# Land use history and botanical changes in the calcareous hillsides of Upper-Normandy (north-western France): new implications for their conservation management

Thierry Dutoit<sup>a,*</sup>, Elise Buisson<sup>a</sup>, Philip Roche<sup>a</sup>, Didier Alard<sup>b</sup>

<sup>a</sup>Université de Provence, Institut Méditerranéen d'Ecologie et de Paléoécologie, UMR-CNRS 6116 IMEP, FST Saint-Jérôme, case 461, F-13397 Marseille Cedex 20, France

<sup>b</sup>Université de Rouen, Laboratoire d'Ecologie, UPRES EA 1293 ECODIV, UFR Sciences, 76821 Mont-Saint-Aignan Cedex, France

## Abstract

Historical studies were made of the changing land use (260–1995) of 14 calcareous hillsides in Upper-Normandy to investigate botanical changes over the last 200 years. Before the French Revolution in 1789, most of these hillsides were used as vineyards mixed with common sheep walks. After the French Revolution, they were distributed among inhabitants for cultivation ( cereals) but they were abandoned in the nineteenth century and grazed again. Our results show that this history is correlated with important changes in the floristic composition of the calcareous hillsides between 1816 and 1995. The species richness of arable weeds and fallow-land species of former cultivated fields or vineyards decrease faster than that of chalk grassland species. The consequences of these long-term dynamics are discussed with regards to the ecological requirements and the rarity status of the species that have disappeared since 1935. We conclude with the need to take account of land use change processes, e.g. the alternation of arable fields and grasslands, to maintain the present species-richness of calcareous hillsides.

Keywords: Chalk grasslands; Arable weeds; Historical ecology; Biological conservation; Upper-Normandy; France

## 1. Introduction

In Upper-Normandy as in the rest of Europe, chalk grasslands can be considered as local hot spots of bio-diversity with regards to their botanical and entomo-logical composition (Prendergast et al., 1993; Wallies DeVries et al., 2002). Since the beginning of the twentieth century, European chalk grasslands have been the subject of numerous ecological studies seeking to explain their great biological richness (Hillier et al., 1990; Dutoit and Alard, 1996). However, as a result of the modern agricultural revolution since World War II, these grasslands have completely lost their former sig-nificance as a food resource for domestic livestock (Willems, 1990). After grazing stops, natural succession

leads to a plagioclimax such as scrub, woodland and/or species-poor coarse grasslands (Bobbink and Willems, 1987). Today, these species-rich grasslands are considerably smaller than they were at the beginning of this century. For example, in England, only 20% of the total area of chalk grasslands existing in 1939 survived until the 1980s with their former botanical richness (Black-wood and Tubbs, 1970; Ratcliffe, 1984).

Nowadays, the ecology of North European chalk grasslands is thoroughly familiar (Smith, 1980; Hillier et al., 1990) and many of them have been managed for conservation purposes since the 1960s (Wells, 1969). Most management plans aim at conserving the flora and fauna of chalk grassland habitats using different regimes of scrub clearing, mowing or grazing (Willems, 2001; Wallies DeVries, et al., 2002). Management plans for these areas are based on the need to protect and preserve endangered or rare species, communities, ecosystems or landscapes (Franklin, 1993). However, rarity
status or degrees of threat are mainly based on present
day inventories and only fragmented information is
available for the long-term dynamics of endangered
species such as orchids (Wells and Willems, 1991) or
certain species like *Gentianella germanica* (Fischer and
Stöcklin, 1997) or *Pulsatilla vulgaris* (Wells and Barling,
1971).

Much research using historical methods has been
already carried out to investigate human effects on the
vegetation of chalk grasslands (Cornish, 1954; Wells et
al., 1976; Gibson and Brown, 1991; Dutoit and Alard,
1995; Poschlod and Wallies DeVries, 2002). These his-
torical studies have revealed previous external con-
straining or enabling influences, as well as the order of
events and system states that have played an important
role in the creation of present chalk grassland vegeta-
tion patterns and dynamics (Poschlod and Bonn, 1998;
Poschlod et al., 1998). Nevertheless, none of these stud-
ies has focused on changes in botanical composition
over several centuries because of a lack of old botanical
inventories. As a consequence, no definitive data are
available on the historical dynamics of calcareous hill-
side species since the beginning of the 19th century
(Sheail, 1982). This lack of information is mainly due
to the lack of precise and regular botanical field data
during the 19th century. Nevertheless, investigations on
land use history and vegetation of chalk grassland
ecosystems provide a background for understanding
the development of the present landscape, which is
essential for any ecological study. Moreover, these
investigations provide perspectives that may be useful
in the preservation of rare plant communities and spe-
cies (Dolman and Sutherland, 1991; Swetnam et al.,
1999).

As in most calcareous areas of continental Europe (Wilmanns, 1997), little is known about the former vegetation of calcareous hillsides before the early Medieval period (260–1000 BC), unlike studies on the downlands of Eastern England (Bush and Flenley, 1987; Bush, 1993). Paleolithic (80 000–9000 BC), Mesolithic (9000–4000 BC) and Neolithic (4000–1800 BC) periods for these continental areas cannot be investigated with paleoecological methods because of a total lack of wetlands and peat bogs suitable for sampling pollen rain and taking paleoecological samples (wood charcoals, macrofossils, etc.) or archaebotanical samples from the calcareous hillsides. Concerning the impacts of sheep grazing, some flocks have been present in Upper-Normandy since 5000 BP (Habasque, 1990) but no data are available on their presence and the grazing systems used on calcareous hillsides before the early Medieval period.

In this study, we used historical ecology methods in
order to correlate past land uses with the present bota-
nical composition of calcareous hillsides (Russel, 1997;
Egan and Howell, 2001). To achieve this objective, our

paper is divided into four parts in order to: (1) identify
the historical trajectory of former land uses in 14 cal-
careous hillsides of the Seine valley from the early
Medieval period to the end of the twentieth century, (2)
correlate the dynamics observed with changes in the
floristic composition of the calcareous hillsides between
1816 and 1995, (3) discuss our results with regards to
the ecological requirements and the rarity status of the
species that have been lost for two centuries and (4)
apply our results to the biological management of man-
made landscapes such as calcareous hillsides.

## 2. Methods

### 2.1. *Study area*

In Upper-Normandy, chalk and calcareous grasslands
occupy the hillsides of the Seine Valley, its affluents
(Risle, Eure) and the edge of the cuesta of the Bray
region (Fig. 1). These grasslands are underlain by Cre-
taceous chalk (Turonian, Senonian), which usually out-
crops at the surface. The soil types developed on these
substrates are Protorendzina and Rendzina under the
grassland communities and Brown Rendzina, Drift
Brown Rendzina and Brown Calcareous soil under
shrub and wood communities (Boullet, 1986). The cli-
mate of the lower Seine valley is temperate, with an
annual rainfall of 800 mm and a mean annual tempera-
ture of 10 °C (Chaïb, 1989).

The vegetation is typical of chalk grassland forma-
tions which are classified under the *Xerobromion* and
*Mesobromion* communities in phytosociology (Boullet,
1986). The south- facing slopes of the Seine Valley and
the Eure Valley are characterised by the phytosocio-
logical *Pulsatillo-Seslerietum* association, while the
north-facing slopes are characterised by the *Festuco-
Seslerietum* association. More details on the phytosocio-
logical organisation of chalk grasslands in Upper-Nor-
mandy can be found in Boullet (1986), De Foucault and
Frileux (1988) and Alard (1990).

In this study, the 14 sites were chosen because they
represented a wide range of the main ecological and
historical variables that characterise the vegetation of
calcareous hillsides in Upper-Normandy (Table 1).
These sites are located in the provinces of Seine-Mar-
itime (7 sites), Eure (6 sites) and in the Ile-de-France
Region, province of Val-d'Oise (1 site), along a climatic
gradient (Fig. 2) revealed by a previous study on the
climate of Upper-Normandy (Chaïb, 1989). A decrease
in xerophilous species between inland sites and the
estuary sites was correlated with an increase in pre-
cipitation and humidity (Liger, 1952; Duckworth et al.,
2000). Four groups of calcareous hillsides were dis-
tinguished according to climatic characteristics and
slope aspects (Table 1).
<figure><img src="image_1.png" /><figcaption>Fig. 1. Location of the 14 study sites in the calcareous hillsides of Upper-Normandy, northwestern France.</figcaption></figure>

## 2.2. Land use and vegetation history

To study the history of past land uses, we used historical ecology methods based on the analysis of printed and manuscript documents (Sheail, 1980).

Former land uses were assessed by analysing French land surveys. These documents provide a map (1/5000 to 1/10 000 scale) indicating the land use of each agricultural plot at the time of survey (around 1820, 1914 and 1990): cropland, pasture, meadow, fallow land or forest. Previous land uses were also investigated using archives and historical manuscripts from the provincial record offices in Seine-Maritime and Eure. Local farmers and villagers were extensively questioned to add hints on the agricultural history of each site investigated.

Old postcards (1900–1950) and pictures (1585–1850) were used to illustrate past agricultural practices on calcareous hillsides. When the precise location where old postcards or pictures had been taken could be identified,

photographs were taken in the same location to show today's scenery on the same tracts (Debussche et al., 1999). Aerial photographs were used to quantify the extent of forest canopy since 1947 (1957, 1963, 1973, 1991). Cassini's map (1750–1830), military survey maps (1810, 1880) and other maps (1930, 1960, 1970, 1990 *Institut National de Géographie*) (1/25 000) with toponyms were also used.

Historical sources were checked against botanical surveys carried out since the beginning of the nineteenth century (old county and regional floras). Regional scientific societies (*Société Linnéenne de Normandie*, *Société Botanique du Nord de la France*, *Société Elbeuvienne des Sciences Naturelles*, *Société des Amis du Muséum des Sciences Naturelles*, etc.), created at the beginning of the nineteenth century, frequently published botanical surveys of calcareous hillsides in their bulletin. At the beginning of the twentieth century, field data were also available in phytosociological surveys undertaken for PhD theses (Allorge, 1922; De Litardière, 1928).
<table><caption>Table 1
Mean characteristics of the 14 calcareous hillides where inventories have been obtained between 1816 and 1995</caption><thead><tr><th>Towns</th><th>Calcareous hillides</th><th>Geographic co-ordinates</th><th>Aspect</th><th>Geological substratum</th><th>Surface (ha)</th><th>Mean annual temperature °C</th><th>Annual rainfall mm/year</th><th>Climatic groups</th><th>Phytosociological appartenance</th><th>Total inventories</th><th>Species richness</th></tr></thead><tbody><tr><td>1 Saint-Vigor-d'Ymonville</td><td>Les Communaux</td><td>0° 22' E, 49° 29' N</td><td>S-SW</td><td>Cenomanian/Turonian</td><td>43</td><td>10.7</td><td>832.7</td><td>1</td><td>Festuco-Seslerietum</td><td>10</td><td>61</td></tr><tr><td>2 Hénouville</td><td>La Côte de la Fontaine</td><td>0° 56' E, 49° 28' N</td><td>S-SW</td><td>Coniacian/Santonian</td><td>26</td><td>12.2</td><td>919.3</td><td>1</td><td>Pulsatillo-Seslerietum</td><td>5</td><td>48</td></tr><tr><td>3 Saint-Léger-du-Bourg-Denis</td><td>La Côte Sainte-Catherine</td><td>1° 10' E, 49° 26' N</td><td>S-SE</td><td>Turonian/Coniacian</td><td>21</td><td>10.1</td><td>854.2</td><td>1</td><td>Pulsatillo-Seslerietum</td><td>6</td><td>72</td></tr><tr><td>4 Bonsecours</td><td>La Roches de Saint-Adrien</td><td>1° 7' E, 49° 26' N</td><td>W-SW</td><td>Turonian</td><td>10</td><td>10.1</td><td>814.2</td><td>1</td><td>Pulsatillo-Seslerietum</td><td>3</td><td>97</td></tr><tr><td>5 Belbeuf</td><td>Les Roches d'Orival</td><td>1° 7' E, 49° 23' N</td><td>W-SE</td><td>Coniacian</td><td>32</td><td>10.1</td><td>804.7</td><td>II</td><td>Pulsatillo-Seslerietum</td><td>11</td><td>107</td></tr><tr><td>6 Orival</td><td>Les Roches d'Orival</td><td>1° 1' E, 49° 20' N</td><td>S-SE</td><td>Senonian</td><td>90</td><td>10.9</td><td>753.9</td><td>II</td><td>Pulsatillo-Seslerietum</td><td>31</td><td>187</td></tr><tr><td>7 Freneuse</td><td>La Côte du Bois-du-Val</td><td>1° 5' E, 49° 19' N</td><td>S-SE</td><td>Coniacian/Santonian</td><td>32</td><td>10.1</td><td>744.2</td><td>II</td><td>Pulsatillo-Seslerietum</td><td>16</td><td>82</td></tr><tr><td>8 Amfreville-sous-les-Monts</td><td>La Côte des Deux-Amants</td><td>1° 16' E, 49° 8' N</td><td>W-NW</td><td>Coniacian</td><td>80</td><td>10.1</td><td>767.5</td><td>III</td><td>Pulsatillo-Seslerietum</td><td>8</td><td>133</td></tr><tr><td>9 Saint-Pierre-du-Vauvray</td><td>Les Vals</td><td>1° 14' E, 49° 13' N</td><td>N-NE</td><td>Campanian</td><td>180</td><td>10.9</td><td>701.7</td><td>III</td><td>Festuco-Seslerietum</td><td>11</td><td>63</td></tr><tr><td>10 Tosny</td><td>La Côte de Tosny</td><td>1° 22' E, 49° 12' N</td><td>S-SE</td><td>Campanian</td><td>40</td><td>11.1</td><td>676</td><td>III</td><td>Festuco-Seslerietum</td><td>9</td><td>53</td></tr><tr><td>11 Les Andelys</td><td>La Côte Saint-Jacques</td><td>1° 23' E, 49° 15' N</td><td>SW</td><td>Coniacian</td><td>43</td><td>11.1</td><td>719.8</td><td>IV</td><td>Astragalo-Seslerietum</td><td>25</td><td>240</td></tr><tr><td>12 Vernon</td><td>La Côte Saint-Michel</td><td>1° 33' E, 49° 5' N</td><td>SW</td><td>Coniacian/Santonian</td><td>90</td><td>10.4</td><td>643.2</td><td>IV</td><td>Astragalo-Seslerietum</td><td>12</td><td>225</td></tr><tr><td>13 La Roche-Guyon</td><td>Les Roches</td><td>1° 40' E, 49° 5' N</td><td>SW</td><td>Santonian/Campanian</td><td>300</td><td>10.8</td><td>669.4</td><td>IV</td><td>Astragalo-Seslerietum</td><td>13</td><td>179</td></tr><tr><td>14 Cocherel</td><td>La Côte aux Brebis</td><td>1° 21' E, 49° 3' N</td><td>SW</td><td>Campanian</td><td>43</td><td>10.4</td><td>643.9</td><td>IV</td><td>Astragalo-Seslerietum</td><td>16</td><td>175</td></tr></tbody></table>

Because the knowledge of each botanist varies, only the species inventoried in several places in Upper-Normandy during the sampling period (1816–1995) were kept. Species found only once in the inventories, with ecological requirements incompatible with the climate of Upper-Normandy (Mediterranean or mountainous species) or the calcareous substrate of hillsides, were considered as determination errors (e.g. confusion between *Cistus* and *Helianthemum* genera).

The former nomenclature used in the old field inventories and county flora was updated using the Synonymic Index of French Flora (Kerguélen, 1998). Then, all plant names were harmonised following Lambinon et al.'s (1993) nomenclature. Between 1816 and 1995, several hundred surveys of chalk grassland vegetation in Upper-Normandy were taken and all these references are available in an electronic database in the Rouen Natural History Museum (Chaïb, 1991). The rarity status of inventoried species were based on the national protected species list for France (Danton and Baffray, 1995) and on the regional protected species list for Upper-Normandy (Chaïb et al., 1991).

Detailed field work could not be undertaken on all 14 calcareous hillside sites in Upper-Normandy. Instead, a picture of each site as well as field notes on topography (altitude, slope, aspect, geological substratum, etc.), ethnobotany (former cultivated species, arable weeds) and agricultural and pastoral practices (strip lynchets, land organisation, etc.) were taken. All vascular species present in homogeneous areas of the 14 chalk grasslands were recorded in June 1995 with additional visits in order to add the species flowering earlier or later in the year.

## 2.3. Data analysis

Certain difficulties may be met when using historical data: a date may not be given for a record, field inventories may be incomplete, looking only at a part of the vegetation (rare species) or at an area bigger than the calcareous hillside sites. Moreover, the quantitative exploitation of these data may not be directly possible because different values may have been used to estimate the extent of each species: rarity indices at site level for local inventories, rarity indices at regional and provincial levels for the regional county floras and values of abundance-dominance for several hundred square metres grassland in phytosociological inventories according to the standard procedure of Braun-Blanquet (Guinochet, 1973). Moreover, the sites investigated may differ in their size and number of botanical inventories, which strongly influences the results for the period considered. All these circumstances require very careful interpretation (Rösch, 1998).

For every inventory and site, the presence of a taxon was noted. Then, three 60-year time periods were
<figure><img src="image_2.png" /><figcaption>Fig. 2. Location of the 14 study sites (●) in the local climate areas of Upper-Normandy defined by Chaïb (1989). (1) Maritime climate, (2) Attenuated maritime climate, (3) Maritime climate with continental impression, (4) Maritime climate with continental impression and meridian influence, (5) Meridian influences.</figcaption></figure>

defined, [A] (1816–1875), [B] (1876–1935) and [C] (1936–1995) between the first botanical inventory available (Le Turquier De Longchamp, 1816) and present day inventories (1995). We chose these three periods to be long enough to collect a significant number of inventories. The botanical inventories belonging to each period were grouped together. The number of inventories and species for each site are given in Table 1. This preliminary evaluation did not take into account that the sites considered differ very greatly in their structure and in the methods used for their botanical investigations in the past. For instance, both the number of taxa inventoried and the number and size of sites investigated differ. However, the exclusion of less intensively investigated sites would have reduced the number of botanical inventories for some periods so much that the calculation of presence values would not have been possible. The number of botanical inventories for the period investigated (180 years) is between five (site 2) and 31 (site 6) (Table 1).

The different plant communities found in the 14 calcareous hillsides and revealed by the botanical inventories realised since 1816 were classified at the rank of “alliance” in the phytosociological classification (Guinochet, 1973) according to their ecological demands and their resulting phytosociological behaviour. Fifteen ecological sub-groups were distinguished (Appendix) based on the previous ecological and phytosociological survey of Bournérias (1984) and personal communications (Pr. J. Lambinon University of Liège, Belgium): However, to simplify matters, only three sets of species were distinguished for the three periods and the four groups of sites. These sets of species are (1) arable weed species (sensu *lato*) which are the sum of sub-groups: *Caucalion* + *Polygono-Chenopodion* + adventive species + species formerly cultivated; (2) fallow-land species (sensu *lato*): *Potentillion* + *Chenopodion* + *Onopordion* + *Sisymbrion* + *Leontodion* and (iii) grassland species (sensu *lato*): *Xerobromion* + *Mesobromion* + *XeroMesobromion* + *Arrhenatherion* + *Koelerion* + *Geranion* (Appendix).
All these groups have been identified (sites, species, time periods) in order to analyse quantitatively the dynamics of species richness for all species groups through time and according to the position of each site along the environmental gradient (identified between the inland sites to the estuary sites). After grouping the species in ecological (15), temporal (3) and geographical groups (4), a correspondence analysis (CA) was performed on the [15 rows × 12 columns] matrix using ADE-4 software (Thioulouse et al., 1997). Because most of our data were not normally distributed, they were analysed with the Kruskall–Wallis test (Statview software) to find an overall difference between the number of inventories realised for each period or group of sites and the species richness of the three species groups.

**3. Results**

**3.1. History of land use and management**

Direct evidence of land use in the 14 study sites is lacking until the early Medieval period, when large tracks of calcareous hillsides were ploughed for vine cultivation. From the early Medieval period to early modern times (1500–1800 AD), they were some vineyards in 13 of our study sites with the exception of the

calcareous hillsides of Saint-Vigor-d'Ymonville (site 1) in the estuary of the Seine River. These hillsides were too close to the marine influence of the Channel (Fig. 2). The maximum vine cultivation in Upper-Normandy occurred from 911 (independence of Normandy) to the middle of the fifteenth century. From the sixteenth century, vine cultivation decreased, mainly due to (1) climatic conditions (little ice age 1600–1800 AD), (2) the poor taste of the product, (3) wine taxes from Charles IX to Louis XV and (4) diseases and predators like the pigeon invasions of the sixteenth century and phylloxera at the end of the nineteenth century. Nevertheless, for the warmest calcareous hillsides like those of La-Roche-Guyon (site 13), vineyards were still grown during all the nineteenth century (Fig. 3).

In that period, tracts of calcareous hillsides were also ploughed up for other crops (wheat, dyes, orchards, etc.) and even the steeper slopes were at times cultivated following the slope (Fig. 4) or in the form of strip lynchets, in order to provide an increasing human population with sufficient food. Old estate maps provide evidence that the plough had, in those times, covered extensive areas of most of the 14 calcareous hillsides studied. The abandonment of arable fields is difficult to date and seems to be spread throughout the nineteenth century depending on problems of cultivation (low fertility, soil dryness, etc.) and on the state of erosion of arable fields, which was relatively rapid because of the

<figure><img src="image_3.png" /><figcaption>Fig. 3. Map showing the existence of numerous plots of vineyards in La-Roche-Guyon (site 13) around 1809 (T. Dutoit, personal collection).</figcaption></figure>
<figure><img src="image_4.png" /><figcaption>Fig. 4. Crop systems in the sense of the steepest slopes of La-Roche-Guyon (site 13) around 1900 (T. Dutoit, personal collection).</figcaption></figure>

steep slopes. This explains the fact that chalk grasslands dated between 100 and 200 years old, and established on former cultivated land, can coexist with very old grasslands which may have been grazed since the Paleolithic Age.

In places where cultivation was strictly impossible (absence of soils, cliffs, etc.), the 14 calcareous hillsides studied were grazed by sheep. During the early Medieval period, flocks of sheep on the calcareous hillsides were constantly tended by a shepherd who moved them about the farm virtually every single day of the year. An essential feature of this type of sheep husbandry was that grazing took place for only a few hours a day on chalk grasslands. During the remaining time the animals were kept in a special kind of stable from which the sheep dung was removed once a year, in early spring, to the arable fields. Thus, meat and wool were not the only important products of sheep husbandry in former days; for a long period of time dung was even more important. The grazing area of these flocks included chalk grasslands and wet meadows but also roadside and river verges, cereal stubble, fallow fields, wet meadow and, especially during winter time, woodlands.

After the French Revolution, common land was abolished and sheep walks possessed by noble families were distributed to the farmers of each parish. During this period, most of the chalk grasslands used as sheep walks were distributed to the inhabitants and cultivated for several decades. As a consequence, important changes in grazing patterns occurred in this period: at the beginning of the nineteenth century, shepherds with large flocks of sheep abandoned itinerant grazing on the common sheep walks. Then, small flocks owned by farmers and tended by their wives or children appeared

in each plot of grassland they possessed. Sheep wool also began to be imported from Australia and New Zealand. As a consequence, sheep breeding in the calcareous hillsides continuously declined during the eighteenth and nineteenth century. At the beginning of the twentieth century, itinerant sheep grazing still existed in most of the calcareous hillsides sampled in this study. While most of the cultivation on calcareous hillsides disappeared at the beginning of the twentieth century (Fig. 5), sheep grazing stopped in the 1960s as a consequence of the modern agricultural revolution. Nowadays, none of the calcareous hillsides studied in this survey is used for agriculture except small private kitchen gardens and small paddocks grazed by a few sheep.

In addition to agricultural practices, the hillsides have also been quarried for chalk at least since the Roman times. There were two types of quarries: underground quarries for blocks for building and open-air quarries for chalk used for the improvement of clay soils. This second type of quarry has provided many habitats (scree, cliffs, shelf, etc.) for calcicolous species. In the early Medieval period, many caves have been transformed into troglodyte dwellings and in the nineteenth century, troglodyte villages were still occupied by humans in the calcareous hillsides of Orival (site 6) and La Roche-Guyon (site 13). This permanent human settlement in the calcareous hillsides also provided a lot of habitats (stone walls, kitchen or ornamental gardens, fallow-land, etc.) for calcicolous species and could be responsible of the introduction of new species (aromatic and ornamental species). Most of the quarries and troglodyte villages were abandoned at the beginning of the twentieth century as the use of cement and concrete became more popular.
<figure><img src="image_5.png" /><figcaption>Fig. 5. (a) A view of the calcareous hillsides of La-Roche-Guyon around 1900 (T. Dutoit, personal collection). (b) The same view taken in 1950 by plane showed the abandonment of cultivation and spontaneous colonization by trees (T. Dutoit, personal collection).</figcaption></figure>

## 3.2. *Botanical changes between 1816 and 1995*

The first two axes of the CA carried out on the 15 group×12 period matrix (Fig. 6) account for more than 73% of total variation, with a contribution of 53.13% for axis 1 (variance 0.2091) and 20.36% for axis 2 (variance 0.0801). The first axis is strongly influenced (Table 2) in its negative part by species of *Polygono-Chenopodion*, *Onopordion*, adventive species, *Potentil-lion* and species formerly cultivated, which were mainly

found in the inventories for the 1816–1875 period in group 3 sites (3A). The positive part of axis 1 is influenced (Table 2) by species of the *XeroMesobromion*, which were inventoried mainly in group 1 during the 1876–1935 period (1B). Axis 2 is more closely correlated (Table 2) in its negative part with species of *Caucalion*, *Sisymbrion* and *Chenopodion* which were mainly inventoried during the 1876–1935 period in group 3 (3B), group 4 (4B) and group 2 (2B). The changes over time of the four groups of sites are represented by arrows in
<figure><img src="image_6.png" /><figcaption>Fig. 6. Correspondence analysis carried out on 15 groups of species (arranged in phytosociological groups at the rank of “Alliance” following Bournérias (1984) and Lambinon (personal communication), inventoried in 4 groups (1-4) of calcareous hillsides in Upper-Normandy during three periods of 60 years (A: 1816-1875, B: 1876-1935, C: 1936-1995). The arrows indicate the temporal direction for each group of sites.</figcaption></figure>

the scatter plot defined by axes 1 and 2 of the CA. The same changes can be observed in groups 2, 3 and 4, dominated by species from cultivated land (*Polygono-Chenopodion, Caucalion*), fallow land (*Onopordion, Sisymbrion, Chenopodion*), mineral substrates like chalk cliffs or stone walls (*Potentillion*) and scree in abandoned quarries (*Leontodion*) during the 1816-1875 and 1876-1935 periods. Between 1936-1995, these sites were dominated by grassland species (*XeroMesobromion, Mesobromion, Arrhenatherion, Geranion*). Only the group 1 sites showed contrasting changes in the opposite direction, which is mainly due to the absence of arable weeds and fallow-land species in the inventories for the first two periods.

The 176 inventories made between 1816 and 1995 were used in this study to estimate botanical changes in calcareous hillside plant communities. Fifty-four per cent of the inventories were made in 1876-1935, 28% in 1936-1995 and 18% from 1816 to 1875. The results of Kruskall-Wallis tests showed a significant difference between the number of inventories realised in each period (H = 10.238 for P < 0.01) which is mainly due to a higher number of inventories realised in 1876-1935 in comparison with the two other periods.

Three-hundred and seventy-nine species were found in our study (Appendix). Among these species, 27% can be considered as arable weeds, 18% as fallow-land species and 55% as grassland species. Because the first period was under-inventoried, the disappearance of a species was only considered from the end of the second period (1935). This means that a species is considered to be lost if not mentioned in an inventory for the last 60 years in the 14 calcareous hillsides studied. Overall, of the 379 species, 37.5% have not been inventoried during the third period. However, this value masks a strong disparity between the three groups of species: there is a disappearance of 64.35% of the arable weed species; 62.70% of the fallow-land species and only 16.60% of the grassland species. For the arable weed group, the adventive species sub-group and *Caucalion* sub-group showed the highest rate of disappearance since 1935: 91 and 74.5%, respectively. For the fallow-land species group, the highest rate of disappearance are among the *Chenopodion* (83.3%) and *Sisymbrion* (81.8%) alliances. Conversely, a very low rate of disappearance was calculated for the grassland species group with only 9% for the *Xerobromion*, 7.8% for the *XeroMesobromion* and 0% for the *Mesobromion*.
Species richness of arable weeds and fallow-land species decreased between the second and the third period (Fig. 7). However, results for the four groups of sites showed no significant differences between the three periods for the species richness of arable weed, fallow-land

<table><caption>Table 2
Factorial contributions of phytosociological groups and geographical-temporal groups on the first two axes extracted from the correspondence analysis. In bold, the highest contributions to axis 1 and 2. See the caption of Fig. 1 for codes</caption><thead><tr><th></th><th>Axis 1</th><th>Axis 2</th></tr></thead><tbody><tr><td><strong>Phytosociological groups</strong></td><td></td><td></td></tr><tr><td>Caucation</td><td>102</td><td><strong>7393</strong></td></tr><tr><td>Adventive species</td><td><strong>8881</strong></td><td>448</td></tr><tr><td>Polygono-Chenopodion</td><td><strong>9471</strong></td><td>1</td></tr><tr><td>Species formerly cultivated</td><td>8148</td><td>101</td></tr><tr><td>Potentillion</td><td><strong>8329</strong></td><td>319</td></tr><tr><td>Chenopodion</td><td>13</td><td><strong>6260</strong></td></tr><tr><td>Onopordion</td><td><strong>9436</strong></td><td>31</td></tr><tr><td>Sisymbrium</td><td>136</td><td><strong>6722</strong></td></tr><tr><td>Leontodion</td><td>3355</td><td>3623</td></tr><tr><td>Xerobromion</td><td>166</td><td>359</td></tr><tr><td>Mesobromion</td><td>2648</td><td>25</td></tr><tr><td>Xeromesobromion</td><td><strong>8579</strong></td><td>731</td></tr><tr><td>Arrhenatherion</td><td>1173</td><td>999</td></tr><tr><td>Geranion</td><td>5940</td><td>897</td></tr><tr><td>Koelerion</td><td>701</td><td>5195</td></tr><tr><td><strong>Geographical-temporal groups</strong></td><td></td><td></td></tr><tr><td>1A</td><td>2461</td><td>531</td></tr><tr><td>1B</td><td><strong>6894</strong></td><td>1588</td></tr><tr><td>1C</td><td>2938</td><td>3297</td></tr><tr><td>2A</td><td>110</td><td>863</td></tr><tr><td>2B</td><td>2080</td><td><strong>4730</strong></td></tr><tr><td>2C</td><td>3597</td><td>1276</td></tr><tr><td>3A</td><td><strong>8864</strong></td><td>1096</td></tr><tr><td>3B</td><td>396</td><td><strong>7629</strong></td></tr><tr><td>3C</td><td>5017</td><td>238</td></tr><tr><td>4A</td><td>3533</td><td>2195</td></tr><tr><td>4B</td><td>1185</td><td><strong>6020</strong></td></tr><tr><td>4C</td><td>1554</td><td>220</td></tr></tbody></table>

<figure><img src="image_7.png" /><figcaption>Fig. 7. Mean values (±1 S.E.) of arable weeds, fallow-land and grassland species for the three periods investigated.</figcaption></figure>

and grassland species. Kruskall-Wallis tests showed significant differences between species richness of arable weeds (H=9.842 for P<0.05) and fallow-land species (H=8.077 for P<0.05) between the four groups of sites. This result is mainly due to a high species richness of arable and fallow-land species in group 4.

# 4. Discussion

## 4.1. Land use history and botanical changes

Many studies on past agricultural practices and their effects on present day vegetation of calcareous hillsides carried out in northwestern Europe (characterised by a man-made landscape) have shown that land use history is the most important factor determining the evolution of the landscape since their abandonment (Dolman and Sutherland, 1991). A clear relationship between past land use and the dynamics of present day vegetation has been identified. Sometimes, natural successions from arable land to grassland take so long that former arable land is still recognisable 130 years after abandonment (Wells et al., 1976; Dutoit and Alard, 1995). Indeed, sites formerly disturbed by cultivation characteristically lack typical and rare calcicolous species and are mainly covered with more common species (Cornish, 1954).

At the plot scale, the enduring effects of past cultivation (ploughing, fertilising, manure), which indicate a major soil disturbance in comparison with sheep grazing, also seems to increase the dominance of tussock grasses and leads to a tall grassland vegetation (Mesobromion, XeroMesobromion, Arrhenatherion). Ploughing has also been considered as destructive and responsible for deflected plant succession after abandonment (Gibson and Brown, 1992), with changes in botanical composition and decrease in species richness (Austrheim and Olsson, 1999) in comparison with very old calcareous grasslands that have been managed by grazing for centuries (Xerobromion).

Despite these results, very few data are available on the changes of the floristic composition in the calcareous hillsides of northwestern Europe because botanical inventories and documentary evidence are missing. As a consequence, no relationship between the history of past land use (sheep grazing, mowing, arable fields, vineyards, etc.) and changes in the floristic composition of chalk grassland plant communities since their abandonment has been demonstrated, except for particular species that clearly indicate past land use, such as *Muscari comosum* for vineyards (Dutoit, 1997; Poschlod and WallisDeVries, 2002) or flagship species for biological conservation like orchids (Wells and Willems, 1991).

Our study, focusing on 14 calcareous hillsides along the Seine and Eure River in northwestern France, confirms that past complex systems of exploitation, combining
different agricultural land uses (cultivation, grazing), industrial land uses (open air quarries and underground quarries) and permanent human settlement (troglodyte villages), have existed as observed in other places in Europe where similar research has been carried out: Belgium (Tihon, 1984; Bruynseels and Vermander, 1984), England (Wells et al., 1976; Smith, 1980), Estonia (Pärtel et al., 1999), Germany (Beinlich and Plachter, 1995; Poschlod and Wallies DeVries, 2002), Netherlands (Hillegers, 1993; Willems, 1990) and Sweden (Rosén and Borgegärd, 1999). Nevertheless, the relationship between changes in land use and the botanical composition throughout the nineteenth and twentieth centuries is obscured by problems in interpreting old data (e.g. botanical inventories).

For example, in our work, the 1876–1935 period is over-represented with a high number of inventories because botanical societies were very active at the end of the nineteenth century, identified as the “naturalist period” in France. Their activity decreased at the beginning of the twentieth century with the First World War (Stott, 1970). Also, some sites were over-inventoried because they were close to some of the members’ houses or because they were attractive for their rare or beautiful species like orchids. In our study, this is particularly the case for group four sites between 1876 and 1935: the biggest site, La Roche-Guyon (300 ha), is easily accessible by public transport from Paris and, as a consequence, well-known by botanists in the nineteenth century (Cosson and De Saint-Pierre, 1861; Dutoit et al., 1999a).

The CA results illustrate the effect of changes in land use on the botanical composition of the four groups of sites from 1816 to 1995. The 1816–1875 period is mainly characterised for group 3 sites by species growing in vineyards (*Polygono-Chenopodion*), kitchen or ornamental gardens (species formerly cultivated), scree (*Leontodion*), stone walls (*Potentillion*) and young fallow lands on sunny slope (*Onopordination*). The 1876–1935 period is characterised for groups 2, 3, and 4 by species growing in calcareous arable fields (*Caucalion*), and dry sandy short grasslands (*Koelerion*). Conversely, the 1936–1995 period is characterised for the same sites by species growing in tall calcareous (*Mesobromion*) or mesophilous (*Arrhenatherion*) grasslands and in zones between scrub and grasslands (*Geranion*). The group 1 sites do not follow this pattern because of the almost total absence of cultivated areas in the nineteenth century shown in the 1876–1935 period by documentary evidence (Dutoit et al., 1994).

This discrimination of the botanical composition of calcareous hillsides from 1816 to 1995 may be linked with the different types of land use and their changes as showed by the documents used during the historical investigation (maps, postcards, prints, documents, etc.) The fact that arable fields were created and rapidly

abandoned during the nineteenth century is confirmed by the increase in arable weeds and fallow-land species between 1816 and 1935, and their subsequent decrease. The increase in grassland species between 1816 and 1995 shows that calcareous hillsides were either constantly grazed or remained as tall grassland dominated by tussock species (e.g. *Brachypodium pinnatum*, *Bromus erecitus*, *Arrhenatherum elatius*) after abandonment until the second part of the twentieth century. Floristic changes between the three periods are less clear statistically for the three groups of species. This may be because of the ecological gradient between the estuary and inland sites (Chaïb, 1991; Liger, 1952). Precipitation and humidity are higher in the lower valley sites (group 1) than in the inland sites (group 2, 3, 4) (Table 1).

Most of the arable weeds originated from the Eastern Mediterranean Basin (Holzner, 1978) and their ecological preference explains that they are not found in group 1 even if crops and vineyards have existed in the nineteenth century as revealed by archives. Small vineyards were cultivated in Bonsecours (site 4) around 1580 near the monastery of Sainte-Catherine, but disappeared in 1594 during the religious war (Dutoit and Chaïb, 1995). These observations are confirmed by statistical analyses showing significantly higher species richness of arable weeds and fallow-land species in the most inland sites (group 4).

## 4.2. *Species lost in the last two centuries*

Our results showed that most of the species that have disappeared in the 14 calcareous hillsides studied since 1935 are arable weed and fallow-land species. These results confirmed those of Dupont (2000), who showed that 67 arable weeds of northern France (Nord-Pas-de-Calais region) have not been observed in arable fields since 1930. Forty of them were only inventoried in the nineteenth century. Thirty of these species were inventoried in our study sites and most of them have not been observed since 1935 either (Appendix). However, the causes of disappearance are different depending on the study. In northern France as in England (McCollin et al., 2000), most of the arable weeds have disappeared because of the development of intensive agricultural systems using chemical amendments since the 1930s (Le Maignan, 1981; Dupont, 1990). Conversely, our results showed that arable weed species have disappeared from the calcareous hillsides of Upper-Normandy because cultivation ceased in the nineteenth century (Dutoit, 1997; Dutoit et al., 2000).

In France, according to Jauzein (1995), about 24% of arable weed species are endangered and 8% are at risk of extinction. Nevertheless, of the 450 species legally protected in France, only 20 species can be considered as arable weed or fallow-land species (Danton and Baf-fray, 1995). In the 14 calcareous hillsides studied, 21
legally protected species have been inventoried (Appendix), two at the national scale and 19 at the regional scale. Only one of these species is considered as an arable weed (*Consolida regalis*) and two are only found on scree (*Iberis linifolia* subsp. *intermedia* and *Viola hispida*). It must be noted that, with the exception of *Consolida regalis*, all the legally protected species have been inventoried after 1935 (Appendix) in the open calcareous grasslands of the *Xerobromion* (43%) and the tall grasslands of *XeroMesobromion* (29%), *Mesobromion* (10%) and *Arrhenatherion* (4%).

Sixty-four per cent of the arable weeds and 63% of the fallow-land species have disappeared from the 14 calcareous hillsides studied since 1935. However, in 1995, 43% of the regional protected species were observed on tall calcareous grasslands established on formerly cultivated plots (*Mesobromion*, *XeroMesobromion*, *Arrhenatherion*) and 43% on open calcareous grasslands grazed for several centuries (*Xerobromion*). Because calcareous grasslands belong to priority habitats for conservation at the European scale (WallisDeVries et al., 2002), their contrasting agricultural histories must be studied as a basis for all management plans. Indeed, changes in agricultural histories alone can sometimes explain the differences in floristic composition, structure and successional processes of human-disturbed plant communities (Dolman and Sutherland, 1991). For the conservation of arable weed communities, successful 'edge programs' (no fertilisation and herbicide applications in a strip of several metres at the edge of arable fields) have been developed in England (Wilson, 2000), Germany (Eggers and Wicke, 2000) and France (Dutoit et al., 1999b).

## 5. Conclusion

Most management plans for conserving calcareous grasslands on calcareous hillsides use different disturbance regimes, e.g. scrub clearing, mowing, grazing

etc. (Willems, 2001). Although these biological management techniques and regimes have succeeded in conserving calcareous grassland plant and animal communities until now (WalliesDeVries et al., 2002), it is clear that most of the calcareous grasslands of northwestern Europe have been cultivated for some time during the nineteenth century. Consequently, the loss of arable weeds we have observed during the two last centuries in the calcareous hillsides of the Seine Valley is probably more than a simple regional trend.

Our results have shown that calcareous grasslands on which restoration usually focuses, are only one possible reference among others recorded along the complex history of these landscapes (Alard, 2002). Nevertheless, their maintenance remains a European priority for such habitats (WalliesDeVries et al., 2002). A considerable shift is probably needed in order to focus more on the historical variability of ecosystem management. Considering that land-use change processes, such as the mosaic-cycle, were practised in traditional man-made landscapes of northwestern Europe, management plans must be based on a more dynamic and “process-oriented” approach (Poschlod personal communication) This also means that grazing is only one of the many possible ways of restoring this historical variability.

## Acknowledgements

We would like to thank the staff of the "Agence Régional de l'Environnement de Haute-Normandie" for their help in finding the old botanical inventories (Sophie Lecuit, Valérie Ménage and Jérôme Chaïb). Many thanks to Cyrille Gréaume for helping us with the data capture. Thanks also to Olivier Chabrerie for the climatic data of the calcareous hillsides of Upper-Normandy, Marjorie Sweetko for proof-reading and Dr. B.N.K. Davis, Professor Dr. P. Poschlod and an anonymous referee for improving the previous version of the manuscript.
Appendix A. Species inventoried in 14 calcareous hillsides of Upper-Normandy from 1816 to 1995

<table><thead><tr><th>Time period</th><th>1816</th><th>1876</th><th>1936</th><th>Time periods</th><th>1816</th><th>1876</th><th>1936</th></tr><tr><th>Species</th><th>1875</th><th>1935</th><th>1995</th><th>Species</th><th>1875</th><th>1935</th><th>1995</th></tr></thead><tbody><tr><td><strong>Caucalion</strong><br>(=arable weed species)</td><td></td><td></td><td></td><td>Adventive species</td><td></td><td></td><td></td></tr><tr><td>Acinos arvensis</td><td></td><td>4</td><td>3</td><td>Alyssum simplex</td><td></td><td>1</td><td></td></tr><tr><td>Adonis aestivalis*</td><td></td><td>1</td><td></td><td>Amaranthus blitum</td><td>1</td><td>1</td><td></td></tr><tr><td>Adonis annua*</td><td></td><td>2</td><td></td><td>Carduus pycnocephalus</td><td></td><td>1</td><td></td></tr><tr><td>Ajuga chamaepitys*</td><td>3</td><td>4</td><td>2</td><td>Diplotaxis viminea</td><td>1</td><td></td><td></td></tr><tr><td>Asperula arvensis*</td><td></td><td>2</td><td></td><td>Eruca sativa</td><td></td><td>2</td><td>1</td></tr><tr><td>Bifora radians</td><td></td><td>1</td><td></td><td>Erucastrum nasturtiifolium</td><td>1</td><td></td><td></td></tr><tr><td>Bromus secalinus*</td><td></td><td>3</td><td></td><td>Glaucium corniculatum</td><td></td><td>1</td><td></td></tr><tr><td>Bunium bulbocastanum*</td><td></td><td>2</td><td></td><td>Lappula squarrosa</td><td>1</td><td>1</td><td></td></tr><tr><td>Bupleurum rotundifolium*</td><td>2</td><td>2</td><td>1</td><td>Lathyrus sativus</td><td></td><td>1</td><td></td></tr><tr><td>Campanula rapunculoides</td><td></td><td>1</td><td>1</td><td>Polycnemum arvense</td><td>1</td><td></td><td></td></tr><tr><td>Caucalis platycarpos*</td><td>2</td><td>3</td><td></td><td>Silene pendula</td><td></td><td>2</td><td></td></tr><tr><td>Chaenorhinum minus</td><td>1</td><td>1</td><td>2</td><td></td><td></td><td></td><td></td></tr><tr><td>Consolida regalis* RP</td><td>1</td><td>2</td><td></td><td>Polygono-Chenopodion<br>(=arable weed species of vineyards)</td><td></td><td></td><td></td></tr><tr><td>Datura stramonium</td><td></td><td>1</td><td></td><td>Allium sphaerocephalon</td><td>3</td><td>4</td><td>7</td></tr><tr><td>Euphorbia exigua</td><td></td><td>1</td><td></td><td>Allium vineale</td><td>2</td><td>2</td><td>4</td></tr><tr><td>Euphorbia falcata</td><td>1</td><td>1</td><td></td><td>Althaea hirsuta*</td><td>2</td><td>3</td><td>1</td></tr><tr><td>Fumaria densiflora</td><td></td><td>4</td><td></td><td>Aristolochia clematitis</td><td>1</td><td>5</td><td>2</td></tr><tr><td>Fumaria officinalis</td><td></td><td>2</td><td></td><td>Calendula arvensis</td><td>2</td><td>4</td><td>1</td></tr><tr><td>Fumaria parviflora</td><td>2</td><td>3</td><td></td><td>Crepis foetida</td><td></td><td>3</td><td></td></tr><tr><td>Fumaria vaillantii</td><td></td><td>2</td><td></td><td>Crepis pulchra</td><td>1</td><td>5</td><td>2</td></tr><tr><td>Galium spurium*</td><td>2</td><td>1</td><td></td><td>Gagea villosa</td><td>1</td><td></td><td></td></tr><tr><td>Galium tricornutum*</td><td></td><td>1</td><td></td><td>Heliotropium europaeum</td><td>2</td><td>3</td><td></td></tr><tr><td>Herniaria glabra</td><td></td><td>3</td><td>1</td><td>Iberis amara*</td><td>3</td><td>1</td><td>1</td></tr><tr><td>Holosteum umbellatum*</td><td>1</td><td>1</td><td></td><td>Lactuca saligna</td><td>2</td><td>2</td><td>1</td></tr><tr><td>Lathyrus aphaca</td><td></td><td></td><td>2</td><td>Muscari atlanticum*</td><td>3</td><td>5</td><td>4</td></tr><tr><td>Lathyrus tuberosus</td><td></td><td>1</td><td></td><td>Muscari botryoides*</td><td></td><td>2</td><td>3</td></tr><tr><td>Legousia hybrida</td><td></td><td>1</td><td>1</td><td>Muscari comosum*</td><td>1</td><td>4</td><td>4</td></tr><tr><td>Legousia speculum-veneris*</td><td></td><td>4</td><td></td><td>Ornithogalum umbellatum*</td><td></td><td>2</td><td></td></tr><tr><td>Lithospermum arvense</td><td>1</td><td>1</td><td></td><td>Physalis alkekengi</td><td></td><td>2</td><td></td></tr><tr><td>Medicago rigidula</td><td></td><td>1</td><td></td><td>Torilis nodosa</td><td></td><td>2</td><td></td></tr><tr><td>Melampyrum arvense*</td><td>2</td><td>5</td><td>3</td><td></td><td></td><td></td><td></td></tr><tr><td>Melampyrum pratense</td><td></td><td></td><td>1</td><td></td><td></td><td></td><td></td></tr><tr><td>Myosotis ramossissima</td><td></td><td>2</td><td></td><td>Species formerly cultivated<br>(=aromatic, ornamental and dye species)</td><td></td><td></td><td></td></tr><tr><td>Nigella arvensis*</td><td>2</td><td>3</td><td></td><td>Antirrhinum majus</td><td>1</td><td>5</td><td>10</td></tr><tr><td>Papaver dubium</td><td></td><td></td><td>1</td><td>Artemisia absinthium</td><td></td><td>1</td><td></td></tr><tr><td>Papaver hybridum</td><td>1</td><td>2</td><td></td><td>Asparagus officinalis</td><td></td><td>2</td><td>3</td></tr><tr><td>Reseda phyteuma</td><td></td><td>1</td><td></td><td>Borago officinalis</td><td></td><td>1</td><td></td></tr><tr><td>Scandix pecten-veneris</td><td></td><td>2</td><td></td><td>Campanula persicifolia</td><td></td><td>1</td><td></td></tr><tr><td>Setaria verticillata</td><td></td><td>1</td><td></td><td>Centranthus ruber</td><td>1</td><td>2</td><td>2</td></tr><tr><td>Setaria viridis</td><td></td><td>1</td><td></td><td>Cheiranthus cheiri</td><td>2</td><td>5</td><td>5</td></tr><tr><td>Sinapis alba</td><td></td><td>2</td><td></td><td>Foeniculum vulgare</td><td>1</td><td>4</td><td>2</td></tr><tr><td>Teucrium botrys*</td><td>2</td><td>4</td><td></td><td>Hyssopus officinalis</td><td></td><td>1</td><td>1</td></tr><tr><td>Thlaspi arvense</td><td></td><td>1</td><td></td><td>Isatis tinctoria</td><td>3</td><td>4</td><td>4</td></tr><tr><td>Thymelaea passerina</td><td>1</td><td></td><td></td><td>Lunaria annua</td><td>1</td><td>2</td><td>1</td></tr><tr><td>Torilis arvensis</td><td></td><td>1</td><td></td><td>Lycium barbatum</td><td></td><td>2</td><td>1</td></tr><tr><td>Vaccaria hispanica*</td><td>2</td><td>3</td><td>2</td><td>Nigella damascena</td><td></td><td>1</td><td></td></tr><tr><td>Valerianella carinata*</td><td></td><td>1</td><td></td><td></td><td></td><td></td><td></td></tr></tbody></table>

(continued on next page)
Appendix A. (continued)

<table><thead><tr><th>Time period</th><th>1816</th><th>1876</th><th>1936</th><th>Time periods</th><th>1816</th><th>1876</th><th>1936</th></tr><tr><th>Species</th><th>1875</th><th>1935</th><th>1995</th><th>Species</th><th>1875</th><th>1935</th><th>1995</th></tr></thead><tbody><tr><td>Valerianella coronata</td><td></td><td>2</td><td></td><td>Onobrychis viciifolia</td><td></td><td>1</td><td></td></tr><tr><td>Valerianella dentata</td><td></td><td>1</td><td></td><td>Papaver setigerum</td><td></td><td>1</td><td></td></tr><tr><td>Veronica arvensis</td><td></td><td>1</td><td></td><td>Papaver somniferum</td><td>1</td><td>2</td><td>1</td></tr><tr><td>Vicia sativa</td><td></td><td>2</td><td>1</td><td>Pseudofumaria lutea</td><td></td><td>2</td><td>1</td></tr><tr><td></td><td></td><td></td><td></td><td>Rubia tinctorum</td><td>1</td><td>1</td><td></td></tr><tr><td></td><td></td><td></td><td></td><td>Ruta graveolens</td><td>1</td><td>2</td><td></td></tr><tr><td></td><td></td><td></td><td></td><td>Salvia officinalis</td><td></td><td>1</td><td></td></tr><tr><td></td><td></td><td></td><td></td><td>Salvia sclarea</td><td>1</td><td>1</td><td></td></tr><tr><td></td><td></td><td></td><td></td><td>Vitis vinifera</td><td>1</td><td>2</td><td></td></tr><tr><td>Potentillion</td><td></td><td></td><td></td><td>Sisymbrium</td><td></td><td></td><td></td></tr><tr><td>(=species of calcareous cliffs and walls)</td><td></td><td></td><td></td><td>(=species of dry trampling areas)</td><td></td><td></td><td></td></tr><tr><td>Dianthus caryophyllus</td><td>2</td><td>3</td><td></td><td>Artemisia campestris</td><td></td><td>2</td><td></td></tr><tr><td>Festuca pallens</td><td>1</td><td>1</td><td>1</td><td>Bromus tectorum</td><td></td><td>3</td><td></td></tr><tr><td>Mycelis muralis</td><td></td><td></td><td>1</td><td>Calamintha nepeta</td><td></td><td>1</td><td></td></tr><tr><td>Parietaria judaica</td><td></td><td>2</td><td>1</td><td>Capsella rubella</td><td>1</td><td>1</td><td></td></tr><tr><td>Podospermum laciniatum</td><td>1</td><td></td><td></td><td>Hypochaeris glabra</td><td></td><td>1</td><td></td></tr><tr><td>Sedum cepaea</td><td>1</td><td>3</td><td></td><td>Lactuca serriola</td><td></td><td>1</td><td></td></tr><tr><td>Sedum dasyphyllum</td><td>1</td><td>3</td><td></td><td>Petrorhagia prolifera</td><td></td><td>4</td><td>1</td></tr><tr><td>Sedum rupestre</td><td>1</td><td>5</td><td>2</td><td>Picris echioides</td><td></td><td>1</td><td></td></tr><tr><td>Sedum sexangulare</td><td></td><td>3</td><td>2</td><td>Polycnemum majus</td><td></td><td>2</td><td></td></tr><tr><td></td><td></td><td></td><td></td><td>Verbascum densiflorum</td><td>1</td><td>3</td><td></td></tr><tr><td></td><td></td><td></td><td></td><td>Verbascum lychnitis</td><td>1</td><td>3</td><td>1</td></tr><tr><td>Chenopodion</td><td></td><td></td><td></td><td>Leontodion</td><td></td><td></td><td></td></tr><tr><td>(=species of nitrophilous walls)</td><td></td><td></td><td></td><td>(=species of calcareous scree)</td><td></td><td></td><td></td></tr><tr><td>Amaranthus blitoides</td><td></td><td>1</td><td></td><td>Galium fleurotii</td><td></td><td></td><td>4</td></tr><tr><td>Amaranthus retroflexus</td><td>1</td><td>3</td><td></td><td>Iberis linifolia subsp. Intermedia RP</td><td></td><td>1</td><td></td></tr><tr><td>Astragalus cicer</td><td></td><td>1</td><td></td><td>Leontodon hyoseroides</td><td></td><td></td><td>6</td></tr><tr><td>Astragalus glycyphyllos</td><td></td><td>3</td><td>2</td><td>Linaria supina</td><td></td><td>2</td><td>5</td></tr><tr><td>Chenopodium glaucum</td><td></td><td>2</td><td></td><td>Rumex scutatus</td><td></td><td>1</td><td></td></tr><tr><td>Chenopodium polyspermum</td><td></td><td>1</td><td></td><td>Sisymbrium supinum</td><td>3</td><td>1</td><td></td></tr><tr><td>Chenopodium vulvaria*</td><td></td><td>2</td><td></td><td>Viola hispida NP</td><td>3</td><td>3</td><td>2</td></tr><tr><td>Conium maculatum</td><td></td><td>1</td><td>1</td><td></td><td></td><td></td><td></td></tr><tr><td>Fumaria muralis</td><td></td><td>2</td><td></td><td></td><td></td><td></td><td></td></tr><tr><td>Herniaria hirsuta*</td><td></td><td>1</td><td></td><td></td><td></td><td></td><td></td></tr><tr><td>Lactuca virosa</td><td></td><td>2</td><td>3</td><td>Xerobromion</td><td></td><td></td><td></td></tr><tr><td>Leonurus cardiaca</td><td></td><td>3</td><td></td><td>(=species of dry calcareous grasslands)</td><td></td><td></td><td></td></tr><tr><td>Lepidium campestre</td><td>1</td><td></td><td></td><td>Aster linosyris RP</td><td>3</td><td>3</td><td>4</td></tr><tr><td>Marrubium vulgare</td><td></td><td>3</td><td></td><td>Astragalus monspessulanus RP</td><td>2</td><td>3</td><td>3</td></tr><tr><td>Nepeta cataria</td><td></td><td>2</td><td></td><td>Biscutella laevigata</td><td>1</td><td>3</td><td>2</td></tr><tr><td>Rumex pulcher</td><td></td><td>1</td><td></td><td>Biscutella neustriaca NP</td><td></td><td>3</td><td>2</td></tr><tr><td>Salvia verticillata</td><td></td><td>3</td><td></td><td>Carex humilis RP</td><td>2</td><td>4</td><td>3</td></tr><tr><td>Sisymbrium irio</td><td></td><td>2</td><td></td><td>Globularia bisnagarica</td><td>1</td><td>9</td><td>7</td></tr><tr><td></td><td></td><td></td><td></td><td>Coronilla minima</td><td>2</td><td>3</td><td>4</td></tr><tr><td></td><td></td><td></td><td></td><td>Euphorbia seguieriana</td><td>4</td><td>6</td><td>7</td></tr><tr><td></td><td></td><td></td><td></td><td>Fumana procumbens</td><td>3</td><td>2</td><td>2</td></tr><tr><td>Onopordion</td><td></td><td></td><td></td><td>Galium glaucum</td><td>1</td><td>3</td><td></td></tr><tr><td>(=species of calcareous fallow lands)</td><td></td><td></td><td></td><td>Helianthemum apenninum</td><td>3</td><td>6</td><td>7</td></tr><tr><td>Campanula rapunculus</td><td></td><td></td><td>1</td><td>Helianthemum canum RP</td><td>2</td><td>5</td><td>5</td></tr><tr><td>Carduus tenuiflorus</td><td></td><td>1</td><td></td><td>Herminium monorchis RP</td><td>1</td><td>3</td><td>1</td></tr><tr><td>Carthamus lanatus</td><td>2</td><td>4</td><td></td><td>Lactuca perennis</td><td></td><td>2</td><td></td></tr><tr><td>Centaurea calcitrapa</td><td>1</td><td>2</td><td></td><td></td><td></td><td></td><td></td></tr><tr><td>Centaurea decipiens</td><td></td><td>1</td><td></td><td></td><td></td><td></td><td></td></tr></tbody></table>

(continued on next page)
Appendix A. (continued)

<table><thead><tr><th>Time period</th><th>1816</th><th>1876</th><th>1936</th><th>Time periods</th><th>1816</th><th>1876</th><th>1936</th></tr><tr><th>Species</th><th>1875</th><th>1935</th><th>1995</th><th>Species</th><th>1875</th><th>1935</th><th>1995</th></tr></thead><tbody><tr><td><strong>Cirsium eriophorum</strong></td><td></td><td>1</td><td></td><td><strong>Linum tenuifolium</strong></td><td>2</td><td>6</td><td>6</td></tr><tr><td><strong>Cynoglossum officinale</strong></td><td>1</td><td>2</td><td>1</td><td><strong>Melica ciliata</strong></td><td>3</td><td>5</td><td>4</td></tr><tr><td><strong>Diplotaxis muralis</strong></td><td>1</td><td>4</td><td>1</td><td><strong>Ononis pusilla RP</strong></td><td>4</td><td>7</td><td>7</td></tr><tr><td><strong>Diplotaxis tenuifolia</strong></td><td>2</td><td>6</td><td>3</td><td><strong>Stipa pennata RP</strong></td><td>1</td><td>2</td><td>2</td></tr><tr><td><strong>Echium vulgare</strong></td><td></td><td></td><td>11</td><td><strong>Teucrium montanum</strong></td><td>1</td><td>8</td><td>13</td></tr><tr><td><strong>Linaria vulgaris</strong></td><td></td><td>1</td><td></td><td><strong>Thlaspi montanum RP</strong></td><td>2</td><td>4</td><td>4</td></tr><tr><td><strong>Melilotus albus</strong></td><td></td><td>2</td><td></td><td><strong>Trinia glauca</strong></td><td></td><td>2</td><td></td></tr><tr><td><strong>Onopordum acanthium</strong></td><td></td><td>1</td><td></td><td><strong>Veronica prostrata</strong></td><td></td><td>2</td><td>1</td></tr><tr><td><strong>Pastinaca sativa</strong></td><td>1</td><td>2</td><td>3</td><td></td><td></td><td></td><td></td></tr><tr><td><strong>Picris hieracioides</strong></td><td>2</td><td></td><td>11</td><td></td><td></td><td></td><td></td></tr><tr><td><strong>Reseda lutea</strong></td><td>1</td><td>5</td><td>7</td><td></td><td></td><td></td><td></td></tr><tr><td><strong>Reseda luteola</strong></td><td></td><td>1</td><td>3</td><td></td><td></td><td></td><td></td></tr><tr><td><strong>Securigera varia</strong></td><td></td><td>2</td><td></td><td></td><td></td><td></td><td></td></tr><tr><td><strong>Silybum marianum</strong></td><td>1</td><td>2</td><td></td><td></td><td></td><td></td><td></td></tr><tr><td><strong>Verbascum pulverulentum</strong></td><td></td><td>3</td><td></td><td></td><td></td><td></td><td></td></tr><tr><td><strong>Verbascum thapsus</strong></td><td></td><td>1</td><td>1</td><td></td><td></td><td></td><td></td></tr><tr><td><strong>Vicia lutea*</strong></td><td>1</td><td></td><td></td><td></td><td></td><td></td><td></td></tr><tr><td><strong>Mesobromion</strong></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td></tr><tr><td><strong>(=species of tall calcareous grasslands)</strong></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td></tr><tr><td><strong>Aceras anthropophorum RP</strong></td><td>1</td><td>3</td><td>2</td><td><strong>Hieracium pilosella</strong></td><td></td><td>6</td><td>12</td></tr><tr><td><strong>Avenula pratensis</strong></td><td></td><td>4</td><td>6</td><td><strong>Himantoglossum hircinum</strong></td><td>1</td><td>6</td><td>9</td></tr><tr><td><strong>Carex flacca</strong></td><td></td><td>4</td><td>10</td><td><strong>Hippocrepis comosa</strong></td><td></td><td>5</td><td>10</td></tr><tr><td><strong>Dactylorhiza fuchsii</strong></td><td></td><td></td><td>1</td><td><strong>Inula salicina</strong></td><td></td><td>1</td><td></td></tr><tr><td><strong>Euphrasia stricta</strong></td><td></td><td>2</td><td>10</td><td><strong>Iris foetidissima</strong></td><td>2</td><td>1</td><td>3</td></tr><tr><td><strong>Orchis simia RP</strong></td><td>1</td><td>3</td><td>3</td><td><strong>Knautia arvensis</strong></td><td></td><td>1</td><td>9</td></tr><tr><td><strong>Orchis ustulata</strong></td><td></td><td></td><td>1</td><td><strong>Koeleria pyramidata</strong></td><td>1</td><td>6</td><td>10</td></tr><tr><td><strong>Orobanche gracilis</strong></td><td></td><td>5</td><td>8</td><td><strong>Leontodon hispidus</strong></td><td></td><td>5</td><td>7</td></tr><tr><td><strong>Orobanche teucrii</strong></td><td>1</td><td>4</td><td>5</td><td><strong>Listera ovata</strong></td><td>1</td><td></td><td>2</td></tr><tr><td><strong>Polygala calcarea</strong></td><td>2</td><td>5</td><td>9</td><td><strong>Medicago falcata</strong></td><td></td><td>3</td><td></td></tr><tr><td><strong>Prunella laciniata</strong></td><td></td><td>4</td><td>1</td><td><strong>Melampyrum cristatum</strong></td><td>1</td><td>4</td><td>1</td></tr><tr><td><strong>Seseli montanum</strong></td><td>1</td><td>4</td><td>2</td><td><strong>Odontites vernus</strong></td><td></td><td>1</td><td>5</td></tr><tr><td><strong>Trifolium ochroleucum</strong></td><td></td><td>1</td><td>1</td><td><strong>Ononis natrix</strong></td><td>3</td><td>7</td><td>6</td></tr><tr><td></td><td></td><td></td><td></td><td><strong>Ophrys apifera</strong></td><td>3</td><td>4</td><td>12</td></tr><tr><td><strong>XeroMesobromion</strong></td><td></td><td></td><td></td><td><strong>Ophrys fuciflora RP</strong></td><td>1</td><td>7</td><td>11</td></tr><tr><td><strong>(=species of calcareous grasslands)</strong></td><td></td><td></td><td></td><td><strong>Ophrys insectifera</strong></td><td>1</td><td>4</td><td>5</td></tr><tr><td><strong>Anacamptis pyramidalis</strong></td><td>2</td><td>7</td><td>12</td><td><strong>Ophrys sphegodes</strong></td><td>3</td><td>6</td><td>10</td></tr><tr><td><strong>Anthericum liliago</strong></td><td></td><td>1</td><td>2</td><td><strong>Orchis purpurea</strong></td><td>2</td><td>4</td><td>11</td></tr><tr><td><strong>Anthericum ramosum</strong></td><td>3</td><td>7</td><td>7</td><td><strong>Orchis militaris</strong></td><td>2</td><td>5</td><td>10</td></tr><tr><td><strong>Anthyllis vulneraria</strong></td><td></td><td>3</td><td>1</td><td><strong>Orchis morio</strong></td><td></td><td>1</td><td></td></tr><tr><td><strong>Asperula cynanchica</strong></td><td></td><td>3</td><td>10</td><td><strong>Orchis ustulata</strong></td><td></td><td></td><td>1</td></tr><tr><td><strong>Avenula pubescens</strong></td><td>1</td><td>5</td><td>9</td><td><strong>Orobanche alba</strong></td><td></td><td>4</td><td>2</td></tr><tr><td><strong>Blackstonia perfoliata</strong></td><td>1</td><td>5</td><td>10</td><td><strong>Orobanche amethystea</strong></td><td></td><td>5</td><td>1</td></tr><tr><td><strong>Brachypodium pinnatum</strong></td><td></td><td>4</td><td>14</td><td><strong>Orobanche lutea</strong></td><td></td><td>1</td><td></td></tr><tr><td><strong>Brassica oleracea</strong></td><td>1</td><td>1</td><td>1</td><td><strong>Orobanche major RP</strong></td><td>1</td><td>1</td><td>2</td></tr><tr><td><strong>Briza media</strong></td><td></td><td>6</td><td>11</td><td><strong>Orobanche minor</strong></td><td></td><td>1</td><td>2</td></tr><tr><td><strong>Bromus erectus</strong></td><td></td><td>5</td><td>12</td><td><strong>Orobanche picridis RP</strong></td><td></td><td>1</td><td></td></tr><tr><td><strong>Bupleurum falcatum</strong></td><td>2</td><td>6</td><td>12</td><td><strong>Phleum phleoides</strong></td><td></td><td>6</td><td>4</td></tr><tr><td><strong>Campanula glomerata</strong></td><td>1</td><td>4</td><td>4</td><td><strong>Plantago media</strong></td><td></td><td>1</td><td>4</td></tr><tr><td><strong>Campanula rotundifolia</strong></td><td></td><td>7</td><td>12</td><td><strong>Platanthera bifolia</strong></td><td></td><td>1</td><td></td></tr></tbody></table>

(continued on next page)
Appendix A. (continued)

<table><thead><tr><th>Time period</th><th>1816</th><th>1876</th><th>1936</th><th>Time periods</th><th>1816</th><th>1876</th><th>1936</th></tr><tr><th>Species</th><th>1875</th><th>1935</th><th>1995</th><th>Species</th><th>1875</th><th>1935</th><th>1995</th></tr></thead><tbody><tr><td>Carex caryophyllea</td><td>1</td><td>3</td><td>9</td><td>Platanthera chlorantha</td><td>1</td><td>4</td><td>7</td></tr><tr><td>Carlina vulgaris</td><td></td><td>4</td><td>11</td><td>Poa bulbosa</td><td></td><td>4</td><td>3</td></tr><tr><td>Centaurea scabiosa</td><td></td><td>5</td><td>10</td><td>Poa compressa</td><td></td><td>2</td><td>6</td></tr><tr><td>Centaurium erythraea</td><td></td><td>2</td><td>4</td><td>Potentilla neumanniana</td><td>1</td><td>5</td><td>6</td></tr><tr><td>Cephalanthera damasonium</td><td>2</td><td>4</td><td>7</td><td>Prunella grandiflora</td><td>1</td><td>5</td><td>3</td></tr><tr><td>Cephalanthera longifolia</td><td></td><td>2</td><td>3</td><td>Pulsatilla vulgaris</td><td>2</td><td>9</td><td>10</td></tr><tr><td>Cephalanthera rubra RP</td><td>1</td><td>2</td><td>1</td><td>Ranunculus bulbosus</td><td></td><td>4</td><td>10</td></tr><tr><td>Cirsium acaule</td><td></td><td>4</td><td>11</td><td>Salvia pratensis</td><td></td><td>3</td><td>4</td></tr><tr><td>Dianthus carthusianorum</td><td>3</td><td>5</td><td>2</td><td>Sanguisorba minor</td><td></td><td>5</td><td>12</td></tr><tr><td>Epipactis atrorubens RP</td><td>2</td><td>6</td><td>10</td><td>Scabiosa columbaria</td><td></td><td>5</td><td>11</td></tr><tr><td>Eryngium campestre</td><td></td><td>3</td><td>8</td><td>Sedum album</td><td></td><td>1</td><td></td></tr><tr><td>Euphorbia cyparissias</td><td></td><td>2</td><td>4</td><td>Senecio erucifolius</td><td>2</td><td>2</td><td>6</td></tr><tr><td>Festuca gr. ovina</td><td></td><td>1</td><td>1</td><td>Serratula tinctoria</td><td>3</td><td>5</td><td>3</td></tr><tr><td>Festuca lemanii</td><td>2</td><td>5</td><td>12</td><td>Seseli annuum</td><td></td><td>3</td><td></td></tr><tr><td>Festuca marginata</td><td></td><td></td><td>1</td><td>Seseli libanotis</td><td>1</td><td>5</td><td>13</td></tr><tr><td>Galium pumilum</td><td></td><td>2</td><td>10</td><td>Sesleria caerulea</td><td>4</td><td>8</td><td>12</td></tr><tr><td>Genista tinctoria</td><td>4</td><td>7</td><td>11</td><td>Silene nutans</td><td>3</td><td>5</td><td>4</td></tr><tr><td>Gentiana cruciata</td><td></td><td>2</td><td></td><td>Spiranthes spiralis</td><td>1</td><td>3</td><td>1</td></tr><tr><td>Gentianella germanica</td><td>1</td><td>5</td><td>7</td><td>Stachys recta</td><td>3</td><td>9</td><td>11</td></tr><tr><td>Gymnadenia conopsea</td><td>2</td><td>7</td><td>10</td><td>Teucrium chamaedrys</td><td>1</td><td>6</td><td>12</td></tr><tr><td>Gymnadenia odoratissima RP</td><td>3</td><td>4</td><td>3</td><td>Thalictrum minus</td><td></td><td>2</td><td>1</td></tr><tr><td>Helianthemum nummularium</td><td>2</td><td>4</td><td>11</td><td>Thesium humifusum</td><td></td><td>6</td><td>6</td></tr><tr><td></td><td></td><td></td><td></td><td>Thymus praecox</td><td></td><td>4</td><td>11</td></tr><tr><td colspan="8" style="text-align: left; font-weight: bold;">Arrhenatherion</td></tr><tr><td colspan="8" style="text-align: left;">(= species of tall mesophilous meadows)</td></tr><tr><td>Achillea millefolium</td><td>1</td><td>3</td><td>10</td><td>Koelerion</td><td></td><td></td><td></td></tr><tr><td>Arrhenatherum elatius</td><td></td><td>2</td><td>5</td><td>(= species of sandy calcareous grasslands)</td><td></td><td></td><td></td></tr><tr><td>Bromus racemosus</td><td></td><td>1</td><td></td><td>Ajuga genevensis</td><td>1</td><td>3</td><td>1</td></tr><tr><td>Centaurea jacea (s.l.)</td><td></td><td>3</td><td>11</td><td>Alyssum alyssoides</td><td>1</td><td>6</td><td>1</td></tr><tr><td>Coeloglossum viride RP</td><td></td><td>1</td><td>1</td><td>Antennaria dioica</td><td>1</td><td>2</td><td>1</td></tr><tr><td>Dactylis glomerata</td><td></td><td>3</td><td>7</td><td>Arabis hirsuta</td><td>1</td><td>5</td><td>9</td></tr><tr><td>Daucus carota</td><td></td><td>1</td><td>7</td><td>Arabis sagittata</td><td></td><td>3</td><td>4</td></tr><tr><td>Euphorbia esula</td><td>2</td><td>8</td><td>7</td><td>Arenaria serpyllifolia</td><td></td><td>3</td><td>3</td></tr><tr><td>Festuca rubra</td><td></td><td>1</td><td>2</td><td>Bothriochloa ischaemum</td><td></td><td>1</td><td></td></tr><tr><td>Galium mollugo (s.l.)</td><td></td><td>3</td><td>10</td><td>Cardaminopsis arenosa</td><td>2</td><td>4</td><td>4</td></tr><tr><td>Galium verum</td><td></td><td>1</td><td></td><td>Cerastium arvense</td><td>1</td><td>2</td><td>1</td></tr><tr><td>Hieracium murorum</td><td></td><td>3</td><td>8</td><td>Cerastium brachypetalum</td><td></td><td>5</td><td>2</td></tr><tr><td>Hypericum perforatum</td><td></td><td></td><td>9</td><td>Cerastium glomeratum</td><td></td><td></td><td></td></tr><tr><td>Leucanthemum vulgare (s.l.)</td><td></td><td>2</td><td>10</td><td>Cerastium pumilum</td><td></td><td></td><td></td></tr><tr><td>Lotus corniculatus</td><td></td><td>6</td><td>13</td><td>Cerastium semidecandrum</td><td></td><td></td><td></td></tr><tr><td>Medicago lupulina</td><td>1</td><td>1</td><td>5</td><td>Erophila verna</td><td></td><td>2</td><td>5</td></tr><tr><td>Ononis minutissima</td><td>1</td><td>1</td><td></td><td>Filago pyramidata *</td><td></td><td>2</td><td></td></tr><tr><td>Ononis repens</td><td></td><td></td><td>4</td><td>Genistella sagittalis</td><td>1</td><td></td><td></td></tr><tr><td>Ononis spinosa</td><td>1</td><td>2</td><td>8</td><td>Hornungia petraea</td><td></td><td>1</td><td>1</td></tr><tr><td>Orobanche caryophyllacea</td><td>2</td><td>3</td><td>3</td><td>Linaria arvensis</td><td></td><td></td><td></td></tr><tr><td>Phyteuma orbiculare</td><td>4</td><td>5</td><td>8</td><td>Linum catharticum</td><td></td><td>4</td><td>12</td></tr><tr><td>Pimpinella saxifraga</td><td></td><td>2</td><td>10</td><td>Medicago minima</td><td></td><td>4</td><td></td></tr><tr><td>Poa pratensis</td><td></td><td>2</td><td>6</td><td>Medicago polymorpha</td><td></td><td>1</td><td></td></tr><tr><td>Polygala vulgaris</td><td></td><td>3</td><td>7</td><td>Minuartia hybrida*</td><td>1</td><td></td><td></td></tr><tr><td>Prunella vulgaris</td><td></td><td>3</td><td>4</td><td>Orobanche purpurea</td><td>1</td><td>4</td><td></td></tr><tr><td></td><td></td><td></td><td></td><td>Polygala comosa</td><td></td><td>1</td><td></td></tr></tbody></table>

(continued on next page)
Appendix A. (continued)

<table><thead><tr><th>Time period</th><th>1816</th><th>1876</th><th>1936</th><th>Time periods</th><th>1816</th><th>1876</th><th>1936</th></tr><tr><th>Species</th><th>1875</th><th>1935</th><th>1995</th><th>Species</th><th>1875</th><th>1935</th><th>1995</th></tr></thead><tbody><tr><td>Rhinanthus alectorolophus</td><td></td><td>2</td><td></td><td>Salvia verbenaca</td><td>1</td><td>4</td><td>2</td></tr><tr><td>Rhinanthus minor</td><td></td><td>1</td><td></td><td>Saxifraga tridactylites</td><td></td><td>3</td><td>3</td></tr><tr><td>Saxifraga granulata</td><td>1</td><td>3</td><td></td><td>Scleropoa rigida</td><td></td><td>2</td><td>5</td></tr><tr><td>Senecio jacobaea</td><td></td><td>2</td><td>7</td><td>Sedum rubens</td><td></td><td>1</td><td>1</td></tr><tr><td>Silene vulgaris</td><td></td><td>4</td><td>7</td><td>Silene conica</td><td></td><td>3</td><td></td></tr><tr><td>Succisa pratensis</td><td></td><td></td><td>4</td><td>Silene otites</td><td>1</td><td>3</td><td></td></tr><tr><td>Teucrium scorodonia</td><td></td><td>2</td><td>7</td><td>Thlaspi perfoliatum*</td><td>4</td><td>7</td><td>4</td></tr><tr><td>Tragopogon pratensis</td><td></td><td>4</td><td>7</td><td>Thymus pulegioides</td><td></td><td>2</td><td>4</td></tr><tr><td>Trisetum flavescens</td><td></td><td>1</td><td>5</td><td>Trifolium scabrum</td><td></td><td>3</td><td></td></tr><tr><td>Verbascum nigrum</td><td>1</td><td></td><td>4</td><td>Trifolium striatum</td><td></td><td>1</td><td></td></tr><tr><td>Veronica teucrium</td><td>1</td><td>7</td><td>7</td><td>Veronica spicata</td><td>3</td><td>4</td><td></td></tr><tr><td></td><td></td><td></td><td></td><td>Vicia lathyroides</td><td>1</td><td>3</td><td></td></tr><tr><td>Geranion</td><td></td><td></td><td></td><td>Vulpia unilateralis</td><td></td><td></td><td>1</td></tr><tr><td>(= species of edges between grasslands and schrubs)</td><td></td><td></td><td></td><td></td><td></td><td></td><td></td></tr><tr><td>Clinopodium vulgare</td><td></td><td></td><td>2</td><td></td><td></td><td></td><td></td></tr><tr><td>Digitalis lutea</td><td>1</td><td>6</td><td>8</td><td></td><td></td><td></td><td></td></tr><tr><td>Geranium sanguineum</td><td>2</td><td>3</td><td>6</td><td></td><td></td><td></td><td></td></tr><tr><td>Inula conyza</td><td></td><td></td><td>7</td><td></td><td></td><td></td><td></td></tr><tr><td>Lathyrus sylvestris</td><td></td><td>3</td><td></td><td></td><td></td><td></td><td></td></tr><tr><td>Melittis melissophyllum</td><td></td><td>2</td><td>5</td><td></td><td></td><td></td><td></td></tr><tr><td>Origanum vulgare</td><td>1</td><td>5</td><td>11</td><td></td><td></td><td></td><td></td></tr><tr><td>Primula elatior</td><td></td><td>3</td><td></td><td></td><td></td><td></td><td></td></tr><tr><td>Primula veris</td><td></td><td>5</td><td>9</td><td></td><td></td><td></td><td></td></tr><tr><td>Rubia peregrina</td><td>1</td><td>5</td><td>7</td><td></td><td></td><td></td><td></td></tr><tr><td>Solidago virgaurea</td><td></td><td></td><td>4</td><td></td><td></td><td></td><td></td></tr><tr><td>Turritis glabra</td><td>1</td><td></td><td>1</td><td></td><td></td><td></td><td></td></tr><tr><td>Vicia tenuifolia</td><td></td><td>1</td><td></td><td></td><td></td><td></td><td></td></tr><tr><td>Vincetoxicum hirundinaria</td><td>1</td><td>5</td><td>7</td><td></td><td></td><td></td><td></td></tr><tr><td>Viola hirta</td><td></td><td>1</td><td>8</td><td></td><td></td><td></td><td></td></tr></tbody></table>

The species have been arranged in phytosociological groups at the rank of “Alliance” following Bournérias, (1984), and Lambinon (personal communication). NP = species protected at the national scale in France (Danton and Baffray, 1995). RP = species protected at the regional scale in Upper-Normandy (Chaïb et al., 1991). *Arable weeds species which have not been inventoried in northern France since 1930 (Dupont, 2000).

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