Journal of Biogeography (J. Biogeogr.

) (2004) 31, 145157

ORIGINAL ARTICLE

The role of landscape history and persistent biogeographical patterns in shaping the responses of Mediterranean land snail communities to recent re disturbances
de ric Magnin and Franck Torre Laurence Kiss*, Fre

diterrane en dEcologie et de Institut Me oe cologie, U.M.R. 6116 du C.N.R.S., Pale des Sciences et Techniques de St Faculte ro me, France. Je

ABSTRACT

Aims To assess the impact of various re regimes over the past 30 years on land snail communities and to analyse the role of recent landscape history and the inuence of biogeography in shaping the response patterns of gastropod communities following disturbances by re. Location South-eastern France (Provence) and Mediterranean region. Methods Stratied sampling within 12 sites was undertaken with regard to re regime (i.e. number of res, re intervals and age of the last re) occurring over the past 30 years. The study was complemented by a historical analysis using aerial photographs, old maps of vegetation cover and an analysis of the biogeographical composition of malacofaunas. Data were investigated using Correspondence Analysis and Srensen coefcient of similarity. Results When a disturbance regime (land use or re disturbances) has been maintained over decades or centuries, land snail communities appear highly modied and tend to be composed of only Mediterranean and xerophilous species. However, low re regimes, since the 1970s, do not seem to greatly affect the composition of gastropod communities. Indeed, shade-loving, mesophilous and European range species persist even after successive res within some sites. In addition, the malacofaunas have a higher component of European range species with increasing distance from the Mediterranean sea. Main conclusions Analysis of the response patterns of gastropod communities to re shows a response to numerous different factors. The composition of current land snail communities is not only the result of (more or less) recent patterns of re regimes but also of anthropogenic disturbances, of landscape changes over the last centuries and of subsequent structure of the pre-re habitat, as well as of the inuence of a biogeographical gradient. However, the response patterns observed and the persistence of pre-re communities imply the presence of cryptic refuges located within burned areas. Keywords Land snail communities, re regime, response patterns, biogeographical gradient, landscape history.

*Correspondence: Laurence Kiss, IMEP, timent Villemin, Domaine du Petit Arbois, ba Avenue Louis Philibert, BP 80 Cerege 13545, Aix-en-Provence, Cedex 04, France. E-mail: laurence.kiss@univ.u-3mrs.fr

INTRODUCTION Fire is a major disturbance within Mediterranean ecosystems and greatly affects their landscapes (Naveh, 1974; Trabaud, , 2001). In 1976; Blondel, 1995; Whelan, 1995; Lloret & Mar
2004 Blackwell Publishing Ltd www.blackwellpublishing.com/jbi

south-eastern France (Provence), wildre is an obvious problem due to climate conditions (strong wind, high temperatures and drought during summer), ammability of the vegetation and recent land abandonment (Sousa, 1984; rou, 1987; Trabaud, 1987; Amouric, 1992; Whelan, Le Houe 145

L. Kiss et al. 1995). Throughout the Mediterranean basin, re regimes are also related to human intervention (Naveh, 1974) dating to Neolithic times (Guillerm & Trabaud, 1980; Mylonas, 1984; , 2001). Pons & Thinon, 1987; Carcaillet, 1998; Lloret & Mar Since the beginning of the 20th century, land abandonment and decrease in pastoral activities have increased garrigue (calcareous matorral) and forests, and increasing build-up of fuel has led to uncontrolled wildres (Barbero et al., 1987; rou, 1987). Le Houe Although numerous studies treat the impact of re regime on vegetation (Barbero et al., 1987; Trabaud, 1987; Whelan, az-Delgado & Pons, 2001; Vila ` et al., 2001), this 1995; D impact on fauna within Mediterranean ecosystems has been little studied and mainly for the American and Australian continents (Whelan, 1995; Huff & Smith, 2000; Lyon et al., 2000). Previous studies on the impact of re on Mediterranean land snail communities have demonstrated that various response patterns of communities have been obtained for the same re regime (Kiss & Magnin, 2002, 2003). Elements of pre-re gastropod communities seem to persist within post-re communities, whatever the age of the re (Kiss & Magnin, 2002, 2003). Other factors are likely to explain to some extent the malacofauna composition after re. Indeed, recent landscape history (date of land abandonment, type of land use, etc.) is heterogeneous throughout Provence (Amouric, 1992) which, in addition, includes slightly different Mediterranean bioclimates (Emberger, 1971). The main objective of the present study is to understand the variability in response patterns of land snail communities to re disturbance. The aims of this study are thus not only to analyse the impact of various re regimes on land snail communities over the past 30 years, but also to estimate the role of recent site history and the inuence of biogeographical gradient, within a priori a homogeneous area, in shaping the response patterns of gastropod communities. METHODS Study area partement des BouchesThe study area is located in the de ne (Provence, south-eastern France) (Table 1; Fig. 1). du-Rho All the sites are characterized by a true Mediterranean climate (three dry summer months and two cold winter months), annual temperatures averaging between 14 and 11 C and annual rain averaging between 543 and 680 mm year)1 (C.N.R.S., 1975). Although the study area is highly limited in order to obtain a homogeneous sampling area, slight climatic
Table 1 Sampling sites. Each sampling site is named after date of the various res suffered over the sampling period (19732001), the district and the massif where it is located, CC1971 being the reference site.
District Charleval Lambesc Rognes Aurons Lanc on-de-Provence Marseille Allauch Allauch Allauch Peynier Trets Cuges-les-Pins Fire dates 1971 1995 197989 1991 198995 1997 197997 19737997 19798997 1998 1989 1993 Name of massif ne des Co tes Cha ne des Co tes Cha ne des Co tes Cha ne des Co tes Cha ne des Co tes Cha ne de lEtoile Cha ne de lEtoile Cha ne de lEtoile Cha ne de lEtoile Cha Regagnas Regagnas Ste Baume Code CC1971 CC1995 CC197989 CC1991 CC198995 CE1997 CE197997 CE19737997 CE19798997 MR1998 MR1989 SB1993

Figure 1 Location map of study area and of sampling sites. The 12 sampling sites are called after their location and the re dates. ne des Co ne de lEtoile; tes; CE, Cha CC, Cha MR, Regagnas; SB, Ste Baume.

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Response patterns of land snail communities to re disturbances gradients exist and induce different bioclimates (Emberger, 1971) and a slight vegetation gradient from north to south (Dupias & Rey, 1985). All substrates are calcareous and sites range from 140 to 660 m in altitude. Sampling strategy Stratied sampling was carried out according to re regime over the past 30 years, i.e. number of res, re intervals and , age of the last re (Sousa, 1984; Whelan, 1995; Lloret & Mar 2001). Twelve sites of various re ages were selected, including six sites which have burned twice or three times and ve sites which have burned once over the period studied (i.e. 1973 2001) and one last burned in 1971 to act as a reference site (Fig. 2). The sampling sites were sampled during Spring 2000 and 2001 with a total of 120 sampling points. For each site, 10 sampling points were performed in various post-re vegetation stands: grassland (Brachypodium retusum Beauvais), garrigue (Quercus coccifera Linnaeus, Q. ilex Linnaeus, Ulex parviorus Pourret, Rosmarinus ofcinalis Linnaeus and Cistus albidus Linnaeus), oak stand (Q. pubescens Willdenow) and pine stand (Pinus halepensis Miller) and, for the sites which have burned more than once, at the intersection of the burned areas. These sampling points were performed in order to ensure a proper representation of land snail communities from various postre vegetation stands within each sampling site. Each sampling point consisted of square of 25 m2, and data were recorded on oristic variables, environmental variables (Godron et al., 1968) and malacofauna. Site history In order to clarify and to understand the role of the recent site history on the response patterns of communities, a brief history of vegetation structure and change in sampling sites during the last centuries was carried out. For each site studied, the recent site history was compiled using aerial photographs ographique National, 2001) taken between 1950 (Institut Ge and 1998, before each re (Fig. 3) and using old maps dating
Figure 2 Selected sampling sites, which are dened by the number of res suffered over the period studied (i.e. 19732001) and re interval type. In bold, selected re dates. The samplings were performed during Spring 2000 and 2001.

ographfrom the 18th century called Cassini maps (Institut Ge ique National, 1999) (Fig. 4). These maps provide a reliable insight into the vegetation structure and the human uses of droit, 1976). landscape at the end of the 18th century (Dougue

Figure 3 Aerial photographs taken at various dates since 1950 of the sampling site which burned once in 1998. The area sampled is located in the white square.
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147

L. Kiss et al.

Figure 4 Cassini map of the sampling site which burned in 1998. The area sampled is located in the black square.

Sampling of land snails Two different samples of land snails were taken in each sampling point. All land snails over 5 mm in diameter were collected during a standard interval of 30 min within the square of 25 m2. Smaller species were collected in four squares of 25 25 cm including litter and the ve upper centimetres of soil. Soil sample treatment (sieved on a 0.5-mm mesh), snail count and snail identication were performed in the laboratory. Only fresh shells (i.e. with intact periostracum) and living individuals were taken into account because they are representative of current communities. The list of species sampled follows the nomenclature of Kerney et al. (1999) (Appendix 1). All species collected were classied into ve biogeographical groups (Appendix 1). These groups take into account the biogeographical range of species sampled according to Magnin (1991) and Kerney et al. (1999). The rst group is composed of Mediterranean species and the second of species widely distributed in the Mediterranean and in Western Europe. The third group is composed of European species. The fourth group is composed of Holarctic species and the last group of Palearctic species. Data analyses Post-re patterns of responses of land snail communities were analysed, using STATLAB software Ivry-sur-Seine, France (SLP 148

Statistique Jambu, 1994). Site data used in the analyses were compiled using the 10 sampling points from each site. A land snail data matrix, composed of fresh shells and living individuals collected within the 12 sites, was subjected to a Correspondence Analysis (CA) in order to estimate the impact of re regime on land snail communities. We also compared community composition within the area studied taking into account the roles of re regime, of recent site history and of various bioclimates. Thus, similarities between species composition of communities at each sampling site were estimated using Srensen index (Legendre & Legendre, 1998): Sx1 ; x2 2a=2a b c; where x1 and x2 are the two sites to be compared, a number of species shared by the two sites, b and c number of species unique to each of the sites. As a variant of the best-known similarity coefcient, i.e. Jaccards coefcient, this coefcient of community gives double weight to double presence of species (Jongman et al., 1995; Legendre & Legendre, 1998), which is a strong indication of resemblance (Legendre & Legendre, 1998). A Srensen matrix was compiled using Srensen indices calculated between sampling sites taken two at a time to estimate similarities in species composition among all sites. A simulation of the distribution of the observed indices, under equirepartition hypothesis of species between the rst and the
Journal of Biogeography 31, 145157, 2004 Blackwell Publishing Ltd

Response patterns of land snail communities to re disturbances second site and in both sites, allowed us to test the signicance of observed values (randomization test, n 1000 permutations) (Manly, 1991). The signicance thresholds were *P 0.05, **P 0.01 and ***P 0.001. RESULTS Impact of recent re regimes on land snail communities Correspondence Analysis was performed on the land snail data matrix (46 species, 12 sampling sites) to study the structure of communities that have been exposed to various re regimes (Fig. 5). Axis 1 (26.88% of total inertia) puts fallow land species (Xeropicta derbentina), grassland or garrigue species (Candidula gigaxii, Candidula unifasciata) and more mesophilous species (Monacha cantiana) on the positive end, and pine stand species (Vallonia costata, Microxeromagna armillata and Lauria cylindracea) on the negative end (Table 2; Fig. 5a). Except for species characteristic of garrigue that are clustered with sampling sites which have been exposed to two successive res within a 6-year interval (CC198995 and CE19737997) (Fig. 5a,b), the species distribution is somewhat chaotic because open habitat species and shade-loving species are randomly represented on both ends of this axis. However, the sampling sites are ordered by their location from north (CC1995 and CC197989), on the positive end, to south (CE19798997) on the negative end (Table 3; Fig. 5b). Axis 2 (19.18% of total inertia) isolates synanthropic and Mediterranean species on the negative end (Fig. 5a). Species of open and dry habitats (Trochoidea trochoides, T. pyramidata and Cernuella virgata) and pine stand or garrigue species (M. armillata and C. unifasciata) are clustered on the negative end (Table 2). Litter-dwelling species and more closed garrigue species (V. costata and Granopupa granum) are located on the positive end (Table 2). This axis discriminates between the sampling sites (i.e. CE19798997 on the positive end, and SB1993 and CE1997 on the negative end), located in the southeastern part of the study area, according to a northsouth gradient (Table 3; Fig. 5b). This CA demonstrates neither a re age gradient nor a re regime gradient. The sampling sites are ordered according to a site location gradient from north-west to south-east and the land snail communities seem to be dependent on a gradient from north to south, representing the distance inland from the coast. A Srensen matrix was also compiled to compare composition of gastropod communities within the study area. Globally, Srensen indices calculated (Table 4) separate the sampling sites into two groups according to a north-west/ south-east gradient. Moreover, all the sampling sites at the same location, taken two at a time, have high Srensen indices with high signicance (P 0.001). However, when sampling sites of these two groups are compared, Srensen indices are
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Figure 5 CA of the land snail data matrix (46 species, 12 sampling sites). (a) Ordination diagram of land snail species; (b) Ordination diagram of sampling sites. AAC, Acanthinula aculeata; APO, Abida polyodon; CAC, Cecilioides acicula; CAS, Cryptomphalus aspersus; CAV, Chondrina avenacea; CGI, Candidula gigaxii; CNE, Cepaea nemoralis; CPA, Cochlostoma patulum; CRU, Clausilia rugosa; CUN, Candidula unifasciata; CVI, Cernuella virgata; EFU, Euconulus fulvus; EVE, Eobania vermiculata; GGR, Granopupa granum; GVA, Granaria variabilis HBO, Hypnophila boissyi; HLA, Helicigona lapicida; JQA, Jaminia quadridens LCY, Lauria cylindracea; MAR, Microxeromagna armillata; MAR/XCO, Microxeromagna armillata/Xerotricha conspurcata; MCA, Monacha cantiana; MCR, Monacha cartusiana; MOB, Merdigera obscura; OAL, Oxychilus alliarius; ODR, Oxychilus draparnaudi; OHY, Oxychilus hydatinus; PEL, Pomatias elegans; PMA, Phenacolimax major; PPY, Punctum pygmaeum; PSO, Papillifera solida; PSP, Pseudotachea splendida; SCA, Sphincterochila candidissima; SSI, Solatopupa similis; TCA, Truncatellina callicratis; TEL, Trochoidea elegans; TPY, Trochoidea pyramidata; TTR, Trochoidea trochoides; VCO, Vitrea contracta; VCY, Vitrea crystallina; VNA, Vitrea narbonensis; VCS, Vallonia costata; ZAL, Zonites algirus; XCE, Xerosecta cespitum; XCO, Xerotricha conspurcata; XDE, Xeropicta derbentina. See Table 1 for sampling site codes.

low, i.e. S 0.62 0.06 on average, with low or no signicance (i.e. P 0.05 and P 0.01). Furthermore, two northern sampling sites are special cases: one of them burned in 1989 and in 1995 (CC198995) and the other burned in 1991 (CC1991). The sampling site, which has burned twice with a short re interval (CC198995), has low indices with all the northern sampling sites but high index with a southern site, which has burned three times, also with a short re interval 149

L. Kiss et al.
Table 2 Signicant species variables on axes 1 and 2 of the CA (46 species, 12 sampling sites). Ctr., contribution.

Positive end Axis 1 Candidula gigaxii (Ctr. 0.06) Candidula unifasciata (Ctr. 0.03) Monacha cantiana (Ctr. 0.03) Xeropicta derbentina (Ctr. 0.28) Granopupa granum (Ctr. 0.03) Vallonia costata (Ctr. 0.26)

Negative end Lauria cylindracea (Ctr. 0.08) Microxeromagna armillata (Ctr. 0.13) Vallonia costata (Ctr. 0.17) Candidula unifasciata (Ctr. 0.05) Cernuella virgata (Ctr. 0.03) Microxeromagna armillata (Ctr. 0.38) Trochoidea pyramidata (Ctr. 0.06) Trochoidea trochoides (Ctr. 0.27)

Axis 2

Table 3 Signicant sampling site variables on axes 1 and 2 of the CA (46 species, 12 sampling sites). Ctr., contribution. See Table 1 for sampling site codes.
Positive end Axis 1 Axis 2 CC1995 (Ctr. 0.26) CC197989 (Ctr. 0.16) CE19798997 (Ctr. 0.10) Negative end CE19798997 (Ctr. 0.24) SB1993 (Ctr. 0.28) CE1997 (Ctr. 0.33)

Biogeographical composition of malacofaunas under different re regimes Proportions of species number from different biogeographical ranges were calculated for all the sampling sites (Fig. 6). European species are more numerous in the two most northern sampling sites (CC1971 and CC1995), i.e. 29.5 3.2% on average, than in all other sites, i.e. 21.2 2.9%. Moreover, the sampling sites at the same location have similar biogeographical compositions. For example both sites of the Regagnas massif, respectively burned in 1989 and 1998 (MR1989 and MR1998), have a comparable proportion of the ve groups. Mediterranean species and Mediterranean/ West European species are highly predominant in the high re regime sampling site (i.e. CC198995). In fact, this site is composed of 50% Mediterranean species and of 30% Mediterranean/West European species against, respectively, 44.4 4.5% and 23.2 4.3% among all other sites. Moreover, widely distributed species are not represented in this sampling site. With the exception of the high re regime sampling site which contains a higher proportion of Mediterranean species than the other sites, the various re regimes occurring since the beginning of the 1970s do not seem to have affected biogeographical composition of malacofaunas. Thus, the

(CE19737997). Thus, its communities are closer to the malacofaunas of a southern sampling site, which has been exposed to high re regime than those of northern sites. The sampling site, which has burned once (CC1991), has high Srensen indices with all sampling sites whatever their location and, thus, has species composition comparable with all the other sites. As in the CA, these results seem to demonstrate that land snail communities are organized according to a biogeographical gradient. Nevertheless, a high re regime (i.e. short re interval and high number of res) seems to modify malacofauna composition in a comparable direction, whatever their geographical location.

Table 4 Matrix of Srensen indices calculated between sites taken two at a time. The distribution of indices observed was tested under null hypothesis of equirepartition of species between sites and within sites. *P 0.05, **P 0.01, ***P 0.001, respective thresholds of index are P 0.6268, P 0.6667 and P 0.7042. See Table 1 for sampling site codes.
CC1971 CC1995 CC197989 CC1991 CC198995 MR1989 MR1998 CE1997 CE197997 CE19737997 CE19798997 SB1993 CC1971 CC1995 CC197989 CC1991 CC198995 MR1989 MR1998 CE1997 CE197997 CE19737997 CE19798997 SB1993 1 0.78*** 0.74*** 0.67* 0.56 0.62 0.75*** 0.64* 0.60 0.61 0.57 0.54

1 0.74*** 0.74*** 0.56 0.58 0.67* 0.60 0.64* 0.65* 0.61 0.58

1 0.63* 0.60 0.62 0.71*** 0.64* 0.64* 0.70** 0.52 0.62

1 0.75*** 0.72*** 0.71*** 0.79*** 0.82*** 0.74*** 0.81*** 0.71***

1 0.76*** 0.62 0.72*** 0.76*** 0.74*** 0.78*** 0.71***

1 0.85*** 0.78*** 0.85*** 0.80*** 0.83*** 0.77***

1 0.69** 0.76*** 0.71*** 0.71*** 0.68**

1 0.84*** 0.88*** 0.79*** 0.77***

1 0.87*** 0.93*** 0.80***

1 0.82*** 0.79***

1 0.71***

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Response patterns of land snail communities to re disturbances

Figure 6 Percentages of the number of species of the ve different biogeographical groups within each sampling site. Sampling sites are ordered on axis according to their location within the sampling area from north to south. See Table 1 for sampling site codes. Table 5 Landscape and vegetation cover changes analysed using aerial photographs taken at various dates since the 1950s. (a) History of the reference sampling site which burned once in 1971, and (b) of sampling sites which burned once over 19732001 period. See Table 1 for sampling site codes.
(a) CC1971 1950s Homogeneous landscape of open garrigue, open wood and grassland 196070 Homogeneous landscape of high and closed wood

(b) MR1989

1950s Heterogeneous landscape of garrigue, open and closed wood Heterogeneous landscape of open wood, closed garrigue with scattered trees and grassland Heterogeneous landscape of open wood, closed garrigue and garrigue with scattered trees Homogeneous and open landscapes of garrigue and grassland Heterogeneous landscape of garrigue, closed and open wood Heterogeneous landscape of garrigue and open wood

1970s Heterogeneous landscape of open and closed wood Heterogeneous landscape of open wood, closed garrigue with scattered trees Heterogeneous landscape of open wood, garrigue and garrigue with scattered trees Heterogeneous landscape of garrigue and closed wood Heterogeneous landscape of open and closed wood, and closed garrigue with scattered trees Heterogeneous landscape of open and closed wood

Pre-re (around 1 year before re) Homogeneous landscape of closed wood Heterogeneous landscape of open wood, closed garrigue with scattered trees Heterogeneous landscape of closed and open wood, garrigue and garrigue with scattered trees Heterogeneous landscape of garrigue, closed and open wood Homogeneous landscape of closed wood and closed garrigue with scattered trees Homogeneous landscape of closed wood

CC1991

SB1993

CC1995

CE1997

MR1998

land snail communities seem to depend on a northsouth biogeographical gradient, i.e. a Mediterranean one. Site history The recent history of vegetation changes (Tables 5 & 6), reconstructed using aerial photographs taken over the past 50 years, demonstrates that landscapes of sampling sites have evolved according to four different patterns.
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Certain sampling sites, that were composed of closed and wooded landscapes in the beginning of the 1950s, have retained a garrigue landscape since the rst re during the period studied (i.e. CE19737997 and CE19798997) (Table 6). Other sampling sites composed of fragmented and open landscapes in 1950 have evolved to more wooded, closed and stratied landscapes before their rst re (i.e. CC1991, SB1993, CC1995, CC197989 and CE197997) (Tables 5 & 6). 151

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Table 6 Landscape and vegetation cover changes analysed using aerial photographs taken at various dates since the 1950s and before each re. (a) History of the sampling sites, which have burned twice and, (b) of the sampling sites which have burned three times over 19732001 period. See Table 1 for sampling site codes.
(a) 1950s 1949 Heterogeneous landscape of open and closed garrigue, grassland and low wood 1949 Heterogeneous landscape of low and open wood, grassland with scattered trees 1950 Heterogeneous landscape of open and closed wood, and closed garrigue 196070 1964 Heterogeneous landscape of closed garrigue with scattered trees, and closed and low wood 1968 Homogeneous, low and open landscape with some areas of high garrigue 1969 Heterogeneous landscape of closed garrigue with scattered trees and open garrigue Before the rst re 1969 Homogeneous landscape of high and closed garrigue Before the rst re 1978 Heterogeneous landscape of open and closed woods Before the second re 1989 Heterogeneous landscape of closed wood and garrigue with scattered trees 1993 Homogeneous, low and open landscape with some areas of high garrigue 1997 Heterogeneous landscape of closed and high garrigue, and open garrigue

CC197989

CC198995

1989 Heterogeneous landscape of closed and highly open garrigue 1978 Heterogeneous landscape of open wood, closed garrigue and grassland

CE197997

(b)

1950s 1950 Homogeneous landscape of low and closed wood

Before the second re 1978 Homogeneous landscape of low and open garrigue

Before the third re 1997 Heterogeneous landscape of high and open garrigue with scattered trees Before the third re 1997 Homogeneous landscape of high and closed garrigue

CE19737997

1950s 1950 Homogeneous landscape of high and closed wood

196070 1969 Homogeneous landscape of high and open wood

Before the rst re 1978 Homogeneous landscape of high and closed wood

Before the second re 1988 Homogeneous landscape of high and closed garrigue

CE19798997

Some sampling sites which were composed of open and wooded landscapes after the Second World War, have evolved to more closed and wooded landscapes since their rst re (CC1971, MR1989 and MR1998) (Table 5). Only the sampling site, that has been exposed to high re regime (i.e. CC198995), has a different vegetation history (Table 6). Indeed, this sampling site has retained, since the 1950s, a landscape of more or less open and low garrigue. The recent history of vegetation change was also compiled using detailed geographical maps called Cassini maps, which date from the end of the 18th century. ne de lEtoile Southern sampling sites, located in the Cha massif, were composed of low landscapes of garrigues and grasslands with few scattered trees (Table 7). Most of the ne des Co tes north-western sampling sites, located in the Cha massif, were composed of forests, woods and of some cultivated areas, but the sampling site that has been exposed to high re regime (i.e. CC198995) was composed, as nowadays, of homogeneous and open landscapes of garrigues. South-eastern sampling sites, located in the Regagnas and Ste Baume massifs, were composed respectively of wooded and closed landscapes and of fragmented landscapes including woods and garrigues. 152

The different massifs have been exposed to various uses and some of them have been more extensively cultivated. This was particularly the case of the high re regime site (i.e. CC1989 95) which appears to have been composed of garrigue since the 18th century. DISCUSSION Impact of recent re regimes on land snail communities As demonstrated before (Kiss & Magnin, 2002, 2003; Nekola, 2002), response patterns of land snail communities to re regimes are not clear. Indeed, communities are not organized according to a re number gradient, a re age gradient or a re interval gradient in the analyses. Only the high re regime sampling sites (CC198995 and CE19737997) seem to have common features in their land snail compositions (Srensen matrix and CA) and they evolve to more Mediterranean malacofaunas (xerophilous and open habitat species) than the other sites. Although the short-term impact of re has a drastic effect on land snail communities (Kiss & Magnin, 2002, 2003), the
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Response patterns of land snail communities to re disturbances

Table 7 Vegetation cover and vegetation-and-use-types of the 12 sampling sites at the end of the 18th century, analysed using Cassini maps. Sampling sites are ordered according to the last re date and to their location within sampling area, i.e. (a) north-western sites located in the ne des Co ne de lEtoile massif (CE), and (c) south-eastern sites located in the tes massif (CC), (b) southern sites located in the Cha Cha Regagnas massif (MR) and in the Ste Baume massif (SB). The nomenclature is according to legends of Cassini maps. See Table 1 for sampling site codes.
(a) Vegetation cover Types of vegetation or uses (b) Vegetation cover Types of vegetation or uses (c) Vegetation cover Types of vegetation or uses CC198995 Open Heath CE1997 Open vineyard and scattered trees MR1998 Closed Forest CC1995 Closed Forest CC1991 Open and closed Cultivated area and forest CE197997 Open Heath SB1993 Open Wood and heath CC197989 Open and closed Cultivated area and forest CE19798997 Open Heath CC1971 Closed Forest CE19737997 Open Heath MR1989 Closed Forest

analyses performed (CA and Srensen matrix) show that these communities are particularly resilient to low re regimes over the period 19732001, as the biogeographical composition of malacofaunas does not change. In fact, some sampling sites which have suffered one re over the sampling period (i.e. MR1989, MR1998, CE1997, CC1993 and CC1971) and the sampling site which has burned twice with a long re interval (i.e. CE197997) are composed of the ve biogeographical groups, whatever the age of the last re. Thus, other factors, i.e. biogeographical or historical, seem to affect the response patterns of gastropod communities. Persistent biogeographical patterns As expected, re regime does not appear to be the main factor controlling malacofauna composition. In general, sampling sites are separated according to a south-east/north-west gradient in the analyses performed (CA and Srensen matrix). Although the most northern sampling sites have been exposed to various re regimes and have different last re ages, they have comparable species composition (high Srensen indices) and similar percentages of the different biogeographical groups. Moreover, their communities are composed of a higher percentage of European range species than the other malacofaunas. Moreover, although some north and south sampling sites have shown similar vegetation cover history and vegetation structure changes (i.e. CC1971, MR1998 and MR1989) since the 18th century, and before the rst re suffered over the period studied, they are separated by the different analyses (CA and Srensen matrix). It is clear that whatever the re regime, the vegetation cover history and the human uses, land snail communities are comparable with the northern sampling sites. Therefore, the communities seem to be controlled by a biogeographical gradient from north to south representing
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the distance inland. Indeed, the sampling area is characterized by various precipitation regimes from north to south and by cooler climate in its northern part (Emberger, 1971; C.N.R.S., 1975) and it has different bioclimates (Emberger, 1971), which induce a oristic composition gradient at micro-scale (Molinier, 1974; Dupias & Rey, 1985). Thus, gastropod communities are organized according to a gradient which can be due to combined or distinct inuences of a climate gradient and of a oristic composition gradient. On the one hand, land snails are highly sensitive to the structure and the micro-climate of their habitat (Boycott, 1934; Cameron & Redfern, 1976; Bishop, 1977). On the other hand, in Mediterranean ecosystems of Crete, malacofauna diversity is dependent on precipitation regime, which induces a biogeographical gradient at the microregional scale (Cameron et al., 2000). Moreover, malacofaunas vary widely between northern and southern Provence (Magnin, 1991; Pfenninger et al., 2003), although at a higher scale, i.e. 100 km against 30 km in our sampling area. In consequence, a Mediterranean gradient seems to exist within our restricted sampling area and to inuence at one and the same time both ora and malacofaunas. Another hypothesis based on historical biogeography could also explain, to some extent, this gradient. Generally, Mediterranean land snail communities are greatly diversied even within a province (Cameron et al., 2000; Cameron et al., 2003). Although communities have been sampled within a restricted area in our study, it was composed of four more or ne des Co ne de tes, Cha less connected massifs (i.e. Cha lEtoile, Regagnas and Ste Baume) where relict and persistent communities could have been maintained during historical times (Cameron et al., 1980; Blondel, 1995). Role of landscape history Aerial photographs, taken at different dates since the 1950s, and Cassini maps provide valuable historic insight into landscape 153

L. Kiss et al. structure and vegetation cover. The sampling site history was compiled in order to clarify and to understand variability in response patterns of gastropod communities, which have been exposed to various re regimes. However, chronological breaks should be taken into account in the reconstructed landscape changes, which is based on these point data and not on continuous information. This reconstruction has, nevertheless, provided an overall picture of vegetation cover changes and of the pre-re habitat structure (forest, garrigue and grassland). Although the sampling site (i.e. CC198995), which has burned twice (in 1989 and 1995), is geographically close to the other northern sampling sites, it is always isolated from them in all analyses performed (CA, biogeographical diagram and Srensen matrix). This site is mainly composed of Mediterranean and Mediterranean/West European species and its malacofauna does not seem to be controlled by biogeographical inuence like communities of the most northern sites. Indeed, its landscapes are denitely composed of garrigue since 1950 (aerial photographs) and probably even before the 18th century, because the sampling site landscape is described in open and low habitats of garrigue by Cassini maps. Moreover, the area where this sampling site is located has been exposed to old anthropogenic disturbances. In previous eras, historical landscapes were maintained in garrigue by frequent pastoral res (Guillerm & Trabaud, 1980) from the 15th century to modern time (Amouric, 1993) to some extent in order to favour and to collect a parasite insect (Kermococcus vermilio Planchon) of Q. coccifera which provided red dye for wool (Delmas, 1958). In consequence, the composition of current gastropod communities (xerophilous and Mediterranean species) is the result of various ancient and high regimes of anthropogenic disturbances, which seem to erase biogeographical roles. Moreover, the persistence of Q. coccifera landscapes over centuries, due to these various disturbances, probably induces a consistent land snail community composition in this sampling site. While long-term historical explanations of differences in fauna are generally acceptable on a biogeographical scale (Cameron et al., 1980), historical processes on a local scale, as it is demonstrated in this study, also seem to inuence current land snail communities. This history consists of re regimes, sometime human-induced, and of anthropogenic disturbances, which control vegetation cover and landscape changes. Thus, response patterns of gastropod communities are controlled by the more or less recent history of sampling sites (Cameron et al., 2000). Combined inuences of landscape history and of biogeography Both the composition of the malacofauna and the response patterns of gastropod communities to res seem to be shaped by the combined inuences of landscape history and biogeographical gradient. In fact, numerous factors seem to control response patterns of post-re gastropod communities in several sampling sites. 154 In the Regagnas massif both sampling sites located had comparable land snail communities including shade-loving species, although one of them was the site most recently burned. The communities from these sites have comparable structure (CA) and composition (high Srensen indices) and similar percentages of the ve biogeographical groups. Moreover, these sites show identical vegetation cover history before the rst re during the study period, and landscape and vegetation cover changes have been comparable since the 18th century. We believe that these sites have been maintained in defens over several centuries, i.e. sheltered from wood cutting and pastoral activities (Amouric, 1992). More recently, despite a high re regime suffered in the Regagnas massif since 1973 (12 res of more than 10 ha) (Centre informatique de la fecture des Bouches-du-Rho ne, 2002), these sites seem to Pre have been spared and forest cover has partially been maintained over the last decades. Thus, recent history and geographical position seem to inuence the community composition of these sites. ne de lEtoile massif have All sampling sites of the Cha malacofaunas comparable with those of the southern sites (high Srensen indices). They are also exposed to the Mediterranean gradient. Moreover, although these sampling sites have similar species compositions (high Srensen indices), they are not grouped in the CA. Thus, although re regime affects community composition, it seems that it is not the main factor. Indeed, these sampling sites have not only been exposed to various re regimes but they have also shown different vegetation cover and landscape histories over decades. These vegetation changes probably explain the differences observed in malacofauna composition. The community composition of this massif is thus the result of the recent vegetation history, of re regimes over the last three decades and of biogeography. However, biogeographical and historical inuences can be observed in community response patterns because of malacofauna persistence which must, in turn, be due to cryptic refuges within burned areas. Numerous species are able to survive even within a priori unavailable habitats by using micro-refuges, such as trunks, stumps or logs within dry and open forest after clear cutting (Shikov, 1984). Variability in re severity and land snail location at the time of re probably induce numerous cryptic refuges (Kiss & Magnin, 2003). The presence of these refuges permit land snails, rst, to survive during re event and, secondly, to persist following (and between) successive res. Indeed, the persistence of shadeloving and mesophilous species, whatever re age in the Regagnas massif, indicates land snail survival within cryptic refuges located in burned areas (Kiss & Magnin, 2002, 2003). ne de lEtoile sites (CE197997 For example, two of the Cha and CE19798997), which are < 1 km away from each other, have the highest Srensen index (S 0.93), although sampling sites have been exposed to various re regimes and landscape histories. Finally in some cases, the response patterns of communities to various re regimes also seem to be inuenced by the
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Response patterns of land snail communities to re disturbances structure of pre-re habitats and by post-re human uses (Kiss & Magnin, 2003). The land snail communities from one of the northern sites that burned once in 1991 (i.e. CC1991), have particular species compositions because they are similar not only to the northern sampling sites but also to most of the southern ones (i.e. signicant Srensen indices with all sites). There are two possible explanations for this result: rst, although all the southern sampling sites, which have high Srensen index with this northern site, have different vegetation cover histories, they all show identical pre-re vegetation structure, i.e. garrigue. Thus, the same pre-re landscape structure seems to inuence species composition but also indicates a certain persistence of malacofaunas after disturbance. Secondly, although this site has less signicant indices with the sampling sites located in the northern part of the area studied, it is clustered with them in the CA. Its geographical location and the Mediterranean gradient may explain partially these observations. However, some of these sites (CC1971, CC1995) are exposed to passage of animal herds, that constitutes dispersal vector for the invasive species Xeropicta derbentina (Labaune & Magnin, 1999). Thus, post-re disturbances also affect community composition and response patterns. CONCLUSIONS Gastropod communities appear to be highly resilient to re (Frest & Johannes, 1995; Theler, 1997; Kiss & Magnin, 2003), as to various other anthropogenic disturbances (Shikov, 1984; Cameron & Greenwood, 1991), provided that disturbance regime is not maintained over years or that time elapsed between two successive disturbances is long (Nekola, 2002). However, no clear trends in response patterns of gastropod communities after re have been pointed out. Current postre gastropod communities seem to be determined by numerous factors and are simultaneously subjected to biogeographical inuences (bioclimatic gradient and/or historical biogeography) and to recent history, including re regime, past and present anthropogenic disturbances and landscape changes. Nevertheless, the response patterns obtained must be due to cryptic refuges located within burned areas that allow malacofaunas to remain for years after successive res. Moreover, current communities are the product of the past, particularly of Neolithic times, and of current human activities, involving Mediterranean species introduction and landscape modications in the Mediterranean basin (Mylonas, 1984; Magnin, 1992; Welter-Schultes & Williams, 1999; Martin et al., 2003). In conclusion, these features should not be neglected in the interpretation of ecological responses of less mobile faunas like gastropod communities subsequent to disturbances. ACKNOWLEDGMENTS This study was carried out in the framework under the GIS cologique et paysager des incendies sur les garrigues Impacts e
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ts pe ri-marseillaises (IMEP-CNRS/CEMAGREF) and et les fore ne ral des was supported by funds provided by the Conseil Ge ne and by Unitas Malacologica. We would Bouches-du-Rho like to thank Isabelle Girard, Cinderella Grout, Virginie Libois and Sylvie Marguerier for assistance in the eldwork. We are indebted to Marjorie Sweetko for her careful re-reading and her help in translation, and to Markus Pfenninger for his judicious and helpful comments on the manuscript. REFERENCES ` le preuve du temps. Narration, Amouric, H. (1992) Le feu a Aix-en-Provence. Amouric, H. (1993) Calissanne et Merveille; deux domaines conomie de lEtang de Berre (de la n du Moyen Age dans le ` cle). Arche ologie et Environnement: de la Ste au XVIIIe sie Victoire aux Alpilles (ed. by P. Leveau and M. Provansal), pp. de Provence, Aix-en315373. Publications Universite Provence. Barbero, M., Bonin, G., Loisel, R., Miglioretti, F. & Quezel, P. (1987) Impact of forest res on structure and architecture of Mediterranean ecosystems. Ecologia Mediterranea, 13, 3950. Bishop, J. (1977) Approaches to the quantitative description of terrestrial mollusc populations and habitats. Malacologia, 16, 6166. ographie, approche e cologique et e voluBlondel, J. (1995) Bioge tive. Masson, Paris. Boycott, A.E. (1934) The habitats of land mollusca in Britain. Journal of Animal Ecology, 22, 138. taille e de la France, MarC.N.R.S. (1975) Carte Climatique de seille. Centre National De La Recherche Scientique. Edn. Ophrys, Gap. Cameron, R.A.D. & Greenwood, J.J.D. (1991) Some montane and forest molluscan faunas from eastern Scotland: effects of altitude, disturbance and isolation. Proceeding of the Tenth International Malacological Congress, Tubingen, 1989, 437442. Cameron, R.A.D. & Redfern, M. (1976) British land snails. Mollusca: Gastropoda. Academic Press, London. Cameron, R.A.D., Down, K. & Pannett, D.J. (1980) Historical and environmental inuences on hedgerow snail faunas. Biological Journal of the Linnean Society, 13, 7587. Cameron, R.A.D., Mylonas, M. & Vardinoyannis, K. (2000) Local and regional diversity in some Aegean land snail faunas. Journal of Molluscan Studies, 66, 131142. Cameron, R.A.D., Mylonas, M., Triantis, K., Parmakelis, A. & Vardinoyannis, K. (2003) Land-snail diversity in a square kilometre of Cretan maquis: modest species richness, hight density and local homogeneity. Journal of Molluscan Studies, 69, 8793. Carcaillet, C. (1998) A spatially precise study of Holocene re history, climate and human impact within the Maurienne valley, North French Alps. Journal of Ecology, 86, 384396. fecture des Bouches-du-Rho ne Centre informatique de la Pre the e: La banque de donne es sur les incendies de (2002) Prome ts en Re gion me diterrane enne en France. 1997. http:// Fore www.promethee.com, Marsaille. 155

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Response patterns of land snail communities to re disturbances Theler, J.L. (1997) The modern terrestrial gastropod (land snail) fauna of western Wisconsins hill prairies. The Nautilus, 110, 111121. et combustibilite des prinTrabaud, L. (1976) Inammabilite ` ces des garrigues de la re gion me diterrane enne. cipales espe Acta Oecologica-Oecologica Plantarum, 11, 117136. Trabaud, L. (1987) Dynamics after re of scerophyllous plant communities in the Mediterranean basin. Ecologia Mediterranea, 13, 2537. ` , M., Lloret, F., Ogheri, E. & Terradas, J. (2001) Positive Vila re-grass feedback in Mediterranean Basin Woodlands. Forest Ecology and Management, 147, 314. Welter-Schultes, F.W. & Williams, M.R. (1999) History, island area and habitat availability determine land snail species richness of Aegean islands. Journal of Biogeography, 26, 239 249. Whelan, R.J. (1995) The ecology of re. Cambridge University Press, Cambridge.
Appendix 1 List of species sampled within the 12 sampling sites and classied into ve biogeographical groups. E, European range; H, Holarctic range; M, Mediterranean range; M/WE, Mediterranean and West-European range; P, Palearctic range.
Biogeographical range Abida polyodon (Draparnaud 1801) ller 1774) Acanthinula aculeata (O. F. Mu Candidula gigaxii (L. Pfeiffer 1850) Candidula unifasciata (Poiret 1801) ller 1774) Cecilioides acicula (O. F. Mu Cepaea nemoralis (Linnaeus 1758) Cernuella virgata (da Costa 1778) ` re 1972) Chondrina avenacea (Bruguie russac 1807) Clausilia rugosa (A. Fe Cochlostoma patulum (Draparnaud 1801) ller 1774) Cryptomphalus aspersus (O. F. Mu ller 1774) Eobania vermiculata (O. F. Mu ller 1774) Euconulus fulvus (O. F. Mu Helicigona lapicida (Linnaeus 1758) Hypnophila boissiy (Dupuy 1850) Granaria variabilis (Draparnaud 1801) Granopupa granum (Draparnaud 1801) ller 1774) Jaminia quadridens (O. F. Mu Lauria cylindracea (da Costa 1778) ller 1774) Merdigera obscura (O. F. Mu Microxeromagna armillata (Lowe 1852) Monacha cantiana (Montagu 1803) ller 1774) Monacha cartusiana (O. F. Mu Oxychilus alliarius (Miller 1822) ssler 1838) Oxychilus hydatinus (Rossma Oxychilus draparnaudi (Beck 1837) Papillifera solida (Draparnaud 1805) russac 1807) Phenacolimax major (A. Fe ller 1774) Pomatias elegans (O. F. Mu Pseudotachea splendida (Draparnaud 1801) Punctum pygmaeum (Draparnaud 1801) ` re 1792) Solatopupa similis (Bruguie Sphincterochila candidissima (Draparnaud 1801) Trochoidea elegans (Gmelin 1791) Trochoidea pyramidata (Draparnaud 1805) Trochoidea trochoides (Poiret 1789) Truncatellina callicratis (Scacchi 1833) ller 1774) Vallonia costata (O. F. Mu Vitrea contracta (Westerlund 1871) ller 1774) Vitrea crystallina (O. F. Mu Vitrea narbonensis (Clessin 1877) Xeropicta derbentina (Krynicki 1836) Xerosecta cespitum (Draparnaud 1801) Xerotricha conspurcata (Draparnaud 1801) Zonites algirus (Linnaeus 1758) M P E E M/WE E M/WE E E M M/WE M H E M M M M/WE M/WE E M M/WE M/WE E M M/WE M E M/WE M H M M M M M M H E E E M M M M

BIOSKETCHES Dr Laurence Kiss is a researcher in landscape and commu des Sciences et Techniques de St nity ecology at the Faculte ro me (Marseille, France) with main research interest in the Je impact of re on Mediterranean land snail communities and in their patterns of post-re recolonization. de ric Magnin is a researcher in landscape ecology at Dr Fre des Sciences et Techniques de St Je ro me (Marseille, the Faculte France). His main research topics are response patterns of land snail communities to anthropogenic disturbances and climate change, paleoecology, historical biogeography and invasion biology. Dr Franck Torre is a biostatistical researcher at the Faculte ro me (Marseille, France) des Sciences et Techniques de St Je whose research topics include ecological modelling and multivariate statistical analysis.

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157