
represented by Cakiletea maritimae, is spatially linked
to a sand-dune vegetation represented by Ammophi-
letea. A similar pattern was described, among others,
by Diaz Garretas (1982) on the Almeria coast, by
Mijovic´ et al. (2006) on the sandy beaches of the
Montenegro coast and by Stambouli-Meziane et al.
(2009) on the Algerian coast, confirming the peculiar
character of psammophilous vegetation and its
tendency to become established with similar patterns
in different areas due to its high specificity (Van der
Maarel et al. 1985; Studer-Ehrensberger et al. 1993;
Van der Maarel & Van der Maarel-Versluys 1996).
Acosta et al. (2003) proposed a similar vegetation
series to define the potential natural vegetation along
the sandy coasts of Lazio, Italy. Therefore, different
times of intervention allow to follow in parallel the
different phases of development of a plant commu-
nity, given a sufficient level of homogeneity of
conditions along the coast. Interestingly, the pattern
of species expansion/retreat shows that the commu-
nities have rearranged themselves in the time elapsed
from the restoration to our releve´s, as a response to the
ecological gradient operating from the windward face
to the leeward face of the dune system. The
rearrangement has occurred through a complex
pattern of species expansion and retreat, consistent
with the species’ auto- and synecological preferences.
Although single species showed generally the same
behaviour – expansion, retreat or stability – across
the different beaches, many of them showed
preferences for specific dune faces, where they were
able to increase their cover while retreating from less
suitable area. So, Calystegia soldanella increased its
cover in WW, maintained it in CS, but decreased it in
LW; Salsola kali, not included in the repopulation,
appeared in all beaches and in all dune faces but with a
marked preference for LW and CS; Achillea maritima
increased its cover in WW, maintained it in CS and
disappeared from LW; Malcolmia littorea retreated
from WW plots while expanding in CS and LWones.
Even species with a generalized decrease showed
a preference in that their decrease was more
pronounced in specific dune faces: Cakile maritima
subsp. maritima and Echinophora spinosa decreased to
low levels in LW; Eryngium maritimum and Cyperus
capitatus decreased in WW; Pancratium maritimum
decreased in WW and CS.
The good performances of Calystegia soldanella
and Achillea maritima can find an explanation in their
high fitness to the ecological conditions of coastal
sand dune systems, due to leaf morpho-anatomical
adaptations like orientation, roll, non-glandular and
glandular trichomes, general morphology of epider-
mis, morphology and localization of stomata,
hydathodes, aerenchyma and water-storage parench-
yma (Ciccarelli et al. 2009).
Elytrigia juncea has been already demonstrated to
have a good capacity of colonization and regenerative
potential (Harris & Davy 1986); Malcolmia littorea
has pioneer, thermophilous and oligotrophic charac-
teristics that favour its spreading on the back of the
dunes (Calvo Sendıń et al. 2000).
Carbobrotus acinaciformis, an alien invasive
species, was recorded in few plots, with a low cover.
Maltez-Mouro et al. (2010) showed how the
destabilizing effects of the congeneric C. edulis on
dunal plant communities in the north of Portugal
were very weak, pointing out the strong resilience to
invasions of the sand dune systems analysed. We may
hypothesize that also the Albufera plant communities
are resilient to invasions, probably due to their recent
implant and dynamic stage. However, climatic
factors such as low rainfall might affect the success
of invasion (Carboni et al. 2010).
Comparison with historical data
The increase in floristic diversity recorded over the
time elapsed from the restoration projects is
consistent with a succession from a reconstituted
community to a more natural one. At the start of the
project, the reconstitution of communities had to
deal with some compromises, such as for instance the
availability of plant material, its amenability to
massive propagation and its response to transplants
in situ. For these reasons, the number of species used
for the restoration was lower than the actual floristic
diversity of the natural communities. Despite this
shortcoming, it seems that the restoration has paved

the way for the subsequent colonization by native
species, while retaining at the same time a good
ability to contain alien invasives.
The complex pattern of new colonization,
expansion, decline or disappearance of introduced
species cannot be fully explained in the light of our
data. We may only speculate that it is the likely result
of complex ecological processes connecting the biotic
and abiotic components of this ecosystem, including
the unknown contribution of seeds in the soil seed
bank of the sand used in shaping the dunal bar.
Unravelling such processes would require, inter alia,
a study of soil microorganisms and mycorrhizae,
allelopathic effects, pollinators, reproductive fitness,
seed germination and facilitation systems.
Conclusions
In conclusion, it is possible to state that the
vegetation of the first dunal bar of the beaches
composing Devesa de la Albufera, after a time
ranging between 6 and 22 years from the first
interventions of repopulation, is in a condition of
relative stability, in which significant floristic differ-
ences have developed through a rearrangement of
plant species in the three sectors of front, crest and
back of the dune. The future developments will be
probably represented by a further spreading of those
species that already showed along the years a marked
tendency to propagate and a further “marginaliza-
tion” of the ones that reduced their areas of
distribution and coverage values.
Acknowledgements
The Authors would like to thank Dr Katarıńa
Hegedu¨sˇova´, Institute of Botany SAS, Bratislava,
Slovak Republic, for useful help with multivariate
statistical analysis, and Dr Lorenzo Peruzzi, Depart-
ment of Biology, Pisa University, Italy, for precious
assistance with nomenclatural issues.
Notes
* Email: dimivilli@hotmail.com
** Email: antoni.aguilella@uv.es
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