Sand Dune Restoration in North Brittany, France - Global ...
Sand Dune Restoration in North Brittany, France - Global ...
Sand Dune Restoration in North Brittany, France - Global ...
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<strong>Sand</strong> <strong>Dune</strong> <strong>Restoration</strong> <strong>in</strong> <strong>North</strong> <strong>Brittany</strong>, <strong>France</strong>:<br />
A 10-Year Monitor<strong>in</strong>g Study<br />
Françoise Rozé 1,2 and Servane Lemauviel 1<br />
Abstract<br />
<strong>Dune</strong>s, which account for 13% of the Ille et Vila<strong>in</strong>e north<br />
<strong>Brittany</strong> coast, <strong>France</strong>, were degraded by high tourist<br />
pressure, and they were restored from 1988 onward. Ten<br />
years after commenc<strong>in</strong>g work an assessment of the<br />
restoration was made on three dunes: Les Chevrets,<br />
L’Anse Du Guescl<strong>in</strong>, and Le Verger. Annual monitor<strong>in</strong>g<br />
of the vegetation and dune morphology provided an<br />
opportunity to study the restoration process. The dune<br />
front, which is similar to a reference pioneer dune, lies <strong>in</strong><br />
front of the mobile dune. The highest part of the dune is an<br />
advanced stage of mobile dune. The restoration of the<br />
fixed rear dune was accomplished neither <strong>in</strong> terms of<br />
vegetation composition nor <strong>in</strong> terms of species richness.<br />
The vegetation parameter study allows differentiation<br />
between the dune front, the dune summit, and the rear<br />
dune. The variation <strong>in</strong> species richness and floristic<br />
composition from one zone to another can be expla<strong>in</strong>ed<br />
by abiotic factors such as sal<strong>in</strong>ity and the accretion of sand.<br />
The restoration was satisfactory <strong>in</strong> terms of the geomorphology.<br />
Marram grass is a good tool for restor<strong>in</strong>g the<br />
topography, but it will take a very long time to restore the<br />
conservation value of the dune.<br />
Key words: dune, marram grass, reference ecosystems,<br />
restoration, vegetation.<br />
Introduction<br />
In Ille et Vila<strong>in</strong>e, northern <strong>Brittany</strong>, dunes only account<br />
for 13% of the total coastl<strong>in</strong>e and cover 200 ha (Meur et al.<br />
1992). These are unusual ecosystems that can be def<strong>in</strong>ed as<br />
boundary dunes (Paskoff 1985) or residual dunes (Olson &<br />
Van der Maarel 1989). They lie between two rocky promontories<br />
and have special features because of their small<br />
size.<br />
Western <strong>France</strong>, and particularly <strong>Brittany</strong>, is a region<br />
that has attracted very large numbers of tourists s<strong>in</strong>ce the<br />
Second World War (Guilcher & Hallégouët 1991). The<br />
dunes have been greatly disturbed by factors such as sand<br />
quarry<strong>in</strong>g and erosion caused by trampl<strong>in</strong>g and automobile<br />
traffic.<br />
A restoration program has been implemented on the<br />
dunes of the Ille et Vila<strong>in</strong>e s<strong>in</strong>ce 1988. The protection<br />
works have been conducted by the Ille et Vila<strong>in</strong>e local<br />
council. Wooden fences have been <strong>in</strong>stalled, and marram<br />
grass (Ammophila arenaria) has been planted us<strong>in</strong>g standard<br />
methods (Jacamon 1975; Despeyrou 1984; Boucheron<br />
1987; Klomp 1989; Van der Maarel 1997). In some cases<br />
<strong>in</strong>itial landscap<strong>in</strong>g was conducted (Duffaud 1998). The<br />
study area consisted of three dunes: Les Chevrets,<br />
L’Anse Du Guescl<strong>in</strong>, and Le Verger. The dune morphology<br />
and the vegetation were monitored from 1989. The<br />
1 Université de Rennes 1, Equipe ‘‘Dynamique des communautés,’’ U.M.R.<br />
ECOBIO 6553, Complexe scientifique de Beaulieu, 35042 Rennes cedex,<br />
<strong>France</strong>.<br />
2 Address correspondence to F. Rozé, email francoise.roze@univrennes1.fr<br />
Ó 2004 Society for Ecological <strong>Restoration</strong> International<br />
aim of the restoration was to recreate the dune topography<br />
and return to a landscape <strong>in</strong>clud<strong>in</strong>g a pioneer dune, a<br />
mobile dune, and stabilized dune, with all characteristics<br />
of an undisturbed ecosystem. Ten years after the start of<br />
work a first assessment of the restoration program was<br />
made.<br />
Methods<br />
The study sites are situated along a 5-km length of coastl<strong>in</strong>e<br />
(54°10 0 N, 4°70 0 W). They have the same geomorphological<br />
orig<strong>in</strong> and similar sand. After <strong>in</strong>stall<strong>in</strong>g picket<br />
fenc<strong>in</strong>g around the whole of each dune system without<br />
soil prepar<strong>in</strong>g, the enclosures were immediately planted<br />
with marram grass over their entire area. The plants were<br />
kept <strong>in</strong> place and planted dur<strong>in</strong>g the management day<br />
(April 1988) without other treatments.<br />
The Les Chevrets dune, with an area of about 4 ha, was<br />
heavily degraded before 1988 by a campsite and was<br />
re-landscaped before the start of restoration. The L’Anse<br />
Du Guescl<strong>in</strong> dune, also with an area of about 4 ha, is<br />
bounded to the rear by a road and had a high visitor<br />
pressure. The Le Verger dune has an area of about 2.5 ha<br />
and had been subjected to extensive sand quarry<strong>in</strong>g.<br />
Surveys<br />
At each site two l<strong>in</strong>es measur<strong>in</strong>g 50 m <strong>in</strong> length were set up<br />
from the beach to the rear dune. Three permanent parallel<br />
transects were <strong>in</strong>stalled at right angles to each of these<br />
l<strong>in</strong>es as a function of their topographical position <strong>in</strong> 1989:<br />
one <strong>in</strong> front of the dune, the second at the summit, and the<br />
MARCH 2004 <strong>Restoration</strong> Ecology Vol. 12 No. 1, pp. 29–35 29
<strong>Sand</strong> <strong>Dune</strong> <strong>Restoration</strong> <strong>in</strong> <strong>North</strong> <strong>Brittany</strong>, <strong>France</strong><br />
third <strong>in</strong> the rear dune. Topographical measurements were<br />
made from 1989 to 1998 us<strong>in</strong>g a theodolite. These provided<br />
<strong>in</strong>formation on the accumulation of sand from the beach to<br />
the rear dune.<br />
Every year, <strong>in</strong> June, the vegetation was surveyed us<strong>in</strong>g<br />
the permanent transect method (Daget & Poissonet 1971;<br />
Gloaguen & Touffet 1974). This l<strong>in</strong>ear method was chosen<br />
because it is well suited to the vegetation structure and was<br />
more reliable than measurements on quadrats. One hundred<br />
permanent po<strong>in</strong>ts were distributed at 10 cm <strong>in</strong>tervals<br />
along these transects, whose ends were marked by posts.<br />
The vegetation was recorded as the presence or absence of<br />
each species at each po<strong>in</strong>t. The frequency of the species <strong>in</strong><br />
the 100 po<strong>in</strong>ts provided an estimate of their cover.<br />
Statistical Analysis<br />
The accumulation of sand, the frequency of plant species,<br />
and the species richness of the vegetation <strong>in</strong> the three<br />
zones, predeterm<strong>in</strong>ed by their topography, were compared<br />
<strong>in</strong> each year us<strong>in</strong>g one-way analyses of variance<br />
(ANOVA) (Sokal & Rohlf 1969). Means were compared<br />
us<strong>in</strong>g Tukey tests (Sherrer 1984). The normality of the<br />
data was checked by Ryan–Jo<strong>in</strong>er tests (p 5 90%) and<br />
equality of variances by Levene tests (p 5 95%). The species<br />
richness data were log-10 transformed.<br />
Reference Ecosystems<br />
The aim of the restoration was to re-establish the complete<br />
dune landscape, <strong>in</strong>clud<strong>in</strong>g the plant communities of the<br />
pioneer dune, mobile dune, and fixed dune. The orig<strong>in</strong>al<br />
state was not known precisely, because the degradation<br />
had taken place gradually. S<strong>in</strong>ce the start of this operation,<br />
studies on references <strong>in</strong> restoration ecology (White &<br />
Walker 1997; Egan & Howell 2000) encouraged us<br />
to describe our aims <strong>in</strong> terms of European habitats. The<br />
‘‘Mobile pioneer dunes,’’ adopted <strong>in</strong> version Eur 15 of<br />
the Natura 2000 habitats document (Romao 1996) under<br />
code 2110, is equivalent to the association Euphorbio<br />
3 Agropyretum (Géhu 1994) (association characterized<br />
by Euphorbia paralias and Agropyrum junceum or Elymus<br />
farctus). It is an open, perennial grassland with E. farctus<br />
(Géhu & Tuxen 1975).<br />
The ‘‘Mobile coastal dunes with Ammophila arenaria,’’<br />
listed as code 2120 <strong>in</strong> Romao (1996), is equivalent to the<br />
association Euphorbio 3 Ammophiletum (Géhu 1994)<br />
(characterized by E. paralias and Ammophila arenaria).<br />
This is a community that colonizes dunes after the pioneer<br />
dune stage and that persists with heavy accretion of sand<br />
(Géhu 1969).<br />
The fixed dunes described under the term of ‘‘<strong>North</strong>ern<br />
fixed dunes with a vegetation of Galio-Koelerion albescens,<br />
Corynephorion canescentis’’ (2131) (Romao 1996) belong<br />
to the association Hornungio 3 Tortuletum (Géhu & De<br />
Foucault 1978). Also called gray dunes or mossy dunes,<br />
they are typified by a carpet of bryophytes dom<strong>in</strong>ated by<br />
Tortula ruralis ssp. ruraliformis and colonize relatively<br />
unstable parts of rear dunes (Géhu & De Foucault 1978).<br />
The associations of the pioneer dunes and mobile dunes<br />
(Géhu & Tuxen 1975) and the fixed dune association<br />
(Géhu & De Foucault 1978), described <strong>in</strong> our study area<br />
by detailed phytosociological tables, served as reference <strong>in</strong><br />
our work.<br />
Results<br />
Build-up of the Topography<br />
Figure 1 shows that there was a progressive build-up of the<br />
topography <strong>in</strong> the front and at the summit of the dune. The<br />
very light sand accumulation <strong>in</strong> the rear dune significantly<br />
differentiated it from the front of the dune from 1992 and<br />
from the dune summit from 1993 (Fig. 1; Table 1). The<br />
front of dune and the summit of dune became dist<strong>in</strong>ct <strong>in</strong><br />
terms of sand accumulation from 1993, this be<strong>in</strong>g greater<br />
at the front of the dune than at the summit. This amounted<br />
to 2 m <strong>in</strong> 9 years and could <strong>in</strong> time have formed a new<br />
dune ridge <strong>in</strong> front of the <strong>in</strong>itial summit.<br />
Vegetation Changes by Analysis of the Permanent Transects<br />
The Front of the <strong>Dune</strong>. The front of the dune was dom<strong>in</strong>ated<br />
by two perennial grasses Elymus farctus and Ammophila<br />
arenaria (Fig. 2). From 1993, E. farctus became<br />
dom<strong>in</strong>ant, despite a brief decl<strong>in</strong>e <strong>in</strong> 1998.<br />
The <strong>Dune</strong> Summit. Ammophila arenaria dom<strong>in</strong>ated the<br />
dune summit. Its cover <strong>in</strong>creased until 1995 and then<br />
stabilized at about 60% (Fig. 3). Elymus farctus <strong>in</strong>creased<br />
<strong>in</strong> the early stages of restoration but decl<strong>in</strong>ed from 1993 to<br />
1994. The vegetation became more diverse <strong>in</strong> the later<br />
stages, with the appearance of Festuca arenaria, Calystegia<br />
soldanella, and Vulpia membranacea.<br />
The Rear <strong>Dune</strong>. The rear dune saw the greatest changes<br />
dur<strong>in</strong>g the 10 years (Fig. 4). Ammophila arenaria persisted<br />
Figure 1. Evolution of sand accumulation, regardless of the <strong>in</strong>itial state of<br />
1989, <strong>in</strong> the front dune (F), the dune summit (S), and the rear dune (R).<br />
Values are means and standard errors. Letter <strong>in</strong>dicate the results of the<br />
mean comparison—two different letters <strong>in</strong>dicate a significant difference at<br />
p < 0.05.<br />
30 <strong>Restoration</strong> Ecology MARCH 2004
<strong>Sand</strong> <strong>Dune</strong> <strong>Restoration</strong> <strong>in</strong> <strong>North</strong> <strong>Brittany</strong>, <strong>France</strong><br />
Table 1. Effects of the topographic position (front of the dune, summit,<br />
and rear dune) on the sand accumulation, analyzed with ANOVA.<br />
Year F p<br />
1990 2.04 Not significant<br />
1991 3.39 Not significant<br />
1992 4.01
<strong>Sand</strong> <strong>Dune</strong> <strong>Restoration</strong> <strong>in</strong> <strong>North</strong> <strong>Brittany</strong>, <strong>France</strong><br />
Figure 3. Dynamics of the vegetation <strong>in</strong> the dune summit. Species with cover less than 5% are not represented. Values are means and standard errors.<br />
Figure 4. Dynamics of the vegetation <strong>in</strong> the rear dune. Species with cover less than 10% are not represented. Values are means and standard errors.<br />
elsewhere (Jacamon 1975; Despeyrou 1984; Boucheron<br />
1987; Klomp 1989; Van der Maarel 1997). The result<strong>in</strong>g<br />
dune morphology <strong>in</strong>cluded low dune ridges along the seafront,<br />
followed by flatter areas. This corresponds to the<br />
description of <strong>Brittany</strong> dunes made by Duffaud (1998).<br />
In addition to the geomorphological structur<strong>in</strong>g of the<br />
landscape, the front of the dune, the dune summit, and the<br />
rear dune were dist<strong>in</strong>guished by their plant composition<br />
and species richness.<br />
The plant composition reflected spatial variations <strong>in</strong><br />
environmental factors. The front of the dune is characterized<br />
by strong sal<strong>in</strong>ity constra<strong>in</strong>ts, <strong>in</strong>dicated by both the<br />
dom<strong>in</strong>ance of lyme grass and the scarcity of marram grass<br />
that is much less tolerant of salt stress (Boucheron 1987).<br />
Marram grass is strongly related to sand accretion (Jungerius<br />
& Van der Meulen 1997). Its loss of vigor <strong>in</strong> the rear<br />
dune can be expla<strong>in</strong>ed by the low rates of sand accumulation<br />
that were recorded. In the rear dune the constra<strong>in</strong>ts<br />
Table 2. Effects of the topographic position (front of the dune, summit, and rear dune) on the cover and the number of species, analyzed with ANOVA.<br />
Cover<br />
Number of Species<br />
Year F p F P<br />
1989 1.41 Not significant 0.95 Not significant<br />
1990 1.4 Not significant 0.95 Not significant<br />
1991 0.57 Not significant 1.07 Not significant<br />
1992 0.98 Not significant 17.3
<strong>Sand</strong> <strong>Dune</strong> <strong>Restoration</strong> <strong>in</strong> <strong>North</strong> <strong>Brittany</strong>, <strong>France</strong><br />
Figure 5. Evolution of the vegetation cover <strong>in</strong> the front dune (F), the dune summit (S), and the rear dune (R). Values are means and standard errors.<br />
related to be<strong>in</strong>g close to the sea and the <strong>in</strong>stability of the<br />
soil decrease. Less specialist species appear, and the<br />
species richness <strong>in</strong>creases.<br />
The landscape units identified <strong>in</strong> 1988 changed little <strong>in</strong><br />
their positions, but the restoration operation led to an<br />
advance of the dune ridge toward the sea. At the dune<br />
fronts, the pioneer dunes, similar to those described <strong>in</strong><br />
Géhu and Tuxen <strong>in</strong> 1975, lay <strong>in</strong> front of the mobile dune<br />
and reta<strong>in</strong>ed a high cover of marram grass that was related<br />
to the <strong>in</strong>itial plantation. The dune summits corresponded<br />
to the advanced dynamic stage of mobile dunes as<br />
described by Géhu and Tuxen <strong>in</strong> 1975. They became<br />
enriched with species that frequently occur <strong>in</strong> fixed<br />
dunes. Both <strong>in</strong> terms of diversity and structure, the rear<br />
dune rema<strong>in</strong>ed clearly different from the association<br />
def<strong>in</strong>ed by Géhu and De Foucault (1978). Tortula ruralis<br />
ssp. ruraliformis was present but with a low cover that did<br />
not result <strong>in</strong> the formation of a moss and lichen carpet,<br />
characteristic of fixed dunes. A species richness of about<br />
13 species <strong>in</strong> a few square meters was described by<br />
Géhu (1997), but species richness did not exceed 11<br />
species on average on all our transects. At this part of<br />
the dune profile, the floral composition greatly depends<br />
on the local context. The occurrence of some nitrophilous<br />
plants that do not belong to the reference association<br />
reflects the strong human <strong>in</strong>terference <strong>in</strong> the past.<br />
The lack of species typical of fixed dunes can be<br />
expla<strong>in</strong>ed by the depleted seed bank after many years<br />
of degradation and isolation from seed sources, that is,<br />
<strong>in</strong>tact fixed dunes.<br />
For a long time the aims of dune management were<br />
concentrated on fix<strong>in</strong>g mov<strong>in</strong>g sand. The proximity of<br />
human economic <strong>in</strong>terests made it essential to control<br />
large-scale geomorphological phenomena and, especially,<br />
the drift<strong>in</strong>g of sand <strong>in</strong>to <strong>in</strong>land areas. The fix<strong>in</strong>g of dunes is<br />
now well controlled, but a new challenge has come to the<br />
fore: the conservation of the natural character (Gadgil &<br />
Ede 1998). In dune restoration Sab<strong>in</strong>o et al. (1993) dist<strong>in</strong>guished<br />
two objectives: the restoration of the dune profile<br />
and the restoration of the orig<strong>in</strong>al vegetation. Monitor<strong>in</strong>g<br />
the restoration processes on the dunes of the Ille et Vila<strong>in</strong>e<br />
showed that these two mechanisms are <strong>in</strong>dependent. The<br />
implementation of restoration measures quickly led to a<br />
good plant cover <strong>in</strong> all the landscape zones. On the other<br />
hand, the conservation value and diversity only returned<br />
slowly and the rate of return varied from one site to<br />
another.<br />
Accord<strong>in</strong>g to the term<strong>in</strong>ology used by Aronson et al.<br />
(1993, 1995), the return to reference ecosystems <strong>in</strong>volves a<br />
rehabilitation <strong>in</strong> which the succession passes through a<br />
transitory ecosystem: a plantation of marram grass.<br />
Marram grass provides a quick soil cover and builds up<br />
the dune height by trapp<strong>in</strong>g sand (Kühnholtz-Lordat<br />
1923). It is very effective at restor<strong>in</strong>g dune ridges, but it<br />
seems to slow the vegetation succession <strong>in</strong> the rear dune.<br />
This phenomenon was reported by Lemauviel (2000) and<br />
Lemauviel et al. (2003) <strong>in</strong> <strong>France</strong>, by Webb et al. (2000) <strong>in</strong><br />
Australia, and by Poulson and McClung (1999) <strong>in</strong> Indiana<br />
with the American species, Ammophila breviligulata. The<br />
presence of marram grass is undesirable <strong>in</strong> a semifixed or<br />
Figure 6. Evolution of the species richness (SR) <strong>in</strong> the front dune (F), the dune summit (S), and the rear dune (R). Values are means and standard errors.<br />
Letters <strong>in</strong>dicate the results of the means comparison—two different letters <strong>in</strong>dicate a significant difference at p < 0.05.<br />
MARCH 2004 <strong>Restoration</strong> Ecology 33
<strong>Sand</strong> <strong>Dune</strong> <strong>Restoration</strong> <strong>in</strong> <strong>North</strong> <strong>Brittany</strong>, <strong>France</strong><br />
Table 3. Analysis of the species composition along the permanent transects compared to the reference system.<br />
Reference System<br />
Permanent Transects Géhu and Tuxen (1975) Géhu and De Foucault (1978)<br />
Front of <strong>Dune</strong> <strong>Dune</strong> Summit Rear <strong>Dune</strong> Pioneer <strong>Dune</strong> Mobile <strong>Dune</strong> Fixed <strong>Dune</strong><br />
Elymus farctus X X X X X<br />
Ammophila arenaria X X X X X X<br />
Eryngium maritimum X X X X<br />
Calystegia soldanella X X X X X X<br />
Euphorbia paralias X X X X<br />
Leontodon taraxacoides X X X X<br />
Festuca rubra ssp. arenaria X X X X<br />
Homalothecium lutescens X X X<br />
Tortula ruralis ssp. ruraliformis X X X<br />
Phleum arenarium X X X<br />
Ononis repens X X X<br />
Vulpia membranacea X X X<br />
Cerastium diffusum X X<br />
Carex arenaria X X<br />
Plantago lanceolata X X<br />
Conyza canadensis X X<br />
Bromus rigidus X X<br />
Sonchus oleraceus<br />
X<br />
Lagurus ovatus X X<br />
Diplotaxis tenuifolia<br />
X<br />
Senecio jacobaea<br />
X<br />
Geranium dissectum<br />
X<br />
Elymus repens<br />
X<br />
Honkenya peploides X X<br />
Raphanus raphanistrum<br />
X<br />
fixed dune, where the preservation of conservation value is<br />
paramount. This plant helps ma<strong>in</strong>ta<strong>in</strong> <strong>in</strong>stability by trapp<strong>in</strong>g<br />
sand and by disturb<strong>in</strong>g the soil by the movements of<br />
its leaves. Its presence <strong>in</strong>dicates that the soil is not sufficiently<br />
stabilized.<br />
These restoration procedures were fully justified,<br />
because, as recommended by Hobbs and Norton (1996),<br />
they resulted <strong>in</strong> a vegetation cover on what previously had<br />
been bare soil and gave a conservation value to protected<br />
sites by combat<strong>in</strong>g degradation factors and visitor<br />
pressure. However, a return to a reference state (sensu<br />
Aronson et al. 1993, 1995) has not yet been completed.<br />
At the landscape scale the structures and the communities<br />
of pioneer dunes and mobile dunes have been restored. On<br />
the other hand, at the ecosystem scale, although the<br />
pioneer and mobile dunes have been restored, the fixed<br />
dune restoration is not complete but is <strong>in</strong> a correct<br />
trajectory toward restoration. Cont<strong>in</strong>ued monitor<strong>in</strong>g will<br />
be needed.<br />
Acknowledgments<br />
We are grateful to J. Prod’homme for his participation.<br />
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34 <strong>Restoration</strong> Ecology MARCH 2004
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