# Restoration and rehabilitation of species-rich grassland ecosystems in France: a review

Serge Muller, Thierry Dutoit, Didier Alard, Frédérique Grevilliot

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Serge Muller, Thierry Dutoit, Didier Alard, Frédérique Grevilliot. Restoration and rehabilitation of species-rich grassland ecosystems in France: a review. Restoration Ecology, 1998, 6 (1), pp.94-101. 10.1046/j.1526-100x.1998.06112.x. hal-05471855

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# Restoration and Rehabilitation of Species-Rich Grassland Ecosystems in France: a Review

S. Muller<fnref n="1" />

T. Dutoit<fnref n="2" />

D. Alard<fnref n="3" />

F. Grévilliot<fnref n="1" />

## Abstract

In France the alteration of species-rich grasslands is due mainly to agriculture and civil engineering projects (roadways, dams, ski resorts, etc.). The restoration of such ecosystems has been tested in many regions and after many sorts of degradation. According to the level of initial degradation, different responses can be envisioned. Some variations are presented of a general model of restoration and rehabilitation: (1) rejuvenation of fallow land by grazing with rustic animal breeds in wetlands or chalk grasslands in south and northwestern France, (2) restoration by recovery of extensive agricultural management in intensive agricultural areas such as the Rhône or Meuse Valleys, and (3) rehabilitation by appropriate ecological engineering carried out in grasslands degraded by intensification, ski-track, or civil engineering installations. Despite some positive results, these undertakings have not always had the expected effects and therefore should not provide alibis for the destruction of natural grasslands. In general, the success of such under-

takings depends on the maintenance in the surrounding areas of protected seed source reservoirs and on the persistence of a diversified landscape pattern permitting connectivity between these seed sources and the restoration or rehabilitation sites.

## Introduction

In France, permanent grasslands occupy about 10 million ha and are mainly secondary ecosystems created by centuries of agricultural practices (Pott 1992). The grasslands can present high botanical richness with 30-40 different plant species in an area of only 50 m<fnref n="2" /> (Delpech 1975). Grassland ecosystems have been altered over the past 50 years in various ways (Green 1990), including (1) intensification of agricultural practices such as increased fertilization, liming, drainage, grazing pressure or simple ploughing; (2) land abandonment, which provokes an extension of competitive species and a spontaneous reforestation; and (3) alteration or destruction by civil engineering works, such as the creation of highways, dams, or ski resorts. In France, as in the rest of Europe (Garcia 1992), species-rich grasslands are increasingly impoverished; Delpech (1989) estimated that 300 rare species strictly confined to grasslands are currently threatened.

Restoration or rehabilitation of degraded grasslands have been carried out in order to reconstitute species-rich grasslands (Numata 1992), but different levels of degradation in grassland and meadow influence the choice of interventions required (Bradshaw 1992). A general model has been proposed by Aronson et al. (1993), that can be adapted to this discussion of grassland ecosystems (Fig. 1). Following this approach and the vocabulary of Le Floch & Aronson (1995), several principles can be established: (1) low degradation levels should permit passive restoration; (2) a more important degradation level makes necessary more active intervention aimed at the recovery of functionally similar ecosystems by a rehabilitation process, which can lead to a transitional ecosystem that will then more or less quickly evolve towards the "alternative steady-state," corresponding to the ecosystem present before the human perturbation; (3) if one or more thresholds of irreversibility of degradation have been crossed, a reallocation can be attempted in order to create a new and different kind of grassland ecosystem.

We present a series of French case studies of grassland ecosystem transformations selected to illustrate a range of responses to degradation over a wide geographical range (Fig. 2). For this purpose, we draw upon a bibliographic study and some unpublished studies.

<footnote n="1">¹Laboratoire de Phytoécologie (C.R.E.U.M.), Université de Metz, Ile du Saulcy, F-57045 Metz Cedex 01, France</footnote>

<footnote n="2">²Laboratoire de Biosystématique et Ecologie Méditerranéenne, Université de Provence, FST Saint-Jérôme, URA 1152, case 421 bis, F-13397 Marseille Cedex 20, France</footnote>

<footnote n="3">³Laboratoire d'Ecologie, Université de Rouen, U.F.R. Sciences, F-76821 Mont-Saint-Aignan Cedex, France</footnote>
<figure><img src="image_2.png" /></figure>

Figure 1. General model of grassland ecosystem degradation and the possible pathways of restoration (modified from Aronson et al. 1993).

## Rejuvenation of Species-Rich Grasslands after Land Abandonment

Land abandonment of meadows leads to important modifications of plant communities (Fig. 3). The cessation of mowing or grazing provokes a rapid extension of robust, productive, and highly competitive species, first herbaceous and then woody. Concurrently, their colonization entails the retreat of species that are weaker under direct competition (Prach 1993). This leads to a rapid decrease of plant evenness by competitive dominance, which is followed by local species extinctions. In France, some examples of this process have been studied in central Normandy (Alard et al. 1994) and in the northern Vosges valleys (Muller 1992; Muller et al. 1992).

Several studies have been carried out in France attempting rejuvenation of wet fallow by the use of extensive grazing by a rustic breed such as Highland Cattle (Fig. 4) or Comargue or Konik Polski horses in the Camargue (Bassett 1980; Duncan & d'Herbes 1982; Gordon et al. 1990), the lower Seine River marshes (Lecomte et al. 1981; Lecomte & Le Neveu 1986), the upper Rhône River wetlands (Majchrzak 1992; Morand et al. 1994), and the Brenne marshes (Daudon 1992).

<figure><img src="image_3.png" /></figure>

Figure 2. Distribution of the main grassland rejuvenation, restoration, and rehabilitation experiments in France (modified from Barret 1994).

In a littoral wetland near Trebeurden in Brittany, Rozé (1993) studied successional patterns of reed beds after the reintroduction of extensive grazing by rustic Camargue horses by the permanent line method (20 m) from June 1988 to June 1992. For the three lines, the main results are an increase of species richness (Fig. 5) and a decrease of the dominant species (*Phragmites australis*, *Phalaris arundinacea*, *Calamagrostis epigeios*, *Molinia caerulea*, *Carex acutiformis*, *Cladium mariscus*, etc.).

The rejuvenation of chalk grassland fallow has only rarely been performed in France (Dutoit & Alard 1996) and has usually involved the reintroduction of rustic breeds such as sheep (Arlot & Hesse 1981; Dutoit et al. 1995). Different perturbation regimes involving mowing and ploughing were also tested on chalk grassland over a period of three years (Dutoit 1996). Ten permanent plots (1 m²) were arranged in two rows of five laid
<figure><img src="image_4.png" /><figcaption>Figure 3. Model of grassland rejuvenation after land abandonment.</figcaption></figure>

out in a chalk grassland in the Seine valley in upper Normandy. For each plot, species richness, Shannon index of diversity, and plant evenness were calculated as described by Rozé (1993). Calculation of Shannon's diversity index shows a general increase of species diversity of regularly mown plots (Fig. 6). Differences from

<figure><img src="image_5.png" /><figcaption>Figure 4. Highland cattle used for the rejuvenation and restoration of species-rich grasslands established on a former dumping ground in the wetland of Saint-Martin de Bosherville in the Seine valley. (photo by T. Dutoit.)</figcaption></figure>

<figure><img src="image_6.png" /><figcaption>Figure 5. An example of wet grassland management in a *Phragmites australis* community situation in a littoral wetland near Trébeurden in Brittany. Changes in species richness are recorded along three permanent lines (L1, L2, L3) after the introduction of rustic horses (Camargue) in 1988 (modified from Rozé 1993).</figcaption></figure>

control plots are generally significant from 1992 to 1994. Nevertheless, no significant differences appeared between initial treatments in 1992 and year-to-year changes in species evenness, and in the abandoned plots there was no significant difference between treatments, controls, and year-to-year changes.

Although rejuvenation procedures have just started in France, especially with the partnership of regional conservatories, many such studies have been carried out elsewhere in western Europe (Wells 1969; Willems 1983; Duvigneaud 1983; Hillier et al. 1990). For example, successful rejuvenation of open grassland communities has also been achieved by reducing the importance of competitive species (e.g., *Brachypodium pinnatum*) and stopping shrub and tree encroachment through differ-

<figure><img src="image_7.png" /><figcaption>Figure 6. An example of chalk grassland management in a *Brachypodium pinnatum* community situated in the Seine Valley in upper Normandy. Changes in Shannon index are calculated for 10 quadrats (1 × 1 m square) for control (▲), ploughed (■), or mown (●) plots and with no treatment since 1992; ploughed (□) or mown (○) plots in 1992; and regularly mown in June since this year (modified from Dutoit et al.).</figcaption></figure>
ent kinds of biological management systems (such as extensive grazing by rustic breeds or clearance of scrubs for managing grass-scrub mosaics.

## Restoration of Floristically Rich Meadow Ecosystems by Stopping Intensification of Agricultural Practices

Intensification of agricultural practices, such as liming of acid soils, increasing of fertilization rate, drainage of wet meadows, early cutting of grass, and intensive grazing pressure, generally leads to an increase of grass production (Berendse et al. 1992) but also a drastic impoverishment of botanical and zoological diversity (Delpech 1975; Planteureux et al. 1987; Broyer 1994; Broyer & Prudhomme 1995). Restoration of species-rich grassland communities can be attempted by a passive process (Fig. 7) of stopping the intensification practices (return to lower fertilization, late cutting, more extensive grazing, etc.). This is the principle of article 19 of regulation 797/85 of the European Economic Community (now integrated in the local measures of EEC regulation 2078/92), designed to encourage agricultural methods compatible with environmental concerns and the maintenance of "natural" landscapes. This intensification measure provides subsidies to farmers who accept certain constraints during a period of five years.

In France, this regulation has been applied since 1989 to wet grasslands and since 1993 to chalk grasslands (Dutoit & Alard 1995a). It now concerns about 40 different sites throughout France and a total area of 100,000 ha (Chevalier & Périchon 1993). Table 1 shows the contracts proposed for the Meuse Valley grasslands, where this measure has been applied since 1992 (Grévilliot 1996). Because these types of experiments have just begun, no results of scientific monitoring are yet available. Nevertheless, several Dutch studies (Bakker 1989; Oomes 1990; Berendse et al. 1992) show that even if dry herbaceous biomass decreases rapidly after cessation of fertilization, botanical richness does not increase rapidly. This is linked to the considerable fertilizer reserves in the soil, which must first be used by nutrient-demanding

<figure><img src="image_8.png" /><figcaption>Figure 7. Model of restoration of grasslands impoverished by intensification of management.</figcaption></figure>

species before they will decline, and also to the high density of the meadow vegetation, which inhibits the spontaneous return of new propagules.

## Rehabilitation of Species-Rich Grassland Ecosystems

### Rehabilitation after Intensification of Agricultural Practices

If passive restoration is not successful in the recovery of species-rich grasslands after intensification, rehabilitation must be tried. It has not yet been tested in France as it has in some other European countries (Marrs 1985).

<table><caption>**Table 1.** An example of Article 19 of the European Economic Community regulation: contracts proposed to farmers in the Meuse Valley (from Grévilliot 1996).</caption><thead><tr><th>Level of Contrast</th><th>A</th><th>B</th><th>C</th><th>D</th></tr></thead><tbody><tr><td>Keep in natural grassland</td><td>Yes</td><td>Yes</td><td>Yes</td><td>Conversion of culture in grassland by seeding</td></tr><tr><td>Cutting meadow after the date</td><td>15 July</td><td>01 July</td><td>15 June</td><td>From the second year the area will be exploited like the A, B, or C level</td></tr><tr><td>No use of pesticides and herbicides</td><td>Yes</td><td>Yes</td><td>Yes</td><td>For the first year fertilizers limited to</td></tr><tr><td>Inorganic fertilizers limited to</td><td></td><td></td><td></td><td></td></tr><tr><td>N kg/ha/year</td><td>30</td><td>30</td><td>60</td><td>60 kg/ha</td></tr><tr><td>P kg/ha/year</td><td>15</td><td>15</td><td>30</td><td>30 kg/ha</td></tr><tr><td>K kg/ha/year</td><td>15</td><td>15</td><td>30</td><td>30 kg/ha</td></tr></tbody></table>
Delpech (1991) proposed to seed nutrient-demanding species without destruction of the extant grassland vegetation. Species such as *Lolium multiflorum*, *L. x. hybridum*, or *Zea mays* were found to extract important quantities of fertilizers of the soil. Peeters (unpublished) has started experiments in Belgian meadows on chemical locking of phosphorous by iron oxides in order to increase biodiversity, found to be closely linked with the levels of phosphorus present in the soil. Berendse et al. (1992) studied the effect of removing the topsoil of the grassland. The first year, various ruderal pioneers appeared. They disappeared almost totally in following years. Then they were replaced by typical wet meadow species, including some rare and protected ones such as *Carex hostiana*, *C. pallescens*, and *C. oederi*. Species richness was significantly higher than before sod removal.

Thus stopping fertilization provides, in a few years, a significant decrease of dry matter production but not automatically an increase of plant diversity. This depends on the quantity of fertilizers stocked in the soil, the importance of the seed bank and seed rain coming from neighboring species-rich grasslands, and some perturbations on topsoil which recreate bare areas and therefore allow these seeds to germinate.

## Revegetation as Part of Ski-Station and Civil Engineering Reallocation

Creation of ski tracks often requires new ground-profiling works, entailing important alterations in mountain grassland ecosystems. Rapid reconstitution of the vegetation is desirable, as much for landscape quality as for erosion control (Fig. 8). Many studies along these lines have been carried out in the Alps (Perrussel 1986; Bédécarrats 1988; Bussery 1989; Daburon 1989; Dinger & Antoine 1993; Urbanska 1994, 1995a, 1995b), showing the slowness and irregularity of passive restoration in such an environment (Maman 1985; Perrussel 1986) and the need to sow adapted species (Table 2). This practice leads to the constitution of an artificial meadow (ecosystem of substitution), which can be progressively replaced by native plant communities. For example, Bédécarrats (1991) studied the evolution of such sown meadows and found three main stages in the evolution of these communities (Fig. 9). During the first stage (1-3 years), the sown species and some ruderals dominate. The grassland seems artificial compared to the neighboring spontaneous vegetation; average species richness is about 15 species on a linear sample of 100 points (quadrat point method of Daget & Poissonet 1971). The second stage is transitional, corresponding to colonization by native species such as *Dactylis glomerata*, *Rumex acetosella*, *Trifolium pratense*, *Deschampsia caespitosa*, *Achillea millefolium*, and *Tussilago farfara* at the subalpine level and *Poa alpina* and *Alchemilla vulgaris* at the alpine

<figure><img src="image_9.png" /><figcaption>Figure 8. Model of restoration of mountain grassland after degradation by ski tracks.</figcaption></figure>

level. As a consequence, species richness increases as a whole while the contribution of sown species decreases. Finally, the third stage (6 years and beyond) is a matura-tion stage during which mid-succession species appear, including *Hypericum maculatum*, *Epilobium angustifolium*, and *Leucanthemum vulgare* at low altitudes and *Festuca violacea*, *Carex sempervirens* and *Alchemilla pentaphylla* at higher altitudes. Average species richness rises from 20 in the sixth year to more than 30 in the ninth year. The communities then enter a phase of stabilization and resemble spontaneous grasslands at similar altitudes.

The modifications induced by works such as highways or hydroelectric power stations are generally so drastic that the reconstitution of former ecosystems is

<table><caption>Table 2. Seed mixture used for re-creating grasslands on ski tracks in the Alps (from Dinger 1995).</caption><thead><tr><td>Species and Varieties</td><td>Mixtures % by Weight</td></tr></thead><tbody><tr><td>Phleum pratense Erecta</td><td>20</td></tr><tr><td>Festuca rubra Cibel</td><td>20</td></tr><tr><td>Festuca rubra Echo</td><td>20</td></tr><tr><td>Festuca ovina Bornito</td><td>15</td></tr><tr><td>Lolium perenne Juventus</td><td>15</td></tr><tr><td>Trifolium repens Huia</td><td>10</td></tr></tbody></table>
<figure><img src="image_10.png" /></figure>

Figure 9. Relative contribution of introduced and native species of mountain grasslands after creation of ski tracks (modified from Bédécarrats 1991).

<figure><img src="image_11.png" /></figure>

Figure 10. Model of rehabilitation of ecosystem destroyed by civil engineering works (hydroelectric power station at Chautagne in the Rhône Valley).

no longer possible (Fig. 10). Rehabilitation may be attempted, however, by sowing a different seed mixture in the same way as on ski tracks. For example, Maman (1985) and Coin (1992) studied the evolution of a mixture of five species, *Festuca rubra* (30%), *Phleum pratense* (20%), *Cynosurus cristatus* (20%), *Trifolium repens* (10%), and *Medicago lupulina* (10%) sown in 1980 at the Chautagne power station in the Rhône River valley. Before sowing, the bare substrate was covered by a silt

bed of 20 cm. Seeding was then carried out with the help of a hydroseeder that spreads a mixture of 150 kg seeds, 160 kg organic fixer (cellulose), 50 kg mulch (dehydrated manure), and a chemical fertilizer with nitrogen (20%), phosphorus (30%), and potassium (50%). After eight years, the plant community showed three changes. First, global vegetation cover decreased from 92% in the third year to 72% in the eighth year. Second, the contribution of introduced species decreased from

<figure><img src="image_12.png" /></figure>

Figure 11. A species-rich grassland established on a restored bank of a power station in Gervans (Rhône valley) after 10 years of seeding. This grassland is now colonized by native species such as the conspicuous forb *Helychrysum stoechas*. (Photo by P. N. Frileux.)
93% in 1983 to 46% in 1988; the only species that persisted well after 8 years was *Festuca ruba*. Third, native species largely increased, their overall contribution rising from 7% to 53% between 1983 and 1988 (Fig. 11). Thus as on ski tracks, a spontaneous grassland community progressively replaced the artificially sown mixture. But in this case the newly created grassland was very different from the initial vegetation, much like a creation of a new ecosystem after destruction of a pre-existing one.

## Conclusion

The reconstitution of species-rich grassland ecosystems can be carried out after different sorts of degradation. The maintenance of a high level of resilience permits a passive restoration or, after land abandonment, rejuvenation. A higher level of alteration generally requires more active intervention as in the rehabilitation process. In any case, however, species enrichment results mainly from the germination of new species. Many authors have demonstrated the limits of restoration by the seed bank (Pfadenhauer & Maas 1987; Graham & Hutchings 1988; Bakker et al 1991; Gugerli 1993; Dutoit & Alard, 1995b). Consequently it appears necessary to complement the seed bank with seeds from neighboring grasslands. This seed rain is transported by wind, wild animals, hay-making machinery and grazing cattle (Bakker 1989). Thus, the success of recolonization processes depends on the maintenance of species-rich agricultural meadows and also of linear marginal grassland plant habitats such as road verges, edges, plot boundaries, or even hedgerows (Alard et al. 1994). The ecological networks constituted by patches and corridors improve connectivity between seed reservoirs and restoration plots (Forman & Godron 1986). Restoring species-rich grasslands will depend on the maintenance of landscape diversity by agricultural areas where mainly rejuvenation procedures have to be performed. In contrast, grassland restoration in areas of intensive agriculture or after civil engineering projects in large-scale denuded landscapes appears to be a long-term process.

## Acknowledgments

We thank J. Aronson and E. Le Floch for their helpful comments on the text.

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