RESEARCH ARTICLE

Facilitation and sand burial affect plant survival

during restoration of a tropical coastal sand dune
degraded by tourist cars
Leonardo H. Teixeira1,2 , Wolfgang Weisser3 , Gislene Ganade1

Coastal sand dunes support various ecosystem services, including storm protection and tourism. Restoration programs are
often critical to preserve this ecosystem due to its fragility and high degree of degradation. Dune restoration still suffers from a
general lack of knowledge of the ecological processes controlling tropical dune communities. We investigated if facilitation can
increase restoration success by assisting plant survival in a Brazilian coastal dune degraded by buggies at the Environmental
Protection Area of Jenipabu, RN, Brazil. We performed two field experiments on dune crest sites degraded by buggies. The first
experiment tested how the presence of established vegetation and coconut mesh, mimicking soil-stabilizing effect of vegetation,
facilitates seedling survival and establishment of the early successional dominant dune plant Canavalia maritima (Fabaceae).
The second experiment tested if coconut mesh and initial irrigation would allow the establishment of C. maritima outside the
vegetation, using both seeds and transplants. We found that the presence of established vegetation positively affected seedling
survival. Application of mesh positively affected microclimatic conditions and slightly increased survival, but only for seeds.
Initial irrigation only had a small effect on transplanted seedlings survival. Nevertheless, there was extreme seedling mortality
in the experiment in particular from sand burial. We conclude that restoration of these dune crests is very difficult. Facilitation
by vegetation can reduce plant mortality by sand burial and desiccation, but successful restoration is likely to require a major
effort involving a very high number of seedlings or seeds.
Key words: abiotic conditions, Canavalia maritima, coconut mesh, germination, microclimate, seedling establishment, vege-
tation patches

be considered endangered due to habitat loss (Prisco et al.

Implications for Practice
• Restoring coastal dune vegetation degraded by buggies is
likely to be expensive, time-consuming and to have a high
chance of failure.
• The better management of cars on the dunes would benefit
the tourism economy while avoiding the damage of the
sensitive dune vegetation patches by vehicles.
• Seed sowing seems to be a promising tool to restore dune
vegetation. We suggest that future restoration experiments
should use a large number of scarified seeds from a variety
of plant species.
• Facilitation techniques, such as nucleation, should be
incorporated in coastal dunes restoration programs to
enhance success and reduce long term restoration efforts.
Introduction
2013). Coastal dunes perform important ecosystem services
such as protection against high tides, storm-waves, and sand
invasion (Nordstrom et al. 2002). To combat the demise of dune
vegetation, restoration programs are being applied to stabilize
or increase vegetation cover in dune areas degraded by human
activities.
Dune restoration programs employ a number of techniques
such as the application of dead plant materials to stabilize soil
conditions and release nutrients (Lemauviel et al. 2003), or the
use of wood-fences that rebuild the dune slope and protect
plants from sand burial (Nordstrom et al. 2002). Most of those
techniques are expensive and the restoration of dune vegetation
Author contributions: GG conceived and designed the research; LHT performed the
experiments; LHT, WWW, GG analyzed the data; GG contributed with funding for
materials and equipment; LHT, WWW, GG wrote and edited the manuscript.
1 Departamento de Ecologia, Centro de Biociências, Universidade Federal do Rio

Grande do Norte (UFRN), CEP 59078-900, Natal, Brasil

Coastal dunes around the globe are considered endangered due 2 Address correspondence to L. H. Teixeira, email leonardo.htp@gmail.com

to increasing human impact (Nordstrom et al. 2002; Martínez 3 Terrestrial Ecology, Department of Ecology and Ecosystem Management,

et al. 2004; MEA 2005). Large dune areas have been converted
to private housing as well as touristic resorts, and dune remnants
are often under intense human pressure due to tourism activities,
such as car driving (e.g. Kutiel et al. 2001). In consequence,
many plant and animal species occurring in coastal dunes can
School of Life Sciences Weihenstephan, Technische Universität München,
Hans-Carl-von-Carlowitz-Platz 2, D-85350 Freising, Germany
© 2016 Society for Ecological Restoration
doi: 10.1111/rec.12327
Supporting information at:
http://onlinelibrary.wiley.com/doi/10.1111/rec.12327/suppinfo

Restoration Ecology 1
Restoration of tropical coastal dunes
cover normally takes a long time (Gómez-Pina et al. 2002; Rozé
& Lemauviel 2004; Ruiz-Jaen & Aide 2005). Few attempts to
restore dunes have been conducted in the tropics, where extreme
harsh environmental conditions and constant soil movements
on dune crests require the development of specific techniques
suitable for such environment.
A promising technique for the restoration of tropical coastal
dunes might be the use of established vegetation for nurs-
ing transplanted seedlings, because plant–plant facilitation is
expected to be important for structuring plant dune commu-
nities (Callaway et al. 2002; Franks 2003; Franks & Peterson
2003). The dune vegetation itself can contribute to further plant
colonization and successful establishment (Franks 2003; Franks
& Peterson 2003), leading to higher soil stabilization and veg-
etation cover (Jones et al. 2008). Facilitative interactions can
help to increase restoration success by reducing the negative
effects of unstable soils, harsh microclimatic conditions, and
low resources availability, which compromises plant establish-
ment (Greipsson 2002; Martínez & Moreno-Casasola 1996;
Zheng et al. 2005; Gilbert et al. 2008).
Plant mortality can jeopardize restoration programs. Coastal
dunes naturally have severe abiotic conditions that can lead to
plant death. For example, seedling establishment in bare soil is
hindered by frequent sand movements causing sand burial, tem-
porary droughts, and lack of nutrients (Franco & Nobel 1988;
Maun 2004). In contrast, seedling growth is often higher within
vegetation patches than outside, due to temperature reduction
and moisture enhancement, which decreases plant transpiration
(Martínez 2003; Franks 2003). Other described causes of plant
mortality in dunes are trampling (by humans or vehicles),
desiccation, and herbivory (Bach 2001; Lemauviel & Rozé
2003; Boudreau & Faure-Lacroix 2009; Bagousse-Pinguet
et al. 2013). However, sand burial has been described as the
crucial factor defining patterns of vegetation distribution in
coastal dunes, sometimes working as an environmental filter
for the occurrence of plant species (Maun 2004; Yura & Ogura
2006; Gilbert et al. 2008).
In Northeast Brazil, coastal dunes commonly form high hill
tops with intense sand movements due to strong wind influence
(Trindade 1991; Hesp 2004; Hesp & Martínez 2007; Torres
et al. 2009). In addition, dunes suffer from a particular negative
impact of touristic cars activities that suppress vegetation
patches located at the dune crest. Hence, it is important to deter-
mine if the remaining vegetation patches (i.e. nurse plants and
nucleation effects) could help seeds and seedlings to overcome
the difficulties to colonize degraded sand areas (Franks 2003;
Franks & Peterson 2003; Martínez 2003; Rudgers & Maron
2003; Yura & Ogura 2006).
In this article, we focus on using the potential of facilitation
for restoring dune plant communities. As a model species for
restoring dune crest vegetation, we used Canavalia maritima,
an early successional legume species common to tropical coastal
dunes. Like many other species dominating early successional
dune communities, C. maritima is a fast-growing, prostrate, and
rhizomatous plant species (e.g. Hesp 2004).
This article addresses the following questions: (1) can veg-
etation patches be used to increase seedling establishment
2 Restoration Ecology
for restoration of degraded dune crest areas? (2) can facilita-
tive effects of vegetation patches on seedlings be mimicked
using an artificial substitute, here coconut mesh, to increase
seedling establishment during restoration? (3) can seeds instead
of seedlings be used for restoration? and (4) does initial irriga-
tion further increase seedling survival?
Methods
Study Site
The experiment was carried out in the dunes of Jenipabu beach
in Northeast Brazil that are frequently damaged by tourist cars
(“buggies”) driving on the dunes (Appendix S1, Supporting
Information). For our experiment, we chose vegetation patches
in an area of 20 × 500 m at the top of the dunes that were
damaged by the transit of buggies. These former trails were
vegetation free, that is they have not recovered from the damage
after several years. The dominant plants in these vegetation
patches were cashew trees (Anacardium occidentale), Brazilian
pepper trees (Schinus terebinthifolius), and cocoplum shrubs
(Chrysobalanus icaco), which are common species in this dune
area (Trindade 1991; Torres et al. 2009). The environmental
protection area of Jenipabu where the experiment took place is
bordered by a small fence (maximum height of 1.50 m) that is
frequently buried and does not offer sufficient protection for the
dunes. In fact, during several experimental days, buggies were
encountered inside the protected area.
Experimental Design and Statistical Analyses
Experiment 1: Plant Facilitation versus Application of Plant
Material (Facilitation Experiment). To test for the importance
of facilitation relative to more conventional restoration methods
for improving seedling survival (i.e. placing plants directly on
bare soil), we manipulated the presence of vegetation (facilita-
tion) and the use of coconut mesh. Coconut mesh potentially
mimics some of the effects established plants might impose
on seedling emergence such as: reducing sand movements,
providing protection against desiccation, and releasing nutri-
ents to soil. Seeds of Canavalia maritima were collected in
August 2010 and germinated in the greenhouse to produce
the 4 month-old seedlings that were used in the experiment
(Appendix S1).
At the dune crest area where buggies had partially destroyed
vegetation, seedlings of C. maritima were transplanted into the
sand using a split-plot design (Fig. 1A), on February 2011. Four
pairs of seedlings were planted under the remaining vegetation
or in bare soil, just outside the vegetation (split factor). For
two of these pairs, plants were planted directly in the soil;
for the other two pairs, seedlings were planted under a 1 m2
coconut mesh (Appendix S1). Overall, we had four replicated
blocks × two vegetation treatments (+ −) × two mesh treatments
(+ −) × two subplots per treatment × two seedlings per subplot,
that is 64 seedlings used in the experiment (Fig. 1A). The
experiment was set up over a period of 2 weeks.
 Restoration of tropical coastal dunes

State Rio Grande do Norte

Brazil

EPA
Jenipabu

A Experiment 1 (Facilitation experiment) B Experiment 2 (Restoration experiment)

Inside Outside Outside Vegetation
Vegetation Vegetation (V-)
(V+) (v-)
Subplot Bare Plot
sand

Mesh 10 scarified
Plot
seeds
Block
10 non-scarified
Water seeds
Mesh (M+) Seedlings
(2 in total)
No mesh (M
-) Water
+
Mesh

Block

Figure 1. Experimental design for the facilitation (A) and the restoration (B) experiments. Blocks were implemented inside and outside vegetation (split-plot
design) for the facilitation experiment, testing the effects of vegetation, mesh, and vegetation and mesh combination on plant survival. The restoration
experiment was implemented only outside vegetation and plant survival was tested for irrigation, mesh, and irrigation plus mesh treatments. The use of seeds
for restoring coastal dunes was also tested in the restoration experiment. For this, the efficiency of the thermal scarification method for seed germination was
evaluated. Photo by Jomar Jardim.

Starting in March 2011, we recorded seedling establish-
ment and the causes of mortality for all transplanted seedlings
monthly until July. In three of the four visits, we recorded the
following abiotic variables for each plot at the same day from
10:00 hours to 14:00 hours: air humidity (%), soil surface tem-
perature aboveground (∘ C), and soil temperature at 10 cm depth
(∘ C) using a thermo-hygrometer clock MT-240 (Full Gauge
Controls, Canoas, RS, Brazil) and a digital skewer thermome-
ter (Incoterm Thermometers Industry, Belo Horizonte, MG,
Brazil). If plants were buried under sand, we measured the depth
of sand burial in cm. We also measured plant growth as the
length of the shoot and the number of leaves. Both measures
were taken each time plants were visited.
Experiment 2: Restoration Outside Vegetation Using Plant
Material (Restoration Experiment). The second experiment
tested if the use of plant material (coconut mesh) outside veg-
etation can increase the rate of lateral vegetation expansion.
In contrast to the first experiment, seedling establishment was
tested more than 1 m outside the remaining vegetation. In addi-
tion, we added an irrigation treatment to test if this together with
the properties of the coconut mesh improves restoration success
of dune vegetation.
Outside each of five dune crest vegetation patches degraded
by buggies, four pairs of two seedlings were planted on
bare sand. Next to each seedling pair, we added 20 seeds of
C. maritima directly into the soil, 10 of them scarified using
the thermal method (Appendix S1) and 10 not scarified. The
fast thermal method for scarification was chosen because of
the large number of seeds involved. In a cross-factorial design
we applied two treatments to the four plots of each block:
the presence and absence of initial irrigation and the presence
and absence of coconut mesh. Initial irrigation consisted of
watering a pair of seedlings and seeds with 2 L of water with
the help of a self-built irrigation system that released water
over a period of several days (Appendix S1). Overall, we used
five replicated blocks × four treatments × two seedlings and
20 seeds per treatment, with a total of 40 seedlings and 400
seeds (Fig. 1B). The experiment started on September 2011 and
lasted until February 2012.
We recorded seed germination and seedling establishment
monthly beginning in October 2011. For each plot, we recorded
the number of germinated seeds, number of seedlings survival,
and the causes of plant mortality. We also measured shoot length
and leaf number every time plants were visited.
Statistical Analysis
Abiotic measurements were compared using split-plot repeated
measures analysis of variance (ANOVA) using the software

Restoration Ecology 3
Restoration of tropical coastal dunes
Table 1. Effects of the presence of vegetation and the use of mesh on abiotic factors important for seedling survival in the facilitation experiment. Repeated
measures split-plot ANOVA was carried out using Systat 12. For dependent variables, the table reports the F-values and the level of significance. *p ≤ 0.05;
**p ≤ 0.01; ***p ≤ 0.001.
Variation Source Degrees of Freedom (df) Air Humidity
Vegetation 1 270.449***
Vegetation × block 3
Block 3 5.441
Mesh 1 10.417**
Vegetation × mesh 1 5.315
Error (between subjects) 6
Time 2 1.643
Time × vegetation 2 14.27***
Time × mesh 2 5.87**
Time × vegetation × mesh 2 0.235
Error (within subjects) 12
SYSTAT 12. In the restoration experiment, the effects of
coconut mesh, initial irrigation, and thermal scarification on
seed germination were analyzed by factorial block ANOVA
using the software SYSTAT 12. In all figures, y values are pro-
vided as mean ± standard error (SE).
Seedling survival in both experiments was analyzed using a
generalized linear model with binomial error and mixed effects
using log-likelihood ratio test (LRT) implemented by the pack-
age lme4 (Bates et al. 2014). As we used a split-plot experi-
mental design, the error for testing the effect of vegetation was
determined by the interaction between vegetation and block.
For the analysis of plant mortality causes, we followed the
work of Crawley (2007, p 552–556) in using log-linear mod-
els with Poisson error for the analysis of contingency tables, to
analyze treatment effects on the different causes of plant mortal-
ity. After fitting the full model with the interactions between the
factors, factors were removed to test if they significantly affect
the outcome. Seedling survival and plant mortality causes were
analyzed using the statistical software R version 3.0.2 (R Devel-
opment Core Team 2014).
Results
Experiment 1: Facilitation Experiment
All abiotic variables were affected by the presence of vegetation
(Table 1). Air humidity was higher, and soil surface temperature
and soil temperature at 10 cm depth were lower inside the
vegetation (Fig. 2A–C, respectively). The depths of sand burial
increased over time within and outside the vegetation (Fig. 2D).
The presence of vegetation decreased the depth of sand burial,
but could not prevent plants from sand burial.
The presence of coconut mesh increased air humidity and
decreased sand burial, but there were no effects on soil surface
temperature or soil temperature at 10 cm depth (Table 1; Fig. 2).
The interaction between vegetation and mesh was significant
only for sand burial over time, where the presence of mesh
slightly decreased sand burial outside vegetation (Table 1;
Fig. 2D).
Only 11 out of 64 transplanted seedlings (17.2%) survived
until the end of the experiment. The number of seedlings
4 Restoration Ecology
Soil Surface Temperature Soil Temperature (10 cm depth) Sand Burial
28.48** 11.141** 163.187***
1.399 3.47 152.888
1.362 0.922 13.071**
0.294 1.524 1.025
2.637 6.671** 2.528.962***
6.924** 1.094 67.648***
0.045 0.741 0.701
1.348 2.647 4.566*
survival was higher in the presence of vegetation (𝜒 2 = 5.93;
degrees of freedom (df ) = 1, p = 0.014). The presence of mesh
increased seedling survival (𝜒 2 = 4.86; df = 1, p = 0.027), while
the interaction between vegetation and mesh was not signif-
icant (𝜒 2 = 0; df = 1, p = 1.0; Appendix S2), despite the fact
that increased survival due to mesh treatment was only appar-
ent inside vegetation (Fig. 3A). At the end of the experiment, no
plants were alive outside vegetation.
Overall, the main cause of plant death was sand burial (39.6%
of all 53 deaths). Sand burial was considered the cause of
seedling death if plants were completely covered by sand, and
dry or decomposing at the time of inspection. Other causes of
seeding mortality were desiccation (37.7%, i.e. the plant had
dried out at the time of inspection), overrunning by a car (15.1%,
indicated by tire marks next to the seedling), and herbivory
(7.5%, when all plant leaves were eaten, mostly by leafcutter
ants). Because of the low number of survivors, data on plant
growth (shoot lengths and number of leaves) were not further
analyzed. In fact, average plant growth was negative.
Causes of plant mortality were affected by vegetation pres-
ence. Inside vegetation, the most frequent causes of mortality
were sand burial and herbivory, whereas outside vegetation,
desiccation was the main mortality cause, followed by over-
running by cars and sand burial (Fig. 4A, 𝜒 2 = −51.525;
df = 3, p < 0.001). The presence of mesh did not affect neither
the causes of plant death nor was the interaction between
vegetation and mesh significant (in both cases p > 0.05;
Appendix S2).
Experiment 2: Restoration Experiment
For transplanted seedlings, overall survival was only 7.5%.
The number of seedlings survival was positively affected by
the use of initial irrigation although only slightly (𝜒 2 = 4.58;
df = 1, p = 0.032; Fig. 3B). The presence of mesh did not
affect survival (𝜒 2 = 0.437; df = 1; p = 0.51) and the interaction
between irrigation and mesh also did not affect survival (𝜒 2 = 0;
df = 1, p = 0.99). Due to the low number of survivors, we again
did not analyze data on plant growth (shoot lengths or number
of leaves).
 Restoration of tropical coastal dunes

Soil surface temperature (°C)

65 36

34

Air humidity (%) 60 32

30

55 28

26

50 24
ay

ay
ne

ly

ne

ly
Ju

Ju
M

M
Ju

Ju
Soil temperature - 10 cm (°C)

36 50

34 40

Sand burial (cm)

32 30

30 20

28 10

26 0
ay

ay
ne

ne
ly

ly
Ju

Ju
M

M
Ju

Ju
Figure 2. Measurements (± SE) of air humidity (A), soil surface temperature (B), soil temperature at 10 cm depth (C), and sand burial (D) in four
combinations of vegetation and mesh (− +) treatments in a Brazilian coastal sand dune facilitation experiment.

Figure 3. Survival of Canavalia maritima plants in experiment 1 (A) and experiment 2 (B). Boxplots show the number of plants alive at the end of the
experimental period. In experiment 1, seedlings were transplanted (open bars), in experiment 2, survival was assessed for both transplanted seedlings (open
bars) and for seedlings deriving from seeds (full bars) sown in the field.

The number of germinated seeds (12) was very low (3%), but
scarification increased seed germination compared to no scarifi-
cation (Appendix S3). Seed germination was affected neither by
the presence of mesh or irrigation nor by the interaction between
these treatments (Appendix S3).
Survival of seedlings established from introduced seeds was
only 1.5%. Both the main effects of mesh and irrigation were
not significant, but survival was significantly affected by the
interaction between irrigation and mesh (𝜒 2 = 3.916; df = 1,
p = 0.048); while mesh increased survival in the absence of
irrigation, it tended to decrease survival when irrigation was
applied (Fig. 3B).
Desiccation (approximately 54% of all 37 deaths) and sand
burial (approximately 46%) were the most frequent causes of
seedling mortality (Fig. 4B). Neither the presence of mesh,
initial irrigation nor the interaction between these treatments
affected the relative frequency of these two causes of plant
mortality (for all cases p > 0.05, Appendix S2).
Discussion
The restoration techniques applied to the tropical dune crest
areas showed only limited potential to launch restoration. For

Restoration Ecology 5
Restoration of tropical coastal dunes
Figure 4. Causes of mortality in experiment 1 (A) and experiment 2 (B). Box plots show the number of dead plants of Canavalia maritima seedlings for the
different causes of mortality herbivory, desiccation, sand burial, and overrunning by cars. In experiment 1, seedlings were transplanted and in experiment 2,
mortality was assessed for both transplanted seedlings and for seedlings deriving from seeds sown in the field.
example, we found that the use of coconut mesh and initial
irrigation is unlikely to lead to successful restoration. The high
mortality by burial and the low plant recruitment success in our
experiment emphasizes how dune crest restoration can be very
difficult such that management should primarily aim to avoid
the destruction of intact vegetation. Nevertheless, we also found
important lessons for restoration that should be further improved
in future experiments. First, the presence of established vegeta-
tion facilitates seedling survival thus aiding restoration. Second,
scarification might improve Canavalia maritima seed germina-
tion under field conditions. Third, control of sand burial and
of trampling is a prerequisite for successful restoration of such
dune crest areas.
Sand burial has been described as the most important fac-
tor affecting plant community distribution and composition in
coastal dunes (Martínez & Moreno-Casasola 1996; Maun 2004;
Zheng et al. 2005; Yura & Ogura 2006; Gilbert et al. 2008).
Burial is a key factor controlling germination rates, plant estab-
lishment, and survival (Franks & Peterson 2003), and can also
determine plant species zonation, by acting as an environmen-
tal filter in mobile dunes (Gilbert et al. 2008). Although we
found a positive effect of the presence of vegetation and coconut
mesh on soil stabilization, these factors do not appear to have
been sufficiently strong to prevent high mortality rates by sand
burial. Future studies should, systematically, apply available
techniques such as wood-fences or wind barriers to prevent
strong sand movements in dune crest areas (Greipsson 2002).
Cars (buggy) used for touristic activities on these dunes are
a local source of degradation that negatively affects vegetation
cover and regeneration. Overrunning by cars was detected as
an important cause of seedling death in our experiment, even
though it was much less critical than sand burial. Cars might also
reduce seed germination success, because after being overrun by
a car seeds may be buried too deeply (Sykes & Wilson 1990). In
addition, car traffic on dunes is likely to increase soil instability
and sand movement, which in turn, affects the establishment of
colonizing plants (Kutiel et al. 2001; Lemauviel & Rozé 2003).
6 Restoration Ecology
Thereby, buggy rides create a dynamic situation that increases
adverse conditions for dune regeneration. The frequent intru-
sion of buggies into the protected area during our experiment
shows that successful and sustainable restoration must go along
with stronger law enforcement, participatory management, and
environmental education.
Herbivory was not a very frequent cause of plant mortality
and only affected seedlings located within vegetation. Although
herbivores were not observed in action, the damage inflicted on
plants appeared to be caused by leafcutter ants due to the round
holes observed in leaves. Indeed, leafcutter ant colonies were
observed inside vegetation. Bach (2001) excluded herbivorous
insects using cages in a study of coastal dunes in Michigan
and found that insect herbivory influenced plant community
composition due to reductions in herbaceous plants abundance
(Bach 2001). Thus, the herbivory found in our study might, in
the long term, also shape this dune plant community.
Desiccation, on the other hand, was found to be an important
cause of death for seedlings located outside vegetation only,
on bare soil. Because small seedlings can not tap ground water
they are dependent on rain. Even though precipitation during the
study period corresponded to the normal values for the region
(mean of 269 mm/month from February to July 2011; 10-year
average of 258 mm; CPTEC 2012), plants can suffer water stress
due to high temperatures, high insolation, strong winds, and
sand movements. Inside the vegetation, temperature was lower,
and humidity higher than outside vegetation, thus seedlings
experienced less drought stress. In addition, water availability
was higher as indicated by higher soil moisture, which could
be due to shading or hydraulic lift (Zou et al. 2005; Armas et al.
2010; Prieto et al. 2011). However, because most seedlings were
sand buried and died, it remains unclear if they would have
desiccated at a later stage.
Facilitation theory predicts that established plant individuals
can improve further colonization mostly under adverse condi-
tions (Maestre et al. 2003). Indeed, natural conditions for vege-
tation establishment and survival in coastal dunes, in particular

at the crest, are extremely adverse (Greipsson 2002; Franks &
Peterson 2003; Rozé & Lemauviel 2004; Hesp 2004; Hesp &
Martínez 2007). Previous studies have shown that facilitation
can increase soil stabilization, reduce herbivory, and ameliorate
microclimate conditions and nutrient availability contributing
to plant colonization and survival (Bach 2001; Franks 2003;
Martínez & García-Franco 2004). Our results show that the
vegetation established on dune crests improves microclimatic
conditions for other plants, by increasing humidity and decreas-
ing air and soil temperature, and also by decreasing the like-
lihood of sand burial. Similar positive effects have also been
found in other dune systems (Franks 2003; Franks & Peterson
2003; Martínez 2003; Rudgers & Maron 2003). Our results thus
emphasize the potential use of facilitation for restoring degraded
coastal dunes. In fact, all surviving plants of our first experiment
were only found inside vegetation. However, the degraded dune
crests of Jenipabu present such a harsh environment that a very
large number of seedlings would have to be planted to allow for
effective plant establishment.
Coconut mesh was designed to reduce the likelihood of
sand burial and increase soil moisture, thus mimicking possible
positive effects of established vegetation. Although it slightly
improved microclimatic conditions, reduced sand burial and
improved seedling survival, this small amelioration was not
enough to promote vegetation expansion. This emphasizes the
importance of producing new cutting edge techniques to con-
strain sand burial in dune crests that experience strong patterns
of sand movement.
One additional insight of our study was that seed sowing
might be a cheaper and more effective technique for coastal dune
restoration. This is so because the production of greenhouse
seedlings requires plants to be nursed and transported to remote
restoration sites, which makes the whole process more difficult
and expensive. In addition, by germinating in field conditions
plants can allocate more of their seed reserves in deep roots to
avoid future drought. Greenhouse plants are subjected to poor
root allocation due to pot size constraints. Seed scarification
strongly improved germination rates under natural conditions
and can be used as a valuable tool for dune restoration.
Our results show that dune crests are an extremely harsh
environment and that vegetation loss due to buggy rides further
aggravates dune conservation and restoration efforts. Buggy
rides on the Jenipabu dunes are an important economic activity
for the region. Every year thousands of tourists visit this area
and contribute to employment and income generation. Without
restrictions, these car rides may run over established vegetation
jeopardizing conservation and restoration. The implementation
of a management program restricting buggies circulation to
specific low sensitivity areas should help to maintain vegetation
cover, soil stabilization, and ecosystem services provided by
these dunes. The low survival of seedlings shows that restoration
of the dune crest is likely to require a large and expensive effort,
involving effective protection against sand burial, and a large
number of seedlings or seeds. Although we did not make any
cost estimates, it is likely that the successful restoration of the
dune crest requires a large financial effort, not presently afforded
by the local authorities.
Restoration Ecology
Restoration of tropical coastal dunes
Acknowledgments
This work was made possible by a scholarship granted to L.H.T.
by the Brazilian Committee of Higher Education (CAPES).
G.G. received a PQ-CNPq grant. W.W.W. was supported by
PVE program of CNPq, by the Deutsche Forschungsgemein-
schaft (DFG), and by the program TUMBRA, financed by the
Germany Academic Exchange Service (DAAD). The authors
thank the working team of Restoration Ecology Laboratory
(UFRN) F. Marinho, R. Silva, G. Paterno, and G. Mazzochini
for their valuable contributions to this manuscript. We also
thank the Institute for Sustainable Environmental Development
of Rio Grande do Norte (IDEMA-NUC), which allowed the
development of this work at the Environmental Protection Area
of Jenipabu (APAJ).
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Supporting Information
The following information may be found in the online version of this article:
Figure S1. Field site used for collecting seeds of Canavalia maritima in Ponta Negra
beach, Natal/RN, Brazil (A).
Figure S2. Steps to produce the coconut mesh. First, the coconut leaves must be
collected in nature (A).
Figure S3. Schematic representation of the irrigation system made from plastic
bottles. Materials used to construct the dripping equipment (A).
Figure S4. Number of Canavalia maritima germinated seeds (± SE) with thermal
scarification method (plus presoaking treatment) and no scarification in four combina-
tions of water and coconut mesh treatments (− +), allocated outside vegetation for the
restoration experiment.
Table S1. Effects of the use of mesh and water and the thermal scarification method
on the number of germinated seeds in the restoration experiment.
Appendix S1. Material and methods extended.
Appendix S2. Causes of plant mortality in the experiments.
Appendix S3. Effects of scarification on seed germination.
Received: 5 December, 2014; First decision: 14 February, 2015; Revised: 1
December, 2015; Accepted: 1 December, 2015