Trampling impact on vegetation of embryonic
and stabilised sand dunes in Montenegro
Urban Šilc1,2
& Danka Caković3
& Filip Küzmič1
& Danijela Stešević3
Received: 1 September 2016 /Revised: 3 October 2016 /Accepted: 4 October 2016 /Published online: 18 November 2016
# Springer Science+Business Media Dordrecht 2016
Abstract Trampling is one of the human activities that are
harmful for plant species and communities of sand dune
ecosystems. The aim of this study was to compare the vegetation
of embryonic and shifting Ammophila sand dunes
with and without fencing to limit trampling. Fenced sand
dunes appeared to be richer in species but differences were
more prominent in embryonic sand dunes. Some species
(Cakile maritima, Pancratium maritimum) were missing on
trampled embryonic dunes. The positive impact of trampling
exclusion on embryonic sand dunes was indicated by a
lowered slope in a Whittaker graph as well as by rarefaction
curves that showed higher species richness on the lower
slope. Changes in the vegetation of more stabilised shifting
Ammophila sand dunes due to trampling are not evident,
although species composition is also impoverished.
Fencing of parts of sand dunes proved to be an effective
measure for vegetation conservation. In addition to physical
exclusion of visitors, fences can also have symbolic value
for raising public awareness.
Keywords Habitats directive . Conservation . Fencing .
Trampling
Introduction
Sandy beaches are unique as the world’s single largest dynamic
type of open shoreline and are found at the boundary between
land and sea (McLachlan and Brown 2006). Although
they seem to be abiotic deserts, they provide habitats for specific
biota, determined by three interacting factors: waves,
tides and sand particle size (McLachlan 2001). A specific
vegetation occurs in this naturally highly dynamic environment,
which has developed to be highly resistant to natural
disturbances but is very sensitive to anthropogenic ones
(Andersen 1995; Ciccarelli 2014; Garcia-Mora et al. 1999).
Defeo et al. (2009) summarised the various human stressors to
which dune biota is exposed: recreational seashore activities
(incl. mechanical impact caused by trampling, camping etc.),
beach cleaning, nourishment, pollution, exploitation, biological
invasion, coastal development and engineering, mining
and climate change.
Trampling by beach visitors is commonly considered to be
the most harmful mechanical stressor, identified as being more
harmful for plant development than natural stress (Santoro
et al. 2012). Mechanical stress on plants causes mobilization
of assimilates into storage organs and increases the belowground
biomass, which affects the survival of the plants and
their reproduction, as well as germination and establishment
(Liddle 1975; Puijalon et al. 2008). However, different species
have different sensitivity to trampling and physical damage
(Seer et al. 2015). Trampling does not affect vegetation (and
fauna) only directly but also indirectly through soil compaction,
which leads to differences in its properties (Brown and
McLachlan 2002). Constant trampling pressure was estimated
to be destructive for plant populations even among beach
plants adapted to rough site conditions, so the final outcome
is a reduction of biodiversity, population size and distribution
(Andersen 1995; Scott 1976).
* Urban Šilc
urban@zrc-sazu.si
1
ZRC SAZU (Research Centre of the Slovenian Academy of Sciences
and Arts), Institute of Biology, Novi trg 2, 1000 Ljubljana, Slovenia
2
BC Naklo, Strahinj 99, 4202 Naklo, Slovenia
3
Faculty of Natural Sciences and Mathematics, Biology department,
University of Montenegro, Džordža Vašingtona bb, 81
000 Podgorica, Montenegro
J Coast Conserv (2017) 21:15–21
DOI 10.1007/s11852-016-0468-2
The aim of our study was to focus on diversity changes in
dune vegetation with limited trampling. We compared (i) species
richness and (ii) species change in plant communities
between two habitats of sand dune systems (embryonic and
Ammophila stabilised dune) in fenced and unfenced parts.
Knowledge of the resistance of sand dune communities to
trampling and their ability to regenerate plays a key role in
landscape planning and conservation (Andersen 1995).
Material & methods
Study area
With a length of 12 km, Velika plaža in Ulcinj (Montenegro) is
considered the longest beach on the Adriatic coast. It is located
at the south-eastern end of Montenegro and is bordered by the
channel of Port Milena to the west and the Bojana River to the
east, covering an area of 1270 ha. The average width of the
beach and its hinterland is 1.5 km and it is bordered to the
north by a regional road. The area is frequently exposed to
very strong winds, which carry sand from the shore to the
inland and which consequently covers a major part of the area
(Mijović 1994). According to the Köppen-Geiger system, the
climate of Ulcinj is classified as Csa type – a Mediterranean
climate with hot summers. The mean annual temperature is
15.5°C, while mean precipitation is 1258 mm. The warmest
months are July and August (24.3°C) and the coldest January
and February (6.8°C) (Burić et al. 2014). The psammophytic
vegetation of Velika plaža forms two belts. The one closer to
the sea is represented by 4 xerohalophytic plant communities:
Cakilo-Xanthietum italici (Beg. 1941) Pignatti 1958,
Euphorbio paraliae-Agropyretum junceiformis Tüxen in Br.Bl.
& Tüxen 1952 corr. Darimont, Duvigneaud & Lambinon
1962, Eryngio-Sporoboletum virginici Gehu & Uslu 1989,
Medicagini marinae-Ammophiletum australis Br.-Bl. 1921
corr. F. Prieto & T.E. Díaz 1991, one replacing the other along
the salinity and moisture gradient towards the inland (Mijović
1994; Mijović et al. 2006; Šilc et al. 2016b). The second belt,
which is not directly influenced by seawater, comprises
xeropsammophytic communities of Mediterranean pastures
and dry meadows (Mijović 1994).
Eleven NATURA 2000 habitats are reported for the beach
and its hinterland: Annual vegetation of drift lines (1210),
Embryonic shifting dunes (2110), Shifting dunes along the
shoreline with Ammophila arenaria (white dunes, 2120),
Fixed coastal dunes with herbaceous vegetation (grey dunes,
2130*), Humid dune slacks (2190), Dunes with Euphorbia
terracina (2220), Mediterranean salt meadows (Juncetalia
maritimae) (1410), Brachypodietalia dune grasslands with
annuals (2240), Wooded dunes with Pinus pinea and/or
Pinus pinaster (2270), Mediterranean temporary ponds
(3170), and Salix alba and Populus alba galleries (92A0)
(Petrović et al. 2012). Presence of habitat type 2130* that
has Atlantic distribution (Houston 2008) is questionable.
The most representative 500 ha of psammophytic vegetation
has been recognized as a monument of nature and is
therefore protected from trampling and grazing by cattle with
a wooden fence, which was installed in 2014. The rest of
Ulcinj beach (Velika plaža) is under the major impact of tourism,
illegal dumping, sand exploitation and urbanization
(Petrović and Karaman 2009; Šilc et al. 2016a).
Vegetation sampling
Since vegetation was not sampled prior to fence installation,
we used a synchronic (space-for-time) approach, which enables
the best surrogate for studying vegetation succession/
changes when permanent plots are not available (GlennLewin
and van der Maarel 1992; Walker et al. 2010).
At the beginning of May 2016, we sampled vegetation
within fences, where trampling was limited. Fenced sand
dunes belong to two habitat types: Embryonic shifting dunes
(2110) and Shifting dunes along the shoreline with
Ammophila arenaria (white dunes 2120). In each of the habitat
types (further referred in text as embryonic dunes and
shifting Ammophila dunes), we selected a number of plots,
sized 2 × 2 m, on which we recorded all vascular plant species
and visually determined their cover (specific and in total),
using the Braun-Blanquet scale. Plots were selected approx.
every 30 m to represent microhabitat diversity. For the nonfenced
part of the dune system, we used plots of the same size,
sampled for sand dune monitoring in May 2015. Plots were
laid every 500 m within both habitat types in the most impacted
zone of the beach. The same plot size and sampling method
were used as with the fenced part of the beach and the same
researchers were present. Number of plots sampled is presented
in Table 1.
Data analysis
Basic analysis of vegetation data was done in the Juice program
(Tichý and Chytrý 2006): calculation of frequency and
average plant species cover values. Differences in species
richness between different parts of the sand dune system were
tested with nonparametric tests (Mann-Whitney) in Statistica
10.0 (StatSoft 2011). Similarity Percentages analysis
(SIMPER) (Clarke 1993) was used to identify the taxa mostly
contributing to differences between fenced and non-fenced
sites. Analysis was performed in the PAST program
(Hammer et al. 2001).
Rarefaction curves were used for estimating standardized
species diversity (Gotelli and Colwell 2001); they had already
been used in a similar study on sand dune vegetation (Santoro
et al. 2012). We compared species richness between two sand
dune habitats in fenced and non-fenced parts. We calculated
16 Šilc U. et al.
sample-size based rarefaction and extrapolation (Colwell et al.
2012) using iNext online software (Hsieh et al. 2013).
For determination of species changes in plant communities
under different disturbance regimes, we used a diversity/
dominance (Whittaker) diagram (Magurran 1988; Whittaker
1970). When a community is under shifting disturbance intensity,
the shape and slope of the diagram can show differences
in species composition due to various kinds of stress.
The diversity/dominance diagram was made as a rank/
abundance diagram. For each part of the sand dune system
Table 1 Frequency (in percentage) and rank (based on average cover; in superscript) of plant species in each sand dune habitat (for fenced and
unfenced parts)
Embryonic dunes Shifting Ammophila dunes
Unfenced Fenced Unfenced Fenced
Nr. of plots 13 20 13 22
Alkanna tinctoria 5 8 8 10 55 5
Ammophila arenaria 8 1 14 1
Amorpha fruticosa 8 3
Anagallis arvensis 9 15
Arenaria serpyllifolia agg. 45 5
Arundo donax 5 8
Cakile maritima 30 8
Catapodium rigidum 8 23
Crepis foetida 10 8 38 3 45 15
Cutandia maritima 15 23
Cynodon dactylon 8 6 15 10
Cyperus capitatus 31 1 100 2 54 3 100 8
Dittrichia viscosa 18 15
Echinophora spinosa 31 3 90 5 69 10 77 15
Elymus farctus 23 2 25 8 31 3
Eryngium maritimum 50 8 31 10 41 15
Euphorbia paralias 8 6 8 10
Euphorbia peplis 8 3 15 8 23 23
Euphorbia terracina 9 3
Holoschoenus romanus 8 3
Hypochaeris radicata 9 7
Inula crithmoides 5 8
Lagurus ovatus 8 3 9 14
Linum strictum 5 8 68 8
Lophochloa cristata 8 10
Medicago littoralis 5 2 54 10 100 8
Medicago marina 10 5 8 23 64 1
Oenothera spp. 5 8 15 1 55 15
Onobrychis caput-galli 15 10 14 8
Pancratium maritimum 20 8 23 3 86 15
Polygonum maritimum 20 8 8 10 5 15
Pseudorlaya pumila 8 6 10 5 31 10 100 8
Salsola kali 8 6 20 8 15 10
Senecio vulgaris 14 15
Silene conica 23 15
Valerianella sp. 5 8
Vulpia fasciculata 5 1 31 10 100 4
Xanthium italicum 62 3 80 2 77 10
Trampling impact on vegetation of sand dunes 17
researched, we calculated the average cover for each species,
calculated their relative cover to the total cover of species in all
sampled plots and then ranked them according to the results.
The relative frequency of each species was log transformed.
We then plotted each species rank (from the most to the least
abundant) on the X-axis and the frequency of particular species
on the Y-axis.
Results
Altogether, 41 plant species were found on the researched
sand dunes. On embryonic sand dunes, species richness was
lower on trampled plots than on fenced ones (5.15 vs. 1.85,
Z = 4.13, p < 0.01) and the same was found for shifting
Ammophila dunes (10.64 vs. 6.15, Z = 4.13, p < 0.01).
We performed SIMPER analysis to detect species that contribute
to dissimilarities between fenced and unfenced sites.
Similar differences between sites were found on embryonic
(80.09%) and shifting Ammophila dunes (84.36%). Cyperus
capitatus (38.24%) and Xanthium italicum (18.97%) have a
higher contribution on embryonic dunes. Several characteristic
species of sand dunes are missing (Table 1) on unfenced
embryonic sand dunes (Cakile maritima, Pancratium
maritimum, Eryngium maritimum) or their frequency is reduced
(Salsola kali, Cyperus capitatus). Several characteristic
species are also less frequent on unfenced shifting Ammophila
dunes: Medicago littoralis, Pancratium maritimum, Vulpia
fasciculata, and Cyperus capitatus. Vulpia fasciculata
(17.03%) and Medicago marina (15.74%) make a higher contribution
to dissimilarity on stable dunes.
The pattern of increased species richness is also shown by
the rarefaction curves (Fig. 1). The rarefaction curves between
the two habitats are different, since shifting Ammophila dunes
are more species rich. There are also evident differences between
the rarefaction curves of fenced and unfenced embryonic
dunes, with lower species richness in the unfenced part.
There are no significant differences in species richness and
rarefaction curves of shifting Ammophila dunes.
Embryonic sand dunes are characterized by the presence of
a few dominant species. There were more species present in
the fenced part of the sand dune system, the dominant species
were more frequent and the slope of the regression line was
lower (Table 1, Fig. 2). Shifting Ammophila dunes include
more species and there are almost no differences between
fenced and unfenced parts. Slope and intercept values are very
similar (Table 2).
Discussion
According to our study, there are evident differences in the
vegetation of embryonic and shifting Ammophila dunes due to
trampling impact. Embryonic dunes show low resistance,
while more fixed dunes are less impacted. Human impact on
sand dunes by trampling has been shown in many studies
(Fenu et al. 2013). The impact of trampling is not always
negative, since it can be used as a management tool at moderate
intensities (Andersen 1995; van der Maarel 1971); however,
it is generally considered problematic for conservation
and management of biodiversity.
There is a great difference in the trampling impact on different
plant communities. They vary in their resistance and
resilience to trampling. Meta-analysis of literature related to
trampling of vegetation has revealed that the initial resistance
of a plant community and the length of recovery period are
better predictors of resilience than the intensity of trampling
(Pescott and Stewart 2014). Depending on plant traits, different
plant species become affected to different degrees following
certain impacts, such as trampling (Seer et al. 2015).
Vegetation analyses in trampled foredunes and dunes have
shown a decrease in vegetation cover and an increase in annuals
and trampling-tolerant ruderal and grassy species
(Andersen 1995; Garcia-Mora et al. 1999; Labuz and
Grunewald 2007). In our case species tolerant to trampling
(Cynodon dactylon, Arenaria serpyllifiolia agg.) and also
alien species (Amorpha fruticosa) are found in the unfenced
part of the beach indicating changes in species composition
because of trampling effect. Invasion of alien species and
trampling are characteristic for coasts influenced by the coastal
suburbanization phenomenon (Carboni et al. 2009).
Although we did not perform a BACI (Before–After
Control-Impact) experiment, the results of our study indicate
changes of sand dune vegetation after limitation of trampling.
A special feature of our study is that we analysed the impact of
trampling within two different habitats: Embryonic shifting
dunes (2110) and Shifting dunes along the shoreline with
Ammophila arenaria (2120). There is a rich body of literature
on the trampling of sand dune vegetation but mainly from
Atlantic coasts, while from Mediterranean areas there have
been only a few studies and they concentrate more on embryonic
dunes (Kutiel et al. 2000; Santoro et al. 2012).
The two habitats differ in their response to trampling.
Species diversity is reduced on embryonic dunes because of
trampling, while it remains similar on more stable shifting
Ammophila dunes. It is important to emphasize difference of
substrate between both habitats and consequently the impact
of trampling. Higher impact is visible in the not stabilised
sandy substrate. Damage to vegetation in water saturated, soft
substrates is more likely (Eckrich 2000). One reason is that
vegetation of fixed dunes is more resistant to trampling damage
(Andersen 1995; Lemauviel and Rosé 2003; Rickard et al.
1994), another is that these dunes are more distant from the sea
and less trampled even in an unfenced part of the beach.
Vegetation of more fixed shifting Ammophila dunes is more
abundant and consists of many plant species with spiny fruits
18 Šilc U. et al.
(Onobrychis caput-galli, Medicago spp., Pseudorlaya
pumila). Although plants on embryonic dunes are also spiny
(Echinophora spinosa, Eryngium maritimum) similar plants
on shifting dunes have higher cover. We believe that visitors
tend to avoid them and trampling is also more concentrated on
existing paths.
The speed of recovery of vegetation varies between plant
communities. Both Davenport and Davenport (2006) and
Hylgaard and Liddle (1981) report slow recovery, while it
can be very fast in other cases, since one to two years of
trampling limitation enables recovery of the diversity and
structure of dune plant communities (Santoro et al. 2012).
We did not especially focus on the recovery of sand dune
vegetation but our results suggest that embryonic dune vegetation
recovered very fast, since the fences were only installed
two years ago. Fast recovery of pioneer herbaceous vegetation
is related to adaptation of this vegetation type in the
Mediterranean to the natural disturbances to which it is usually
exposed (Kutiel et al. 2000). It is important to emphasize that
recovery of total cover of vegetation is less important than
enriched species composition and the presence of particular
rare species when recovery is considered (Kerbiriou et al.
Fig. 1 Rarefaction curves of both dune habitats - on the left the embryonic dunes vegetation, on the right the shifting Ammophila dunes (fenced - red and
unfenced - grey)
Fig. 2 Diversity-dominance
(Whittaker) diagram of fenced
and unfenced sand dune habitats.
Legend: S - shifting Ammophila
dunes, E - embryonic dunes, n unfenced,
f - fenced. Dashed line embryonic
dunes, solid line shifting
Ammophila dunes
Trampling impact on vegetation of sand dunes 19
2008). Thus, an area closer to the sea harbouring only few
unique and endangered species could be considered of great
importance (Acosta et al. 2009).
Most trampling studies have been made on fairly narrow
paths (experimental or already established) (for example
Brunbjerg et al. (2015)), while our study takes into account a
larger area, similar to that of Santoro et al. (2012). This is a
better approximation of the real situation on a sand dune system
under recreational impact. Vegetation along paths with
frequent trampling is much damaged, especially less resilient
plant communities, but random trampling (even less intensive)
in the wider area of a beach is less preferred even if it is less
harmful. A management plan with a controlled flow of tourists
is therefore very important.
Our results are important for the consideration of conservation
and management measures of sand dunes. The study confirms
the importance of fence installation for the prevention of
trampling of sand dune vegetation, particularly in highly visited
areas, as has already been pointed out by other researchers
(Grafals-Soto and Nordstrom 2009; Santoro et al. 2012).
Fences are effective in directing visitors and the zonation of
recreation into high and low intensity usage is a better conservation
strategy than less intensive but widespread trampling
(Pescott and Stewart 2014). Another important use of fences
is restoration of sand dune vegetation by limiting access that
leads to improvement of species cover and richness as well as
spatial organisation of plant communities (Acosta et al. 2013).
Fences are a low cost measure for the control of human and
animal access on sand dune systems and effectively prevent
trampling damage to vegetation. In addition to physical prevention,
there is also psychological momentum, indicating vulnerable
habitats worth protecting. In such cases, they can be described
as symbolic fences. A positive side effect of these
fences alone is raising awareness and it becomes even greater
with the erection of information boards. In our case specifically,
more rigid fences were built in order also to prevent cattle
grazing (not only trampling by tourists) but at the same time
to allow aeolian transport of the sand (Nordstrom et al. 2000).
The study area was already legally protected in 1968 as a
natural object (Sl. list SRCG, 30/68) and in 2007 as a natural
monument (Sl. list CG, 30/07). The installation of
fences in 2014 marked additional conservation efforts.
Three habitat types were fenced: embryonic and shifting
Ammophila dunes and dune wetlands. Although the first
habitat is most impacted by trampling and the latter two
are less accessed by tourists due to remoteness from the
sea and unattractiveness, it is also important to protect them.
Because Montenegro has still not established a NATURA
2000 network and the development of the country is directed
towards tourism, conflict between nature conservationists
and landscape planners is expected.
Acknowledgements We thank anonymous reviewer for helpful suggestions
and comments. Martin Cregeen kindly checked our English.
The research was partly financed by Slovenian Research Agency
(ARRS) through a research program (P1-0236) and bilateral project
(BI-ME/16-17-018).
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