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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 />

LITERATURE CITED<br />

Aronson, J., C. Floret, E. Le Floc’h, C. Ovalle, and R. Pontanier. 1993.<br />

<strong>Restoration</strong> and rehabilitation of degraded ecosystems <strong>in</strong> arid and<br />

semi-arid lands. I. A view from the south. <strong>Restoration</strong> Ecology<br />

3:8–17.<br />

Aronson, J., C. Floret, E. Le Floc’h, C. Ovalle, and R. Pontanier. 1995.<br />

Restauration et réhabilitation des écosystèmes dégradés en zones<br />

arides et semi-arides. Le vocabulaire et les concepts. Pages 11–29<br />

<strong>in</strong> R. Pontanier, A. M’Hiri, N. Akrimi, J. Aronson, and E. Le Floc’h,<br />

editors. L’homme Peut Il Refaire Ce Qu’il a Défait? John Libbey<br />

Eurotext, Paris.<br />

Bakker, J. P., A. P. Grootjans, M. Hermy, and P. Poschlod. 2000. How to<br />

def<strong>in</strong>e targets for ecological restoration? Introduction. Applied<br />

Vegetation Science 3:3–6.<br />

Boucheron, C. 1987. Aménagement et gestion des dunes littorales. M<strong>in</strong>istère<br />

de l’environnement, <strong>France</strong>.<br />

Daget, P., and J. Poissonet. 1971. Pr<strong>in</strong>cipes d’une technique d’analyse<br />

quantitative de la végétation des formations herbacées. Doc:<br />

CEPE-CNRS 56, 85–100.<br />

Despeyrou, J. L. 1984. La végétation des dunes littorales du golfe de Gascogne,<br />

Université de Bordeaux III Institut de Géographic, <strong>France</strong>.<br />

Duffaud, M. H. 1998. Végétation des dunes littorales Atlantiques de<br />

l’Adour à Noirmoutier. Revue Forestière Française 4:328–348.<br />

Egan, D., and E. Howell. 2000. The historical ecology handbook. A restorationist’s<br />

guide to reference ecosystems. Island Press, Wash<strong>in</strong>gton.<br />

Gadgil, R. L., and F. J. Ede. 1998. Application of scientific pr<strong>in</strong>ciples to<br />

sand dune stabilization <strong>in</strong> New Zealand: past progress and future<br />

needs. Land Degradation and Development 9:131–142.<br />

34 <strong>Restoration</strong> Ecology MARCH 2004


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