Snails and Fire in the Mediterranean

8/13/2019 Snails and Fire in the Mediterranean

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O R I G I N A LA R T I C L E

The role of landscape history and

persistent biogeographical patterns in

shaping the responses of Mediterranean

land snail communities to recent fire

disturbancesLaurence Kiss*, Frederic Magnin and Franck Torre

INT RO DUCT IO N

Fire is a major disturbance within Mediterranean ecosystems

and greatly affects their landscapes (Naveh, 1974; Trabaud,

1976; Blondel, 1995; Whelan, 1995; Lloret & Mar, 2001). In

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;

Le Houerou, 1987; Trabaud, 1987; Amouric, 1992; Whelan,

Institut Mediterraneen dEcologie et de

Paleoecologie, U.M.R. 6116 du C.N.R.S.,

Facultedes Sciences et Techniques de St

Jerome, France.

*Correspondence: Laurence Kiss, IMEP,

batiment Villemin, Domaine du Petit Arbois,

Avenue Louis Philibert, BP 80 Cerege 13545,Aix-en-Provence, Cedex 04, France.

E-mail: [email protected]

ABS T RACT

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 fireregime (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 Srensen 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.

Journal of Biogeography(J. Biogeogr.) (2004) 31, 145157

2004 Blackwell Publishing Ltd www.blackwellpublishing.com/jbi 145


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1995). Throughout the Mediterranean basin, fire regimes are

also related to human intervention (Naveh, 1974) dating to

Neolithic times (Guillerm & Trabaud, 1980; Mylonas, 1984;

Pons & Thinon, 1987; Carcaillet, 1998; Lloret & Mar, 2001).

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 wildfires (Barbero et al., 1987;

Le Houerou, 1987).

Although numerous studies treat the impact of fire regime

on vegetation (Barbero et al., 1987; Trabaud, 1987; Whelan,

1995; Daz-Delgado & Pons, 2001; Vila et al., 2001), this

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 fire on Mediter-

ranean land snail communities have demonstrated that

various response patterns of communities have been

obtained for the same fire regime (Kiss & Magnin, 2002,

2003). Elements of pre-fire gastropod communities seem to

persist within post-fire communities, whatever the age of thefire (Kiss & Magnin, 2002, 2003). Other factors are likely to

explain to some extent the malacofauna composition after

fire. 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

fire disturbance. 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, but also to estimate the

role of recent site history and the influence of biogeographical

gradient, within a priori a homogeneous area, in shaping the

response patterns of gastropod communities.

M E T HO DS

Study area

The study area is located in the departement des Bouches-

du-Rhone (Provence, south-eastern France) (Table 1; Fig. 1).

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 fires suffered over the sampling period (19732001),

the district and the massif where it is located, CC1971 being the

reference site.

District Fire dates Name of massif Code

Charleval 1971 Chane des Cotes CC1971

Lambesc 1995 Chane des Cotes CC1995

Rognes 197989 Chane des Cotes CC197989

Aurons 1991 Chane des Cotes CC1991

Lancon-de-Provence 198995 Chane des Cotes CC198995

Marseille 1997 Chane de lEtoile CE1997

Allauch 197997 Chane de lEtoile CE197997



Peynier 1998 Regagnas MR1998

Trets 1989 Regagnas MR1989

Cuges-les-Pins 1993 Ste Baume SB1993

Figure 1 Location map of study area and of

sampling sites. The 12 sampling sites are

called after their location and the fire dates.

CC, Chane des Cotes; CE, Chane de lEtoile;

MR, Regagnas; SB, Ste Baume.

L. Kiss et al.

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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

Stratified sampling was carried out according to fire regime

over the past 30 years, i.e. number of fires, fire intervals and

age of the last fire (Sousa, 1984; Whelan, 1995; Lloret & Mar,

2001). Twelve sites of various fire ages were selected, including

six sites which have burned twice or three times and five 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-fire vegetation

stands: grassland (Brachypodium retusum Beauvais), garrigue

(Quercus coccifera Linnaeus, Q. ilexLinnaeus, Ulex parviflorus

Pourret, Rosmarinus officinalis Linnaeus and Cistus albidus

Linnaeus), oak stand (Q. pubescensWilldenow) and pine stand(Pinus halepensisMiller) 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 post-

fire vegetation stands within each sampling site. Each sampling

point consisted of square of 25 m2, and data were recorded

on floristic 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

(Institut Geographique National, 2001) taken between 1950

and 1998, before each fire (Fig. 3) and using old maps dating

from the 18th century called Cassini maps (Institut Geograph-

ique National, 1999) (Fig. 4). These maps provide a reliable

insight into the vegetation structure and the human uses of

landscape at the end of the 18th century (Douguedroit, 1976).

Figure 2 Selected sampling sites, which are defined by the

number of fires suffered over the period studied (i.e. 19732001)

and fire interval type. In bold, selected fire dates. The samplings

were performed during Spring 2000 and 2001.

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.

Response patterns of land snail communities to fire disturbances

Journal of Biogeography31, 145157, 2004 Blackwell Publishing Ltd 147


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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 five upper centimetres of

soil. Soil sample treatment (sieved on a 0.5-mm mesh), snail

count and snail identification were performed in the laboratory.

Only fresh shells (i.e. with intact periostracum) and living

individuals were taken into account because they are represen-

tative of current communities. The list of species sampled

follows the nomenclature of Kerneyet al.(1999) (Appendix 1).

All species collected were classified into five biogeographical

groups (Appendix 1). These groups take into account the

biogeographical range of species sampled according to Magnin

(1991) and Kerneyet al. (1999). The first group is composedof 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-fire patterns of responses of land snail communities were

analysed, using STATLAB software Ivry-sur-Seine, France (SLP

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 fire regime on land snail communities.

We also compared community composition within the area

studied taking into account the roles of fire 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=2abc;

where x1and x2are the two sites to be compared,a number

of species shared by the two sites, b andc number of species

unique to each of the sites.As a variant of the best-known similarity coefficient, i.e.

Jaccards coefficient, this coefficient 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 first and the

Figure 4 Cassini map of the sampling site

which burned in 1998. The area sampled

is located in the black square.

L. Kiss et al.



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second site and in both sites, allowed us to test the significance

of observed values (randomization test, n 1000 permuta-

tions) (Manly, 1991). The significance thresholds were

*P 0.05, **P 0.01 and ***P 0.001.

RE S ULT S

Impact of recent fire regimes on land snailcommunities

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 fire regimes

(Fig. 5).

Axis 1 (26.88% of total inertia) puts fallow land species

(Xeropicta derbentina), grassland or garrigue species (Candi-

dula 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 areclustered with sampling sites which have been exposed to

two successive fires within a 6-year interval (CC198995 and

CE19737997) (Fig. 5a,b), the species distribution is some-

what 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. armillataand C. unifasciata) are clustered on the negative

end (Table 2). Litter-dwelling species and more closed garrigue

species (V. costataand 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 south-

eastern part of the study area, according to a northsouth

gradient (Table 3; Fig. 5b).

This CA demonstrates neither a fire age gradient nor a fire

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 gradientfrom north to south, representing the distance inland from the

coast.

A Srensen matrix was also compiled to compare compo-

sition 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 significance (P 0.001). However, when sampling

sites of these two groups are compared, Srensen indices are

low, i.e. S 0.62 0.06 on average, with low or no signifi-

cance (i.e.P 0.05 andP 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 fire 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 fire interval

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, Cryptom-

phalus 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 nar-

bonensis; VCS, Vallonia costata; ZAL, Zonites algirus; XCE, Xero-

secta cespitum; XCO, Xerotricha conspurcata; XDE, Xeropicta

derbentina. See Table 1 for sampling site codes.




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(CE19737997). Thus, its communities are closer to the

malacofaunas of a southern sampling site, which has been

exposed to high fire 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 biogeographi-

cal gradient. Nevertheless, a high fire regime (i.e. short fire

interval and high number of fires) seems to modify malaco-

fauna composition in a comparable direction, whatever their

geographical location.

Biogeographical composition of malacofaunas under

different fire 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 locationhave 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 five groups. Mediterranean species and Mediterranean/

West European species are highly predominant in the high fire

regime sampling site (i.e. CC198995). In fact, this site is

composed of 50% Mediterranean species and of 30% Medi-

terranean/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 fire regime sampling site

which contains a higher proportion of Mediterranean species

than the other sites, the various fire regimes occurring since the

beginning of the 1970s do not seem to have affected

biogeographical composition of malacofaunas. Thus, the

Table 2 Significant species variables on axes

1 and 2 of the CA (46 species, 12 sampling

sites). Ctr., contribution.

Positive end Negative end

Axis 1 Candidula gigaxii (Ctr. 0.06) Lauria cylindracea (Ctr. 0.08)

Candidula unifasciata (Ctr. 0.03) Microxeromagna armillata (Ctr. 0.13)

Monacha cantiana (Ctr. 0.03) Vallonia costata (Ctr. 0.17)

Xeropicta derbentina (Ctr. 0.28)

Axis 2 Granopupa granum (Ctr. 0.03) Candidula unifasciata (Ctr. 0.05)

Vallonia costata (Ctr. 0.26) Cernuella virgata (Ctr. 0.03)

Microxeromagna armillata (Ctr. 0.38)Trochoidea pyramidata (Ctr. 0.06)

Trochoidea trochoides (Ctr. 0.27)

Table 3 Significant 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 Negative end

Axis 1 CC1995 (Ctr. 0.26) CE19798997 (Ctr. 0.24)

CC197989 (Ctr. 0.16)

Axis 2 CE19798997 (Ctr. 0.10) SB1993 (Ctr. 0.28)

CE1997 (Ctr. 0.33)

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 1

CC1995 0.78*** 1

CC197989 0.74*** 0.74*** 1

CC1991 0.67* 0.74*** 0.63* 1

CC198995 0.56 0.56 0.60 0.75*** 1

MR1989 0.62 0.58 0.62 0.72*** 0.76*** 1

MR1998 0.75*** 0.67* 0.71*** 0.71*** 0.62 0.85*** 1

CE1997 0.64* 0.60 0.64* 0.79*** 0.72*** 0.78*** 0.69** 1

CE197997 0.60 0.64* 0.64* 0.82*** 0.76*** 0.85*** 0.76*** 0.84*** 1

CE19737997 0.61 0.65* 0.70** 0.74*** 0.74*** 0.80*** 0.71*** 0.88*** 0.87*** 1

CE19798997 0.57 0.61 0.52 0.81*** 0.78*** 0.83*** 0.71*** 0.79*** 0.93*** 0.82*** 1

SB1993 0.54 0.58 0.62 0.71*** 0.71*** 0.77*** 0.68** 0.77*** 0.80*** 0.79*** 0.71*** 1

L. Kiss et al.



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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.

Certain sampling sites, that were composed of closed and

wooded landscapes in the beginning of the 1950s, have

retained a garrigue landscape since the first fire 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

stratified landscapes before their first fire (i.e. CC1991, SB1993,

CC1995, CC197989 and CE197997) (Tables 5 & 6).

Figure 6 Percentages of the number of

species of the five 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) 1950s 196070

CC1971 Homogeneous landscape

of open garrigue,

open wood and grassland

Homogeneous landscape

of high and closed wood

(b) 1950s 1970s

Pre-fire

(around 1 year before fire)

MR1989 Heterogeneous landscape

of garrigue, open

and closed wood

Heterogeneous landscape

of open and closed wood


of closed wood

CC1991 Heterogeneous landscape

of open wood, closed garrigue

with scattered trees and grassland



with scattered trees




SB1993 Heteroge neous land scap e


and garrigue with scattered trees


of open wood, garrigue

and garrigue with scattered trees


of closed and open wood,

garrigue and garrigue


CC1995 Homoge neous and open

landscapes of garrigue

and grassland


of garrigue and closed wood


of garrigue, closed and open wood

CE1997 Heterogeneous landscape

of garrigue, closed

and open wood


of open and closed wood,

and closed garrigue



of closed wood and closed

garrigue with scattered trees

MR1998 Heterogeneous landscape

of garrigue and open wood


of open and closed wood


of closed wood




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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 first fire

(CC1971, MR1989 and MR1998) (Table 5).

Only the sampling site, that has been exposed to high fire

regime (i.e. CC198995), has a different vegetation his-

tory (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.

Southern sampling sites, located in the Chane de lEtoile

massif, were composed of low landscapes of garrigues andgrasslands with few scattered trees (Table 7). Most of the

north-western sampling sites, located in the Chane des Cotes

massif, were composed of forests, woods and of some

cultivated areas, but the sampling site that has been exposed

to high fire 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.

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 fire regime site (i.e. CC1989

95) which appears to have been composed of garrigue since the

18th century.

DIS CUS S IO N

Impact of recent fire regimes on land snail

communities

As demonstrated before (Kiss & Magnin, 2002, 2003; Nekola,

2002), response patterns of land snail communities to fireregimes are not clear. Indeed, communities are not organized

according to a fire number gradient, a fire age gradient or a fire

interval gradient in the analyses. Only the high fire 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 fire has a drastic effect

on land snail communities (Kiss & Magnin, 2002, 2003), the

Table 6 Landscape and vegetation cover changes analysed using aerial photographs taken at various dates since the 1950s and before each

fire. (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 196070 Before the first fire Before the second fire

1949 1964 1978 1989


of open and closed garrigue,

grassland and low wood


of closed garrigue

with scattered trees,and closed and low wood


of open and closed woods


of closed wood and garrigue


1949 1968 1989 1993


of low and open wood,

grassland with scattered trees

Homogeneous, low and open

landscape with some areas

of high garrigue


of closed and highly

open garrigue

Homogeneous, low and

open landscape with some

areas of high garrigue

1950 1969 1978 1997

CE197997 Heterogeneous landscape

of open and closed wood,

and closed garrigue


of closed garrigue


and open garrigue



and grassland


of closed and high garrigue,

and open garrigue

(b) 1950s Before the first fire Before the second fire Before the third fire1950 1969 1978 1997

CE19737997 Homogeneous landscape

of low and closed wood


of high and closed garrigue


of low and open garrigue


of high and open garrigue


1950s 196070 Before the first fire Before the second fire Before the third fire

1950 1969 1978 1988 1997

CE19798997 Homogeneous

landscape of high

and closed wood

Homogeneous

landscape of high

and open wood

Homogeneous

landscape of high

and closed wood

Homogeneous

landscape of high

and closed garrigue

Homogeneous

landscape of high

and closed garrigue

L. Kiss et al.



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analyses performed (CA and Srensen matrix) show that thesecommunities are particularly resilient to low fire regimes over

the period 19732001, as the biogeographical composition of

malacofaunas does not change. In fact, some sampling sites

which have suffered one fire over the sampling period (i.e.

MR1989, MR1998, CE1997, CC1993 and CC1971) and the

sampling site which has burned twice with a long fire interval

(i.e. CE197997) are composed of the five biogeographical

groups, whatever the age of the last fire. Thus, other factors, i.e.

biogeographical or historical, seem to affect the response

patterns of gastropod communities.

Persistent biogeographical patterns

As expected, fire 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 fire regimes and have different last fire ages,

they have comparable species composition (high Srensen

indices) and similar percentages of the different biogeographi-

cal 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 first fire suffered over the

period studied, they are separated by the different analyses (CA

and Srensen matrix). It is clear that whatever the fire 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

the distance inland. Indeed, the sampling area is characterizedby 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 floristic 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 influences of a climate gradient

and of a floristic 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 micro-

regional scale (Cameronet 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 influence at one and the same time both

flora and malacofaunas.

Another hypothesis based on historical biogeography could

also explain, to some extent, this gradient. Generally, Medi-

terranean land snail communities are greatly diversified even

within a province (Cameron et al., 2000; Cameron et al.,

2003). Although communities have been sampled within arestricted area in our study, it was composed of four more or

less connected massifs (i.e. Chane des Cotes, Chane de

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

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 fire date and to their location within sampling area, i.e. (a) north-western sites located in the

Chane des Cotes massif (CC), (b) southern sites located in the Chane de lEtoile massif (CE), and (c) south-eastern sites located in the

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) CC198995 CC1995 CC1991 CC197989 CC1971

Vegetation cover Open Closed Open and closed Open and closed Closed

Types of vegetation or uses Heath Forest Cultivated area and forest Cultivated area and forest Forest

(b) CE1997 CE197997 CE19798997 CE19737997

Vegetation cover Open Open Open Open

Types of vegetation or uses vineyard and scattered trees Heath Heath Heath

(c) MR1998 SB1993 MR1989

Vegetation cover Closed Open Closed

Types of vegetation or uses Forest Wood and heath Forest




10/14

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 fire 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-fire 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 Mediter-

ranean and Mediterranean/West European species and its

malacofauna does not seem to be controlled by biogeographi-

cal influence like communities of the most northern sites.

Indeed, its landscapes are definitely 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

fires (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 biogeogra-

phical roles. Moreover, the persistence of Q. coccifera land-

scapes over centuries, due to these various disturbances,

probably induces a consistent land snail community composi-

tion in this sampling site.

While long-term historical explanations of differences in

fauna are generally acceptable on a biogeographical scale

(Cameronet al., 1980), historical processes on a local scale, as

it is demonstrated in this study, also seem to influence current

land snail communities. This history consists of fire 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 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 biogeo-

graphical gradient. In fact, numerous factors seem to control

response patterns of post-fire gastropod communities in

several sampling sites.

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 five biogeographical groups. More-

over, these sites show identical vegetation cover history before

the first fire 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 fire regime suffered in the Regagnas massif since 1973

(12 fires of more than 10 ha) (Centre informatique de la

Prefecture des Bouches-du-Rhone, 2002), these sites seem to

have been spared and forest cover has partially been main-

tained over the last decades. Thus, recent history and

geographical position seem to influence the community

composition of these sites.

All sampling sites of the Chane de lEtoile massif have

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 indi-

ces), they are not grouped in the CA. Thus, although fire

regime affects community composition, it seems that it is not

the main factor. 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.

These vegetation changes probably explain the differences

observed in malacofauna composition. The community com-

position of this massif is thus the result of the recent vegetation

history, of fire regimes over the last three decades and of

biogeography.

However, biogeographical and historical influences can be

observed in community response patterns because of malaco-

fauna 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 fire

severity and land snail location at the time of fire probably

induce numerous cryptic refuges (Kiss & Magnin, 2003). The

presence of these refuges permit land snails, first, to survive

during fire event and, secondly, to persist following (and

between) successive fires. Indeed, the persistence of shade-loving and mesophilous species, whatever fire age in the

Regagnas massif, indicates land snail survival within cryptic

refuges located in burned areas (Kiss & Magnin, 2002, 2003).

For example, two of the Chane de lEtoile sites (CE197997

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 fire regimes and landscape

histories.

Finally in some cases, the response patterns of communities

to various fire regimes also seem to be influenced by the

L. Kiss et al.



11/14

structure of pre-fire habitats and by post-fire 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. significant Srensen indices with all sites).

There are two possible explanations for this result: first,

although all the southern sampling sites, which have high

Srensen index with this northern site, have different veget-

ation cover histories, they all show identical pre-fire vegetation

structure, i.e. garrigue. Thus, the same pre-fire landscape

structure seems to influence species composition but also

indicates a certain persistence of malacofaunas after distur-

bance. Secondly, although this site has less significant 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-firedisturbances also affect community composition and response

patterns.

CO NCLUS IO NS

Gastropod communities appear to be highly resilient to fire

(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 fire have been pointed out. Current post-

fire gastropod communities seem to be determined by

numerous factors and are simultaneously subjected to biogeo-

graphical influences (bioclimatic gradient and/or historical

biogeography) and to recent history, including fire 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 fires.

Moreover, current communities are the product of the past,

particularly of Neolithic times, and of current human activi-

ties, involving Mediterranean species introduction and land-

scape modifications 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.

ACK NO W LE DGM E NT S

This study was carried out in the framework under the GIS

Impacts ecologique et paysager des incendies sur les garrigues

et les forets peri-marseillaises (IMEP-CNRS/CEMAGREF) and

was supported by funds provided by the Conseil General des

Bouches-du-Rhone and by Unitas Malacologica. We would

like to thank Isabelle Girard, Cinderella Grout, Virginie Libois

and Sylvie Marguerier for assistance in the fieldwork. 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.

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B I O S K E T C H E S

Dr Laurence Kiss is a researcher in landscape and commu-

nity ecology at the Faculte des Sciences et Techniques de St

Jerome (Marseille, France) with main research interest in the

impact of fire on Mediterranean land snail communities and

in their patterns of post-fire recolonization.

Dr Frederic Magnin is a researcher in landscape ecology at

the Facultedes Sciences et Techniques de St Jerome (Marseille,

France). His main research topics are response patterns of land

snail communities to anthropogenic disturbances and climate

change, paleoecology, historical biogeography and invasionbiology.

Dr Franck Torre is a biostatistical researcher at the Faculte

des Sciences et Techniques de St Jerome (Marseille, France)

whose research topics include ecological modelling and

multivariate statistical analysis.

Appendix 1 List of species sampled within the 12 sampling sites

and classified into five biogeographical groups. E, European range;

H, Holarctic range; M, Mediterranean range; M/WE, Mediterra-

nean and West-European range; P, Palearctic range.

Biogeographical

range

Abida polyodon (Draparnaud 1801) M

Acanthinula aculeata (O. F. Muller 1774) PCandidula gigaxii (L. Pfeiffer 1850) E

Candidula unifasciata (Poiret 1801) E

Cecilioides acicula (O. F. Muller 1774) M/WE

Cepaea nemoralis (Linnaeus 1758) E

Cernuella virgata(da Costa 1778) M/WE

Chondrina avenacea (Bruguiere 1972) E

Clausilia rugosa(A. Ferussac 1807) E

Cochlostoma patulum (Draparnaud 1801) M

Cryptomphalus aspersus (O. F. Muller 1774) M/WE

Eobania vermiculata (O. F. Muller 1774) M

Euconulus fulvus(O. F. Muller 1774) H

Helicigona lapicida(Linnaeus 1758) E

Hypnophila boissiy(Dupuy 1850) M

Granaria variabilis (Draparnaud 1801) M

Granopupa granum (Draparnaud 1801) M

Jaminia quadridens (O. F. Muller 1774) M/WE

Lauria cylindracea(da Costa 1778) M/WE

Merdigera obscura (O. F. Muller 1774) E

Microxeromagna armillata (Lowe 1852) M

Monacha cantiana (Montagu 1803) M/WE

Monacha cartusiana (O. F. Muller 1774) M/WE

Oxychilus alliarius (Miller 1822) E

Oxychilus hydatinus (Rossmassler 1838) M

Oxychilus draparnaudi (Beck 1837) M/WE

Papillifera solida(Draparnaud 1805) M

Phenacolimax major(A. Ferussac 1807) E

Pomatias elegans (O. F. Muller 1774) M/WEPseudotachea splendida (Draparnaud 1801) M

Punctum pygmaeum (Draparnaud 1801) H

Solatopupa similis (Bruguiere 1792) M

Sphincterochila candidissima (Draparnaud 1801) M

Trochoidea elegans (Gmelin 1791) M

Trochoidea pyramidata (Draparnaud 1805) M

Trochoidea trochoides (Poiret 1789) M

Truncatellina callicratis (Scacchi 1833) M

Vallonia costata (O. F. Muller 1774) H

Vitrea contracta (Westerlund 1871) E

Vitrea crystallina (O. F. Muller 1774) E

Vitrea narbonensis(Clessin 1877) E

Xeropicta derbentina (Krynicki 1836) M

Xerosecta cespitum (Draparnaud 1801) M

Xerotricha conspurcata (Draparnaud 1801) M

Zonites algirus (Linnaeus 1758) M




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Documents

Snails and Fire in the Mediterranean