Journal of Biogeography (J. Biogeogr.

) (2004) 31, 145–157

ORIGINAL ARTICLE

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

´diterrane ´en d’Ecologie et de Institut Me ´oe ´cologie, U.M.R. 6116 du C.N.R.S., Pale ´ des Sciences et Techniques de St Faculte ´ro ˆme, France. Je

ABSTRACT

Aims To assess the impact of various fire regimes over the past 30 years on land snail communities and to analyse the role of recent landscape history and the influence of biogeography in shaping the response patterns of gastropod communities following disturbances by fire. Location South-eastern France (Provence) and Mediterranean region. Methods Stratified sampling within 12 sites was undertaken with regard to fire regime (i.e. number of fires, fire intervals and age of the last fire) 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 Sørensen coefficient of similarity. Results When a disturbance regime (land use or fire disturbances) has been maintained over decades or centuries, land snail communities appear highly modified and tend to be composed of only Mediterranean and xerophilous species. However, low fire 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 fires 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 fire 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 fire regimes but also of anthropogenic disturbances, of landscape changes over the last centuries and of subsequent structure of the pre-fire habitat, as well as of the influence of a biogeographical gradient. However, the response patterns observed and the persistence of pre-fire communities imply the presence of cryptic refuges located within burned areas. Keywords Land snail communities, fire regime, response patterns, biogeographical gradient, landscape history.

*Correspondence: Laurence Kiss, IMEP, ˆ 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 & Marı
ª 2004 Blackwell Publishing Ltd www.blackwellpublishing.com/jbi

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

. 2000).. Le Houe Although numerous studies treat the impact of fire regime on vegetation (Barbero et al. this 1995. Whelan. the district and the massif where it is located. The main objective of the present study is to understand the variability in response patterns of land snail communities to fire disturbance. land abandonment and decrease in pastoral activities have increased garrigue (calcareous matorral) and forests.N. slight climatic Table 1 Sampling sites. Ste Baume. 1987. within a priori a homogeneous area. 1998. Lyon et al. ´ rou.) is heterogeneous throughout Provence (Amouric. in addition. 1987. Trabaud. 2002. 1). 1987. 1984. includes slightly different Mediterranean bioclimates (Emberger.R. 146 Journal of Biogeography 31. type of land use.. CC1971 being the reference site. ˆ tes. Chaı MR. whatever the age of the fire (Kiss & Magnin. Lloret & Marı Since the beginning of the 20th century.S. 1971). Pons & Thinon. etc. 2003). Indeed. Dı impact on fauna within Mediterranean ecosystems has been little studied and mainly for the American and Australian continents (Whelan. The 12 sampling sites are called after their location and the fire dates.L. District Charleval Lambesc Rognes Aurons Lanc ¸ on-de-Provence Marseille Allauch Allauch Allauch Peynier Trets Cuges-les-Pins Fire dates 1971 1995 1979–89 1991 1989–95 1997 1979–97 1973–79–97 1979–89–97 1998 1989 1993 Name of massif ˆne des Co ˆ tes Chaı ˆne des Co ˆ tes Chaı ˆne des Co ˆ tes Chaı ˆne des Co ˆ tes Chaı ˆne des Co ˆ tes Chaı ˆne de l’Etoile Chaı ˆne de l’Etoile Chaı ˆne de l’Etoile Chaı ˆne de l’Etoile Chaı Regagnas Regagnas Ste Baume Code CC1971 CC1995 CC1979–89 CC1991 CC1989–95 CE1997 CE1979–97 CE1973–79–97 CE1979–89–97 MR1998 MR1989 SB1993 Figure 1 Location map of study area and of sampling sites. but also to estimate the role of recent site history and the influence of biogeographical gradient. Mylonas. ª 2004 Blackwell Publishing Ltd . 1995). fire regimes are also related to human intervention (Naveh.. 2001). Other factors are likely to explain to some extent the malacofauna composition after fire. Each sampling site is named after date of the various fires suffered over the sampling period (1973–2001). recent landscape history (date of land abandonment. CE. annual temperatures averaging between 14 and 11 °C and annual rain averaging between 543 and 680 mm year)1 (C. in shaping the response patterns of gastropod communities. Elements of pre-fire gastropod communities seem to persist within post-fire communities. 2002.. and increasing build-up of fuel has led to uncontrolled wildfires (Barbero et al. Previous studies on the impact of fire on Mediterranean land snail communities have demonstrated that various response patterns of communities have been obtained for the same fire regime (Kiss & Magnin. Throughout the Mediterranean basin. 2000. 1975). 2003). ´. south-eastern France) (Table 1. Carcaillet. du-Rho All the sites are characterized by a true Mediterranean climate (three dry summer months and two cold winter months). Vila ` et al. 2001. Huff & Smith. ´az-Delgado & Pons. 1992) which. Kiss et al. Regagnas. SB. Although the study area is highly limited in order to obtain a homogeneous sampling area. 1987). Fig. 1974) dating to Neolithic times (Guillerm & Trabaud. 2001). METHODS Study area ´ partement des BouchesThe study area is located in the ‘de ˆ ne’ (Provence. The aims of this study are thus not only to analyse the impact of various fire regimes on land snail communities over the past 30 years. ˆne des Co ˆne de l’Etoile. 1980. 1995. 1987. Chaı CC. 145–157.

Each sampling point consisted of square of 25 m2. ª 2004 Blackwell Publishing Ltd 147 . 1984. ´ ographfrom the 18th century called Cassini maps (Institut Ge ique National. The sampling sites were sampled during Spring 2000 and 2001 with a total of 120 sampling points. For each site. Site history In order to clarify and to understand the role of the recent site history on the response patterns of communities. and data were recorded on floristic variables. 4). fire intervals and ´. Rosmarinus officinalis Linnaeus and Cistus albidus Linnaeus). Twelve sites of various fire ages were selected. 10 sampling points were performed in various post-fire vegetation stands: grassland (Brachypodium retusum Beauvais). pubescens Willdenow) and pine stand (Pinus halepensis Miller) and. 1968) and malacofauna. 1985). which are defined by the number of fires suffered over the period studied (i. selected fire dates. Sampling strategy Stratified sampling was carried out according to fire regime over the past 30 years. For each site studied. Lloret & Marı 2001). These sampling points were performed in order to ensure a proper representation of land snail communities from various postfire vegetation stands within each sampling site.e. i. ilex Linnaeus. for the sites which have burned more than once.e. number of fires. Ulex parviflorus Pourret. Whelan. 145–157. the recent site history was compiled using aerial photographs ´ ographique National. The area sampled is located in the white square. 1995. a brief history of vegetation structure and change in sampling sites during the last centuries was carried out. All substrates are calcareous and sites range from 140 to 660 m in altitude.e. 3) and using old maps dating Figure 2 Selected sampling sites. 1999) (Fig. 2). 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.Response patterns of land snail communities to fire disturbances gradients exist and induce different bioclimates (Emberger.. including six sites which have burned twice or three times and five sites which have burned once over the period studied (i. 1971) and a slight vegetation gradient from north to south (Dupias & Rey. These maps provide a reliable insight into the vegetation structure and the human uses of ´ droit. Q. Journal of Biogeography 31. environmental variables (Godron et al. The samplings were performed during Spring 2000 and 2001. 1973–2001) and fire interval type. 2001) taken between 1950 (Institut Ge and 1998. oak stand (Q. In bold. before each fire (Fig. garrigue (Quercus coccifera Linnaeus. at the intersection of the burned areas. age of the last fire (Sousa. 1973– 2001) and one last burned in 1971 to act as a reference site (Fig. 1976).

ª 2004 Blackwell Publishing Ltd . A Sørensen matrix was compiled using Sørensen indices calculated between sampling sites taken two at a time to estimate similarities in species composition among all sites. 1998). Site data used in the analyses were compiled using the 10 sampling points from each site. Only fresh shells (i. Thus.L. Sampling of land snails Two different samples of land snails were taken in each sampling point. snail count and snail identification were performed in the laboratory. Smaller species were collected in four squares of 25 · 25 cm including litter and the five upper centimetres of soil. under equirepartition hypothesis of species between the first and the Journal of Biogeography 31. France (SLP 148 Statistique Jambu. 1998): Sðx1 . 1995. (1999) (Appendix 1). was subjected to a Correspondence Analysis (CA) in order to estimate the impact of fire regime on land snail communities. this coefficient of community gives double weight to double presence of species (Jongman et al. composed of fresh shells and living individuals collected within the 12 sites. The fourth group is composed of Holarctic species and the last group of Palearctic species. The area sampled is located in the black square. The third group is composed of European species. of recent site history and of various bioclimates.. 1998). 1994). x2 Þ ¼ 2a=2a þ b þ c. b and c ¼ number of species unique to each of the sites. Legendre & Legendre.5-mm mesh).e. These groups take into account the biogeographical range of species sampled according to Magnin (1991) and Kerney et al. Figure 4 Cassini map of the sampling site which burned in 1998.e. Data analyses Post-fire patterns of responses of land snail communities were analysed. where x1 and x2 are the two sites to be compared. As a variant of the best-known similarity coefficient. similarities between species composition of communities at each sampling site were estimated using Sørensen index (Legendre & Legendre. i. All land snails over 5 mm in diameter were collected during a standard interval of 30 min within the square of 25 m2. a ¼ number of species shared by the two sites. with intact periostracum) and living individuals were taken into account because they are representative of current communities. A land snail data matrix. which is a strong indication of resemblance (Legendre & Legendre. Soil sample treatment (sieved on a 0. All species collected were classified into five biogeographical groups (Appendix 1). We also compared community composition within the area studied taking into account the roles of fire regime. 145–157. The list of species sampled follows the nomenclature of Kerney et al. A simulation of the distribution of the observed indices. The first group is composed of Mediterranean species and the second of species widely distributed in the Mediterranean and in Western Europe. using STATLAB software Ivry-sur-Seine. Jaccard’s coefficient. Kiss et al. (1999).

i. MAR/XCO. AAC. Oxychilus alliarius. Merdigera obscura. APO. Abida polyodon.e. CAC. PEL. Papillifera solida. CRU. Sørensen indices are Journal of Biogeography 31.Response patterns of land snail communities to fire disturbances second site and in both sites. 5b). Candidula gigaxii. Microxeromagna armillata. has low indices with all the northern sampling sites but high index with a southern site. Microxeromagna armillata/Xerotricha conspurcata. EFU. two northern sampling sites are special cases: one of them burned in 1989 and in 1995 (CC1989–95) and the other burned in 1991 (CC1991). PPY. unifasciata) are clustered on the negative end (Table 2). 1991). Acanthinula aculeata. Solatopupa similis.e. pyramidata and Cernuella virgata) and pine stand or garrigue species (M. Vitrea narbonensis. when sampling sites of these two groups are compared. CAS. with low or no significance (i. 5a). Fig. MAR. Xeropicta derbentina. Globally. ODR. CVI. TCA. located in the southeastern part of the study area. PSP. and SB1993 and CE1997 on the negative end). Eobania vermiculata. Cochlostoma patulum. Oxychilus hydatinus. Helicigona lapicida. Cryptomphalus aspersus. P £ 0. PMA. Zonites algirus. the sampling sites are ordered by their location from north (CC1995 and CC1979–89). Xerotricha conspurcata. Xerosecta cespitum.001). the species distribution is somewhat chaotic because open habitat species and shade-loving species are randomly represented on both ends of this axis.01 and ***P £ 0. Sphincterochila candidissima. Granaria variabilis HBO.88% of total inertia) puts fallow land species (Xeropicta derbentina). representing the distance inland from the coast. Sørensen indices calculated (Table 4) separate the sampling sites into two groups according to a north-west/ south-east gradient. S ¼ 0. (a) Ordination diagram of land snail species. low. MOB. Phenacolimax major. EVE.05.001. Punctum pygmaeum. Fig. See Table 1 for sampling site codes. GVA. MCR.05 and P £ 0. CPA. have high Sørensen indices with high significance (P £ 0. Monacha cartusiana. (b) Ordination diagram of sampling sites. 5b). RESULTS Impact of recent fire regimes on land snail communities Correspondence Analysis was performed on the land snail data matrix (46 species. armillata and C. ZAL. Vitrea contracta. Candidula unifasciata. SCA. Trochoidea trochoides. Euconulus fulvus. on the positive end. HLA. VCO. Moreover. allowed us to test the significance of observed values (randomization test. OHY. OAL. Cepaea nemoralis. Granopupa granum.b). TEL. 5a). However. Vitrea crystallina. CAV. Fig. T. VCS. Furthermore.18% of total inertia) isolates synanthropic and Mediterranean species on the negative end (Fig. **P £ 0. JQA. and pine stand species (Vallonia costata. Trochoidea pyramidata.62 ± 0. ª 2004 Blackwell Publishing Ltd Figure 5 CA of the land snail data matrix (46 species. Jaminia quadridens LCY.01). all the sampling sites at the same location. A Sørensen matrix was also compiled to compare composition of gastropod communities within the study area.e. Pomatias elegans. n ¼ 1000 permutations) (Manly. Axis 1 (26. Truncatellina callicratis. Hypnophila boissyi. VCY. MCA. TTR. Cernuella virgata. CNE. Cecilioides acicula. taken two at a time. which has burned twice with a short fire interval (CC1989–95). Axis 2 (19. to south (CE1979–89–97) on the negative end (Table 3. 12 sampling sites) to study the structure of communities that have been exposed to various fire regimes (Fig. Chondrina avenacea. Except for species characteristic of garrigue that are clustered with sampling sites which have been exposed to two successive fires within a 6-year interval (CC1989–95 and CE1973–79–97) (Fig. CE1979–89–97 on the positive end. 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. XCE. XDE. This axis discriminates between the sampling sites (i. CGI. 5). grassland or garrigue species (Candidula gigaxii. according to a north–south gradient (Table 3. PSO. Litter-dwelling species and more closed garrigue species (V. Pseudotachea splendida. Microxeromagna armillata and Lauria cylindracea) on the negative end (Table 2. VNA. SSI. GGR. Candidula unifasciata) and more mesophilous species (Monacha cantiana) on the positive end.06 on average. This CA demonstrates neither a fire age gradient nor a fire regime gradient. also with a short fire interval 149 . Trochoidea elegans. Lauria cylindracea. Oxychilus draparnaudi. 5a. 12 sampling sites). However. CUN. which has burned three times. XCO. Species of open and dry habitats (Trochoidea trochoides. Clausilia rugosa. The significance thresholds were *P £ 0. costata and Granopupa granum) are located on the positive end (Table 2). Monacha cantiana. TPY. 145–157. The sampling site. Vallonia costata.

80*** 1 0.79*** 1 0.05) Cernuella virgata (Ctr.64* 0.6667 and P ¼ 0.61 0.10) Negative end CE1979–89–97 (Ctr. 29.06) Trochoidea trochoides (Ctr. **P £ 0. See Table 1 for sampling site codes. ***P £ 0.05. its communities are closer to the malacofaunas of a southern sampling site.85*** 0.70** 0. ¼ 0.88*** 0. With the exception of the high fire regime sampling site which contains a higher proportion of Mediterranean species than the other sites.26) CC1979–89 (Ctr.65* 0.72*** 0. Nevertheless. ª 2004 Blackwell Publishing Ltd . this site is composed of 50% Mediterranean species and of 30% Mediterranean/West European species against. ¼ 0.77*** 1 0. which has been exposed to high fire regime than those of northern sites.74*** 0. ¼ 0.54 1 0.28) Granopupa granum (Ctr. the (CE1973–79–97).75*** 0. ¼ 0.38) Trochoidea pyramidata (Ctr.5 ± 3. respectively. 44. CC1971 CC1995 CC1979–89 CC1991 CC1989–95 MR1989 MR1998 CE1997 CE1979–97 CE1973–79–97 CE1979–89–97 SB1993 CC1971 CC1995 CC1979–89 CC1991 CC1989–95 MR1989 MR1998 CE1997 CE1979–97 CE1973–79–97 CE1979–89–97 SB1993 1 0.80*** 0..82*** 0. Positive end Axis 1 Candidula gigaxii (Ctr.33) Biogeographical composition of malacofaunas under different fire regimes Proportions of species number from different biogeographical ranges were calculated for all the sampling sites (Fig.58 0.16) CE1979–89–97 (Ctr. 12 sampling sites).2 ± 4.001. contribution.68** 1 0.56 0.79*** 0. widely distributed species are not represented in this sampling site. ¼ 0. which has burned once (CC1991). European species are more numerous in the two most northern sampling sites (CC1971 and CC1995).52 0.3% among all other sites. P ¼ 0. ¼ 0.77*** 1 0. Thus.71*** 0.62 0. the various fire regimes occurring since the beginning of the 1970s do not seem to have affected biogeographical composition of malacofaunas.5% and 23. ¼ 0.6268.e..71*** 1 0. CC1989–95). 6). See Table 1 for sampling site codes. ¼ 0.71*** 1 0.01.81*** 0. Thus.60 0.60 0.93*** 0. i.78*** 0.62 1 0.71*** 0.13) Vallonia costata (Ctr.74*** 0. these results seem to demonstrate that land snail communities are organized according to a biogeographical gradient.62 0. ¼ 0.57 0.60 0.79*** 0. ¼ 0.63* 0. Ctr. ¼ 0.85*** 0. a high fire regime (i. The sampling site.76*** 0. ¼ 0.72*** 0. Kiss et al.78*** 0. ¼ 0.62 0. For example both sites of the Regagnas massif.2% on average.2 ± 2.78*** 0.03) Microxeromagna armillata (Ctr. Ctr.71*** 1 150 Journal of Biogeography 31.08) Microxeromagna armillata (Ctr.e. ¼ 0.4 ± 4.17) Candidula unifasciata (Ctr. short fire interval and high number of fires) seems to modify malacofauna composition in a comparable direction.03) Xeropicta derbentina (Ctr.e.24) SB1993 (Ctr.74*** 0.58 1 0. Table 2 Significant species variables on axes 1 and 2 of the CA (46 species. ¼ 0.56 0. Positive end Axis 1 Axis 2 CC1995 (Ctr. ¼ 0.74*** 0.69** 0.27) Axis 2 Table 3 Significant sampling site variables on axes 1 and 2 of the CA (46 species.64* 0.26) Negative end Lauria cylindracea (Ctr.76*** 0. ¼ 0. *P £ 0.71*** 0. respectively burned in 1989 and 1998 (MR1989 and MR1998). As in the CA. Mediterranean species and Mediterranean/ West European species are highly predominant in the high fire regime sampling site (i.87*** 0.76*** 0. Moreover. has species composition comparable with all the other sites.67* 0.83*** 0. ¼ 0.64* 0. 145–157.28) CE1997 (Ctr. the sampling sites at the same location have similar biogeographical compositions. 12 sampling sites). ¼ 0. whatever their geographical location. thus.7042.03) Monacha cantiana (Ctr. than in all other sites.06) Candidula unifasciata (Ctr.64* 0. i.67* 0. Table 4 Matrix of Sørensen 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. In fact. 21. respective thresholds of index are P ¼ 0.84*** 0.e.03) Vallonia costata (Ctr.74*** 0.71*** 0. Moreover. have a comparable proportion of the five groups.82*** 0.61 0.75*** 0. has high Sørensen indices with all sampling sites whatever their location and. ¼ 0.L. contribution.9%.

e. and closed garrigue with scattered trees Heterogeneous landscape of open and closed wood Pre-fire (around 1 year before fire) Homogeneous landscape of closed wood Heterogeneous landscape of open wood. i. that were composed of closed and wooded landscapes in the beginning of the 1950s. reconstructed using aerial photographs taken over the past 50 years. and (b) of sampling sites which burned once over 1973–2001 period. closed garrigue with scattered trees Heterogeneous landscape of closed and open wood. open wood and grassland 1960–70 Homogeneous landscape of high and closed wood (b) MR1989 1950s Heterogeneous landscape of garrigue. Table 5 Landscape and vegetation cover changes analysed using aerial photographs taken at various dates since the 1950s. ª 2004 Blackwell Publishing Ltd Certain sampling sites. 151 . CC1991. Journal of Biogeography 31. closed garrigue with scattered trees Heterogeneous landscape of open wood. CC1979–89 and CE1979–97) (Tables 5 & 6).Response patterns of land snail communities to fire disturbances Figure 6 Percentages of the number of species of the five different biogeographical groups within each sampling site. CE1973–79–97 and CE1979–89–97) (Table 6).e. CC1995.e. demonstrates that landscapes of sampling sites have evolved according to four different patterns. See Table 1 for sampling site codes. See Table 1 for sampling site codes. open and closed wood Heterogeneous landscape of open wood. closed garrigue with scattered trees and grassland Heterogeneous landscape of open wood. Other sampling sites composed of fragmented and open landscapes in 1950 have evolved to more wooded. closed and open wood Heterogeneous landscape of garrigue and open wood 1970s Heterogeneous landscape of open and closed wood Heterogeneous landscape of open wood. a Mediterranean one. have retained a garrigue landscape since the first fire during the period studied (i. Site history The recent history of vegetation changes (Tables 5 & 6). 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 north–south biogeographical gradient. Sampling sites are ordered on axis according to their location within the sampling area from north to south. closed garrigue and garrigue with scattered trees Homogeneous and open landscapes of garrigue and grassland Heterogeneous landscape of garrigue. 145–157. garrigue and garrigue with scattered trees Heterogeneous landscape of garrigue and closed wood Heterogeneous landscape of open and closed wood. garrigue and garrigue with scattered trees Heterogeneous landscape of garrigue. (a) History of the reference sampling site which burned once in 1971. closed and stratified landscapes before their first fire (i. SB1993. (a) CC1971 1950s Homogeneous landscape of open garrigue.

2003). Although the short-term impact of fire has a drastic effect on land snail communities (Kiss & Magnin. Only the sampling site. have evolved to more closed and wooded landscapes since their first fire (CC1971.e. were composed of forests. located in the Chaı massif. has a different vegetation history (Table 6). CC1989–95) was composed. See Table 1 for sampling site codes. and closed and low wood 1968 Homogeneous. but the sampling site that has been exposed to high fire regime (i. 2002. This was particularly the case of the high fire regime site (i. 2002). Most of the ˆne des Co ˆtes north-western sampling sites. low and open landscape with some areas of high garrigue 1969 Heterogeneous landscape of closed garrigue with scattered trees and open garrigue Before the first fire 1969 Homogeneous landscape of high and closed garrigue Before the first fire 1978 Heterogeneous landscape of open and closed woods Before the second fire 1989 Heterogeneous landscape of closed wood and garrigue with scattered trees 1993 Homogeneous. (a) History of the sampling sites. Table 6 Landscape and vegetation cover changes analysed using aerial photographs taken at various dates since the 1950s and before each fire. and closed garrigue 1960–70 1964 Heterogeneous landscape of closed garrigue with scattered trees. (a) 1950s 1949 Heterogeneous landscape of open and closed garrigue.e. Kiss et al. Indeed. which have burned twice and. The recent history of vegetation change was also compiled using detailed geographical maps called Cassini maps. this sampling site has retained. closed garrigue and grassland CE1979–97 (b) 1950s 1950 Homogeneous landscape of low and closed wood Before the second fire 1978 Homogeneous landscape of low and open garrigue Before the third fire 1997 Heterogeneous landscape of high and open garrigue with scattered trees Before the third fire 1997 Homogeneous landscape of high and closed garrigue CE1973–79–97 1950s 1950 Homogeneous landscape of high and closed wood 1960–70 1969 Homogeneous landscape of high and open wood Before the first fire 1978 Homogeneous landscape of high and closed wood Before the second fire 1988 Homogeneous landscape of high and closed garrigue CE1979–89–97 Some sampling sites which were composed of open and wooded landscapes after the Second World War. woods and of some cultivated areas. that has been exposed to high fire regime (i. a fire age gradient or a fire interval gradient in the analyses. were composed respectively of wooded and closed landscapes and of fragmented landscapes including woods and garrigues. 2003. of homogeneous and open landscapes of garrigues. communities are not organized according to a fire number gradient. grassland and low wood 1949 Heterogeneous landscape of low and open wood. CC1989– 95) which appears to have been composed of garrigue since the 18th century. Indeed. low and open landscape with some areas of high garrigue 1997 Heterogeneous landscape of closed and high garrigue. located in the Chaı massif.L. which date from the end of the 18th century. were composed of low landscapes of garrigues and grasslands with few scattered trees (Table 7). Nekola. Only the high fire regime sampling sites (CC1989–95 and CE1973–79–97) seem to have common features in their land snail compositions (Sørensen matrix and CA) and they evolve to more Mediterranean malacofaunas (xerophilous and open habitat species) than the other sites. South-eastern sampling sites. (b) of the sampling sites which have burned three times over 1973–2001 period. CC1989–95). response patterns of land snail communities to fire regimes are not clear. located in the Regagnas and Ste Baume massifs. DISCUSSION Impact of recent fire regimes on land snail communities As demonstrated before (Kiss & Magnin. and open garrigue CC1979–89 CC1989–95 1989 Heterogeneous landscape of closed and highly open garrigue 1978 Heterogeneous landscape of open wood. 2002. 152 The different massifs have been exposed to various uses and some of them have been more extensively cultivated. a landscape of more or less open and low garrigue. 145–157. ª 2004 Blackwell Publishing Ltd . the Journal of Biogeography 31. since the 1950s. ˆne de l’Etoile Southern sampling sites.e. MR1989 and MR1998) (Table 5). as nowadays. grassland with scattered trees 1950 Heterogeneous landscape of open and closed wood.

In consequence.R. the communities seem to be controlled by a biogeographical gradient from north to south representing Journal of Biogeography 31. they are separated by the different analyses (CA and Sørensen matrix).e. In general. Chaı l’Etoile.S.. biogeographical or historical.. Chaı less connected massifs (i. 1934. 2000).. 2003). in Mediterranean ecosystems of Crete.e. C. 1977). other factors. Moreover. as the biogeographical composition of malacofaunas does not change. MR1998 and MR1989) since the 18th century. although some north and south sampling sites have shown similar vegetation cover history and vegetation structure changes (i. On the other hand. Thus. Sampling sites are ordered according to the last fire date and to their location within sampling area. 1971). CE1997.. 2000. 1975) and it has different bioclimates (Emberger. malacofaunas vary widely between northern and southern Provence (Magnin. The nomenclature is according to legends of Cassini maps. CE1979–97) are composed of the five biogeographical groups.e. although at a higher scale.e. Role of landscape history Aerial photographs. and Cassini maps provide valuable historic insight into landscape 153 . Moreover. Indeed. ª 2004 Blackwell Publishing Ltd the distance inland. (b) southern sites located in the Chaı Chaı Regagnas massif (MR) and in the Ste Baume massif (SB). malacofauna diversity is dependent on precipitation regime. 1991. 1995). 2003). Therefore. some sampling sites which have suffered one fire over the sampling period (i. Another hypothesis based on historical biogeography could also explain. 100 km against 30 km in our sampling area. 1971. analysed using Cassini maps. 1976.N. Although the most northern sampling sites have been exposed to various fire regimes and have different last fire ages. 1980. Dupias & Rey. Pfenninger et al. In fact.e. On the one hand. It is clear that whatever the fire regime. sampling sites are separated according to a south-east/north-west gradient in the analyses performed (CA and Sørensen matrix). fire regime does not appear to be the main factor controlling malacofauna composition. i. See Table 1 for sampling site codes. Although communities have been sampled within a restricted area in our study. they have comparable species composition (high Sørensen indices) and similar percentages of the different biogeographical groups. i. Persistent biogeographical patterns As expected. MR1989. a Mediterranean gradient seems to exist within our restricted sampling area and to influence at one and the same time both flora and malacofaunas. (a) Vegetation cover Types of vegetation or uses (b) Vegetation cover Types of vegetation or uses (c) Vegetation cover Types of vegetation or uses CC1989–95 Open Heath CE1997 Open vineyard and scattered trees MR1998 Closed Forest CC1995 Closed Forest CC1991 Open and closed Cultivated area and forest CE1979–97 Open Heath SB1993 Open Wood and heath CC1979–89 Open and closed Cultivated area and forest CE1979–89–97 Open Heath CC1971 Closed Forest CE1973–79–97 Open Heath MR1989 Closed Forest analyses performed (CA and Sørensen matrix) show that these communities are particularly resilient to low fire regimes over the period 1973–2001. Blondel. whatever the age of the last fire.. CC1993 and CC1971) and the sampling site which has burned twice with a long fire interval (i. which induces a biogeographical gradient at the microregional scale (Cameron et al. (a) north-western sites located in the ˆne des Co ˆne de l’Etoile massif (CE). Bishop. gastropod communities are organized according to a gradient which can be due to combined or distinct influences of a climate gradient and of a floristic composition gradient.e. Mediterranean land snail communities are greatly diversified even within a province (Cameron et al. the vegetation cover history and the human uses. land snails are highly sensitive to the structure and the micro-climate of their habitat (Boycott.e. and before the first fire suffered over the period studied. and (c) south-eastern sites located in the ˆ tes massif (CC). seem to affect the response patterns of gastropod communities. Cameron et al. the sampling area is characterized by various precipitation regimes from north to south and by cooler climate in its northern part (Emberger. this gradient. their communities are composed of a higher percentage of European range species than the other malacofaunas. 145–157. to some extent.. Regagnas and Ste Baume) where relict and persistent communities could have been maintained during historical times (Cameron et al. CC1971. taken at different dates since the 1950s. MR1998. Thus. it was composed of four more or ˆne des Co ˆne de ˆtes. Moreover. Generally. i. Cameron & Redfern. land snail communities are comparable with the northern sampling sites. 1974.Response patterns of land snail communities to fire disturbances Table 7 Vegetation cover and vegetation-and-use-types of the 12 sampling sites at the end of the 18th century. 1985). which induce a floristic composition gradient at micro-scale (Molinier.

which seem to erase biogeographical roles. This reconstruction has. Finally in some cases. 1958). also seem to influence current land snail communities. 1980). In previous eras. 145–157.e. indicates land snail survival within cryptic refuges located in burned areas (Kiss & Magnin. 1993) to some extent in order to favour and to collect a parasite insect (Kermococcus vermilio Planchon) of Q. response patterns of gastropod communities are controlled by the more or less recent history of sampling sites (Cameron et al. probably induces a consistent land snail community composition in this sampling site. This history consists of fire regimes. provided an overall picture of vegetation cover changes and of the pre-fire habitat structure (forest. 2003). to persist following (and between) successive fires. coccifera landscapes over centuries. Kiss et al. the persistence of shadeloving and mesophilous species. structure and vegetation cover. these sites seem to Pre have been spared and forest cover has partially been maintained over the last decades. historical processes on a local scale. Moreover. historical landscapes were maintained in garrigue by frequent pastoral fires (Guillerm & Trabaud. although one of them was the site most recently burned. such as trunks. numerous factors seem to control response patterns of post-fire gastropod communities in several sampling sites. the persistence of Q. these sites show identical vegetation cover history before the first fire during the study period. although these sampling sites have similar species compositions (high Sørensen indices). 2003). In fact. which is based on these point data and not on continuous information. Variability in fire severity and land snail location at the time of fire probably induce numerous cryptic refuges (Kiss & Magnin. although fire regime affects community composition. ª 2004 Blackwell Publishing Ltd . be due to cryptic refuges within burned areas. Indeed. 1980) from the 15th century to modern time (Amouric. Moreover. The communities from these sites have comparable structure (CA) and composition (high Sørensen indices) and similar percentages of the five biogeographical groups. The presence of these refuges permit land snails. nevertheless. 1992). Moreover. to survive during fire event and. ˆne de l’Etoile massif have All sampling sites of the Chaı malacofaunas comparable with those of the southern sites (high Sørensen indices). 154 In the Regagnas massif both sampling sites located had comparable land snail communities including shade-loving species. We believe that these sites have been maintained in ‘defens’ over several centuries. Although the sampling site (i. 2002).L. Indeed.. the area where this sampling site is located has been exposed to old anthropogenic disturbances. whatever fire age in the Regagnas massif. The sampling site history was compiled in order to clarify and to understand variability in response patterns of gastropod communities. which control vegetation cover and landscape changes. recent history and geographical position seem to influence the community composition of these sites. due to these various disturbances. ˆne de l’Etoile sites (CE1979–97 For example. Moreover. which have been exposed to various fire regimes. secondly.. which has burned twice (in 1989 and 1995). Thus. because the sampling site landscape is described in open and low habitats of garrigue by Cassini maps. despite a high fire regime suffered in the Regagnas massif since 1973 (12 fires of more than 10 ha) (Centre informatique de la ´ fecture des Bouches-du-Rho ˆne. it seems that it is not the main factor. In consequence. stumps or logs within dry and open forest after clear cutting (Shikov. This site is mainly composed of Mediterranean and Mediterranean/West European species and its malacofauna does not seem to be controlled by biogeographical influence like communities of the most northern sites. Thus. Numerous species are able to survive even within a priori unavailable habitats by using micro-refuges. garrigue and grassland). More recently. sheltered from wood cutting and pastoral activities (Amouric.93). 1984). biogeographical diagram and Sørensen matrix). although sampling sites have been exposed to various fire regimes and landscape histories. chronological breaks should be taken into account in the reconstructed landscape changes. sometime human-induced. While long-term historical explanations of differences in fauna are generally acceptable on a biogeographical scale (Cameron et al. the response patterns of communities to various fire regimes also seem to be influenced by the Journal of Biogeography 31. They are also exposed to the Mediterranean gradient. two of the Chaı and CE1979–89–97). However. CC1989–95). However. the composition of current gastropod communities (xerophilous and Mediterranean species) is the result of various ancient and high regimes of anthropogenic disturbances. have the highest Sørensen index (S ¼ 0. Indeed. these sampling sites have not only been exposed to various fire regimes but they have also shown different vegetation cover and landscape histories over decades. it is always isolated from them in all analyses performed (CA. and landscape and vegetation cover changes have been comparable since the 18th century. 2000). These vegetation changes probably explain the differences observed in malacofauna composition.e. in turn. of fire regimes over the last three decades and of biogeography. first. Combined influences of landscape history and of biogeography Both the composition of the malacofauna and the response patterns of gastropod communities to fires seem to be shaped by the combined influences of landscape history and biogeographical gradient. its landscapes are definitely composed of garrigue since 1950 (aerial photographs) and probably even before the 18th century. 2002. as it is demonstrated in this study. is geographically close to the other northern sampling sites. Thus. they are not grouped in the CA. i. coccifera which provided red dye for wool (Delmas. which are < 1 km away from each other. biogeographical and historical influences can be observed in community response patterns because of malacofauna persistence which must. and of anthropogenic disturbances. The community composition of this massif is thus the result of the recent vegetation history.

D. Gap. London. However. Nevertheless. 66. (1992) Le feu a Aix-en-Provence. G. particularly of Neolithic times. 1989. R. (2000) Local and regional diversity in some Aegean land snail faunas. 1999. Barbero. (1995) Bioge tive.A. it is clustered with them in the CA. 86.D. Parmakelis. (1977) Approaches to the quantitative description of terrestrial mollusc populations and habitats.. 155 . (1991) Some montane and forest molluscan faunas from eastern Scotland: effects of altitude. CONCLUSIONS Gastropod communities appear to be highly resilient to fire (Frest & Johannes. Journal of Molluscan Studies.J. past and present anthropogenic disturbances and landscape changes. Mylonas. Welter-Schultes & Williams. (1980) Historical and environmental influences on hedgerow snail faunas. North French Alps... 1991). 437–442. CC1991). MarC. 2003).D. Journal of Animal Ecology. 1–38. climate and human impact within the Maurienne valley. Thus.S. Provansal). some of these sites (CC1971. 1992. Magnin. Cinderella Grout. ´taille ´e de la France. H. There are two possible explanations for this result: first. Proceeding of the Tenth International Malacological Congress. Down. these features should not be neglected in the interpretation of ecological responses of less mobile faunas like gastropod communities subsequent to disturbances. Martin et al.Response patterns of land snail communities to fire disturbances structure of pre-fire habitats and by post-fire human uses (Kiss & Magnin. ´ fecture des Bouches-du-Rho ˆne Centre informatique de la Pre ´the ´e: La banque de donne ´es sur les incendies de (2002) Prome ˆts en Re ´gion me ´diterrane ´enne en France. have different vegetation cover histories.A.A. R. Cameron. by P.e. ACKNOWLEDGMENTS This study was carried out in the framework under the GIS ´ cologique et paysager des incendies sur les garrigues ‘Impacts e Journal of Biogeography 31. and to Markus Pfenninger for his judicious and helpful comments on the manuscript. R. (2003) Land-snail diversity in a square kilometre of Cretan maquis: modest species richness. Arche ´ologie et Environnement: de la Ste au XVIIIe sie Victoire aux Alpilles (ed. although this site has less significant indices with the sampling sites located in the northern part of the area studied. Amouric. (1976) British land snails. Loisel. that constitutes dispersal vector for the invasive species Xeropicta derbentina (Labaune & Magnin. i. including fire regime. ´ographie.. 131–142. J. Centre National De La Recherche Scientifique. & Redfern.e. disturbance and isolation. and of current human activities. Bishop. 2003). current communities are the product of the past. garrigue. 16. (1987) Impact of forest fires on structure and architecture of Mediterranean ecosystems. (1934) The habitats of land mollusca in Britain. Moreover. 145–157. 39–50. Its geographical location and the Mediterranean gradient may explain partially these observations. & Quezel. ´ de Provence. P. have particular species compositions because they are similar not only to the northern sampling sites but also to most of the southern ones (i. In conclusion. Amouric.D. Malacologia. 1995. A. CC1995) are exposed to passage of animal herds.. We would Bouches-du-Rho like to thank Isabelle Girard.D. Cameron.. Academic Press. 13. M. the same pre-fire landscape structure seems to influence species composition but also indicates a certain persistence of malacofaunas after disturbance. The land snail communities from one of the northern sites that burned once in 1991 (i. Cameron. Aix-en315–373. (1993) Calissanne et Merveille. Cameron. Tubingen.D. approche e ´cologique et e ´voluBlondel.R. Bonin. (1998) A spatially precise study of Holocene fire history. provided that disturbance regime is not maintained over years or that time elapsed between two successive disturbances is long (Nekola. Cameron & Greenwood. R. 2003). Kiss & Magnin.e.com..promethee. although all the southern sampling sites. http:// Fore www. post-fire disturbances also affect community composition and response patterns. K. Marsaille. Mollusca: Gastropoda. Ophrys.J. C. Thus. M. & Greenwood. the response patterns obtained must be due to cryptic refuges located within burned areas that allow malacofaunas to remain for years after successive fires. pp.. 87–93. Masson. Carcaillet. ª 2004 Blackwell Publishing Ltd ˆ ts pe ´ ri-marseillaises’ (IMEP-CNRS/CEMAGREF) and et les fore ´ ne ´ ral des was supported by funds provided by the Conseil Ge ˆ ne and by Unitas Malacologica. K. Cameron.. Narration. & Vardinoyannis. However. deux domaines ´ conomie de l’Etang de Berre (de la fin du Moyen Age dans l’e ` cle). J. Journal of Molluscan Studies. Paris. 69. Current postfire gastropod communities seem to be determined by numerous factors and are simultaneously subjected to biogeographical influences (bioclimatic gradient and/or historical biogeography) and to recent history. they all show identical pre-fire vegetation structure. 1984. Publications Universite Provence. F. K. A. We are indebted to Marjorie Sweetko for her careful re-reading and her help in translation. involving Mediterranean species introduction and landscape modifications in the Mediterranean basin (Mylonas. R. Biological Journal of the Linnean Society. significant Sørensen indices with all sites). 1999). Boycott. J. & Pannett. D. Triantis. hight density and local homogeneity. 384–396. 1997. Secondly.E. 61–66. R. 22. & Vardinoyannis. (1975) Carte Climatique de seille. K. Virginie Libois and Sylvie Marguerier for assistance in the fieldwork. no clear trends in response patterns of gastropod communities after fire have been pointed out.N. which have high Sørensen index with this northern site. 13. 1997. Journal of Ecology.A. Theler. M. H. 1984. Leveau and M. 2002). Edn. REFERENCES ` l’e ´preuve du temps. Mylonas.A. Miglioretti. 75–87. M. as to various other anthropogenic disturbances (Shikov. Ecologia Mediterranea.

1801) and C.J. Annales de la Socie ´ cial La Garrigue.. (1991) Mollusques continentaux et histoire qua´ diterrane ´ ens (Sud-Est de la France. Leiden. 11–24. 67–74. F. G. Elsevier Science B. Emberger. A. Emberger. M. Solem and A. Backhuys.D. Unpublished Thesis. 15. (1976) Les Paysages Forestiers de Haute ProDougue vence et des Alpes Maritimes. F. A.. J. 611–628. Sauvage. 102–110. R. Long. (1984) The role of disturbance in natural communities.N. Re ´ rou. Magnin. Van Bruggen). (2000) Wildland fire in ecosystems. & Kapler Smith. D. Kiss et al. E. Rocky Mountain Research Station.. L. M. 147. (2003) Evidence for ecological speciation in the sister species Candidula unfasciata (Poiret. A.. New York. R. Z. Kiss. Fire and ecosystems (ed. (1987) The role of fire from paleoecological data. J. (1968) Code ´ me ´thodique de la ve ´ge ´tation et du milieu. Fort Collins. E. 52. Telfer. dite ‘‘Carte de Cassini’’. Centre d’Ecologie et des Ressources Renouvelables.K. (2001) Spatial patterns of forest Dı fires in Catalonia (NE of Spain) along the period 1975–1995. Trabaud and R. E. Wacquant.L. Walla Walla. 1836). pp. & Magnin. & Johannes. ternaire des milieux me ´ d’Aix-Marseille Catalogne). (1974) Carte de la ve Marseille. L. Shikov. 59–70. Van Bruggen). 145–157. (1999) Guide ´. by A. Ecologia Mediterranea. & Legendre. E. M. I. & Smith. (2002) Effects of fire management on the richness and abundance of central North American grassland land snail faunas. Academic Press.P. graphies ae Jongman. Centre National De La Recherche Scientifique. Edn. J. Cameron. pp. O. 13.. Annual Review of Ecolology and Systematics. & Pons. Monspeliensia. A.P. C. (1995) Interior Columbia Basin mollusc species of special concern. Ecologia Mediterranea. Prodon). & Bertrand. Leiden. P. M.. Kozlowski and C. (1995) Data analysis in community and landscape ecology. Interior Columbia Basin Ecosystem Management Project. F. 157–171. Paris. & Trabaud.J. J. Cambridge.F. & Magnin. by A. Forest Ecology and Management. Schreiner. (1991) Randomization and Monte Carlo methods in biology. Forest Ecology and Management. 34. F. WI. 3. 237– 248. (1987) Vegetation wildfires in the MediLe Houe terranean basin: evolution and trends. ´ de la vie animale dans la garDelmas. (1999) Un escargot nouveau venu dans le Luberon et en Provence: Xeropicta derbentina (Krinicki. 353– 391. CO. T. Centre National de la Recherche re Scientifique. Statistiques Logiciel Pe ´ dagogie. 69. Cambridge University Press. Fire and biological processes (ed.. Gastropoda). Nekola. C. 79. 249–259. Analysis of the vegetation recovery after fire. W. F. M. 156 Legendre. by L. (1984) Effects of land use changes on the land mollusc fauna in the central portion of the Russian plain. d’exploration des donne Ivry-sur-Seine. Kerney. Institut Ge Mylonas. Geographie – Ecologie – Histoire. Leiden. Amsterdam. N° spe ´az-Delgado.. rigue. Masson & Cie. (1958) La diversite ´te ´ d’Histoire Naturelle de l’He ´rault. (2000) Wildland fire in ecosystems. Delachaux et Niestle Lausanne. J. (2002) Impact of fire on land snail communities in the French Mediterranean region: preliminary results. Effects of fire on fauna. Ahlgren).. Frest. ´ge ´tation de la France. R.G. X. Edn.F. ´ droit. Martin. (1974) Effects of fire in the Mediterranean Region. Second English edn. Solem and A. 155–163. (2003) The impact of fire on some land snail communities and patterns of post-fire recolonisation. 141..A.H. ´ ographique National (1999) Carte ge ´ome ´trique de la Institut Ge France. Courrier Scientifique du Parc Naturel ´gional de Luberon. Eppenstein. F. Brill.S. Magnin.. London.. 53–66. L. of human impacts and climatic changes. 25. Actes du Colloque: Dynamiques ´diterrane ´ens environnementales et Histoire en domaines me (Paris 2002) (in press). Animal Biodiversity and Conservation. Paris. pour le releve C. ª 2004 Blackwell Publishing Ltd .T. Sousa. pp. Toulouse. N° Hors se Huff. F. Godron. 197–213. Naturalia plus particulie ´ rie. Daget. pp. EDISUD. M. World-wide snails – biogeographical studies on non-marine Mollusca (ed. Journal of Biogeography 31. World-wide snails – biogeographical studies on non-marine Mollusca (ed. Journal of Molluscan Studies.C. ´ ographique National (2001) Campagne de photoInstitut Ge ´riennes 1949 a ` 1998. US Department of Agriculture Forest Service. & Van Tongeren. le logiciel ´es.J. Universite II. L.N. (1971) Travaux de Botanique et d’Ecologie. (2003) Land snails and human impact: the temporal resolution of the Holocene assemblages. Effects of fire on fauna. n° 74 Molinier. (1998) Numerical ecology. 1831) (Helicellinae. Kiss. Paris. & Rey. Aix-en-Provence. & Kiss. ´ centes Guillerm.C. & Thinon.. teer Braak.C. Aix-en-Provence. (1984) The influence of man: a special problem in the study of the zoogeography of terrestrial molluscs on the Aegean islands. Final Report Contract #430E00-4-9112. F. (2001) A comparison of the medieval and Lloret. General Technical Report RMRS-GTR-42. Pons. I. C. H. US Department of Agriculture Forest Service. 3–11. Paris. Lyon. S..G. Dupias.R. L. ´ ographique National. (1992) The dynamics of land gastropod communities in Mediterranean France during the Holocene: effects ´soge ´e. & Magnin.N. Naveh. (1980) Les interventions re ´ ge ´ tation au nord de la me ´ diterrane ´ e et de l’homme sur la ve ` rement dans le sud de la France.. L. 401–434. Le Floc’h. Biological Journal of the Linnean Society.J.N. L.R. P.J. by T. Labaune. M.. Marseille. Rocky Mountain Research Station. Brill. E. CO. Huff. (1985) Document pour un zonage des ´gions phyto-e ´cologiques. J. ´.J.L.. Magnin. Chapman and Hall. & Poissonnet. Pfenninger. R.F.E. P.V. M. Me Manly.G. General Technical Report RMRSGTR-42. Fort Collins. SLP Statistique Jambu (1994) STATlab by slp. G. E. des escargots et des limaces d’Europe. G.H. & Magnin. B. & Marı the current fire regimes in managed pine forest of Catalonia (NE Spain). 13. J. rugosiuscula (Michaud. 43–53.

Fe ¨ ller 1774) Pomatias elegans (O. Mediterranean and West-European range. F. 147. 117–136. Mediterranean range. F.R. Fe Cochlostoma patulum (Draparnaud 1801) ¨ ller 1774) Cryptomphalus aspersus (O. His main research topics are response patterns of land snail communities to anthropogenic disturbances and climate change. island area and habitat availability determine land snail species richness of Aegean islands. 26. (1976) Inflammabilite ` ces des garrigues de la re ´ gion me ´ diterrane ´ enne. F. Pfeiffer 1850) Candidula unifasciata (Poiret 1801) ¨ ller 1774) Cecilioides acicula (O. Cambridge. Mu Lauria cylindracea (da Costa 1778) ¨ ller 1774) Merdigera obscura (O. the Faculte France). 25–37. F. & Williams. Appendix 1 List of species sampled within the 12 sampling sites and classified into five biogeographical groups. P. (1987) Dynamics after fire of scerophyllous plant communities in the Mediterranean basin. Holarctic range. Journal of Biogeography. The Nautilus. 11. F. European range.L..Response patterns of land snail communities to fire disturbances Theler. Mu Helicigona lapicida (Linnaeus 1758) Hypnophila boissiy (Dupuy 1850) Granaria variabilis (Draparnaud 1801) Granopupa granum (Draparnaud 1801) ¨ ller 1774) Jaminia quadridens (O. Welter-Schultes. (1995) The ecology of fire.W. Mu Microxeromagna armillata (Lowe 1852) Monacha cantiana (Montagu 1803) ¨ ller 1774) Monacha cartusiana (O. Journal of Biogeography 31. Ecologia Mediterranea. ª 2004 Blackwell Publishing Ltd 157 . Biogeographical range Abida polyodon (Draparnaud 1801) ¨ ller 1774) Acanthinula aculeata (O. 110. 239– 249. M.. ` . R. 3–14. H. Ogheri. Cambridge University Press. France) des Sciences et Techniques de St Je whose research topics include ecological modelling and multivariate statistical analysis. Lloret. Mu Candidula gigaxii (L. J. E. ´ de ´ ric Magnin is a researcher in landscape ecology at Dr Fre ´ des Sciences et Techniques de St Je ´ ro ˆ me (Marseille. F. Mu ¨ ller 1774) Eobania vermiculata (O. 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 ˆ me (Marseille. L. Mu Vitrea contracta (Westerlund 1871) ¨ ller 1774) Vitrea crystallina (O. M. (2001) Positive Vila fire-grass feedback in Mediterranean Basin Woodlands. M. F. Forest Ecology and Management. ´ Dr Franck Torre is a biostatistical researcher at the Faculte ´ ro ˆ me (Marseille. F. E. 145–157. Mu Cepaea nemoralis (Linnaeus 1758) Cernuella virgata (da Costa 1778) ` re 1972) Chondrina avenacea (Bruguie ´ russac 1807) Clausilia rugosa (A. Trabaud. Mu Oxychilus alliarius (Miller 1822) ¨ ssler 1838) Oxychilus hydatinus (Rossma Oxychilus draparnaudi (Beck 1837) Papillifera solida (Draparnaud 1805) ´ russac 1807) Phenacolimax major (A. Mu ¨ ller 1774) Euconulus fulvus (O. F. F. J. (1997) The modern terrestrial gastropod (land snail) fauna of western Wisconsin’s hill prairies. L. cipales espe Acta Oecologica-Oecologica Plantarum. M/WE. F. paleoecology. 13. 111–121. Palearctic range.J. & Terradas. ´ et combustibilite ´ des prinTrabaud. 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. (1999) History. F. Whelan. France) with main research interest in the Je impact of fire on Mediterranean land snail communities and in their patterns of post-fire recolonization. historical biogeography and invasion biology.