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Global Ecology and Biogeography Letters (1998) 7, 97–113
RESEARCH ARTICLE
Delimitation of biogeographical areas in the Iberian
Peninsula on the basis of Helicoidea species
(Pulmonata: Stylommatophora)
ANA I. PUENTE, KEPA ALTONAGA, CARLOS E. PRIETO and ANA RALLO Laboratory of
Zoology, Department of Zoology and Animal Cell Dynamics, Faculty of Sciences, University of the Basque
Country, P.O. Box 644, E-48080 Bilbao, Spain; e-mail: ggpalsuk@lg.ehu.es (K. Altonaga)
Abstract.
A sectorization of the Iberian Peninsula Baleary. This province is characterized by species of
Mediterranean affinities but also contains some specieshas been obtained on the basis of Helicoidea species
distributions. The distributions were analysed by of Septentrionalaffinities because this province includes
the north of the Iberian mountain range, whichmultivariate methods with several qualitative indices,
applied to the matrix obtained by dividing the study constitutes a very important refuge area for several
species of septentrional origin. (4) Province ofarea into 113 squares of 100×100 km. Dendrograms
were obtained by the agglomerative procedure Extremadura-La Mancha-Andalusia, with forty-six
species and divided into the subprovinces ofUPGMA. The analysis identified five biogeographical
provinces subdivided into subprovinces and sectors. Extremadura-La Mancha-Andalusia and Almerı
´a
(represented by only one square). The first subprovinceThey are as follows. (1) Basque-Pyrenean-Catalonian
province, with seventy-nine species and divided into is divided into the sectors of Andalusia-La Mancha
and Extremadura-La Mancha. (5) Lusitanian province,the sectors Basque Septentrional, Basque Meridional,
Pyrenean and Catalonian-Provenzal. This province with fifty-six species, divided into Galician-Asturian,
Portuguese and Duerense sectors, and characterizedseems to be a contact area between the three
malacofaunistic groupings of western Europe. (2) by species of Lusitanian and Mediterranean affinities.
Province of Castilla la Vieja, with thirty-six species
and divided into Northern and Southern sectors. (3)
Key words.
Pulmonata, Helicoidea, biogeo-
graphical sectorization, endemic species, IberianMediterranean province, with sixty-seven species and
divided into the sectors of Ebro Valley, Levantine and Peninsula.
INTRODUCTION
Characterization of malacogeographical areas
according to the different distribution patterns of the
The first step in the interpretation of faunistic variation species has only recently been undertaken and, in the
within a territory involves characterization of different Iberian Peninsula, has been limited to small or medium-
areas on the basis of taxon distribution maps. Students sized regions (Prieto, 1986; Faci, 1991; Mun
˜oz, 1992;
of vertebrate biogeography have long since blazed a Puente & Prieto, 1992a, c; Prieto et al., 1992; Altonaga
trail for regional analyses and they have also shown et al., 1994; Hermida, Outeiro & Rodrı
´guez, 1994;
the way for statistical investigations on their data, Prieto & Sevillano, 1994). Apart from the work by
made possible thanks to powerful software for data Hidalgo (1875), in which the Iberian Peninsula was
management and statistical analysis (Andre
´, 1984). For divided into various regions depending on the species’
instance, concerning the Iberian Peninsula, see Doadrio distribution patterns, only Sacchi (1957, 1964) and
(1988). However, students of invertebrates, with their Andre
´(1984) have carried out more extensive analyses.
However, both Sacchi and Andre
´restricted theirproblems of sampling, sorting, identification, and
diversity, have not reached that stage of regional analyses to the Mediterranean regions. Sacchi (1957,
1964) set up a sectorization without a hierarchicalanalysis.
97
1998 Blackwell Science Ltd
98 Ana I. Puente et al.
organization, based on more or less subjective the Cantabrian cornice to Cantabria Province; the
second comprises the species which are found alongappraisals, and characterised by the presence of typical
elements or endemic species. Andre
´(1984) carried out the western peninsular edge, from Asturias and Galicia
to central or southern Portugal, including the westerna quantitative and hierarchical analysis based on
similarity indexes between provinces or departments part of the Duero plateau; the third pattern comprises
the species which present an area of distribution whichof 125 species from over 800 localities. The aim of the
present work is to undertake a malacogeographical includes southern Portugal, part of Extremadura and
western Andalusia, and, occasionally, northernsectorization of the whole Iberian Peninsula and
Balearic Islands area based on numerical analysis of Morocco. Lastly, the Mediterranean grouping includes
xerophilous species which are distributed throughoutthe distributions of the Helicoidea species.
the peninsular regions neighbouring the
Mediterranean. This grouping is the best represented
in the Iberian Penisula because the greater part of the
STUDY AREA
peninsular surface is subject to a Mediterranean climate
(Rivas-Martı
´nez, 1987). This vast region has a veryWithin Europe, the Iberian Peninsula and the Balearic
Islands constitute a very attractive malacological study complex fauna (Sacchi, 1964) and there coexist species
whose distribution patterns are diverse.area, not only because of their high number of species
but also because of their singularity (degree of The degree of endemicity is very large in the study
area. In fact, of the 141 species of Helicoidea consideredendemicity) and the diversity of their malacofauna.
This can be explained by the location of the study area, as present in the Iberian Peninsula and Balearic Islands
(Puente, 1994a), eighty-nine are endemic, whichwhich constitutes a bridge between Europe and Africa,
and between the Eurosiberian and Mediterranean represents an endemicity level of 63%. These endemics
are included in twenty-six genera, of which sixteen arebiogeographical regions, and by the fact that it
harbours a great heterogeneity of habitats as a result exclusive to the study area. The zones which harbour
the majority of endemics are the Pyrenees, theof its complex historical, climatic and orographic
characteristics (Sacchi, 1964). The geology and soils Cantabrian mountain range, the southeastern
peninsular mountainous regions, the Algarve, theare also greatly variable.
The Iberian Peninsula harbours representatives of mountainous regions of western Andalusia, the Iberian
mountain range and the Duero plateau (Puente &the Helicoidea superfamily from the three
malacofaunistic groupings pertaining to western Prieto, 1992b).
Europe as described by Germain (1930): Septentrional
or European, Atlantic or Lusitanian and
Mediterranean. The Septentrional grouping is made
MATERIALS AND METHODS
up of hygrophylic elements whose distributions range
throughout central Europe. According to Sacchi (1964), The analyses presented here are based upon the
distribution maps of the species of the Helicoideaits southwestern border is located in the Ebro valley
(see Fig. 1). In the Iberian Peninsula, the species superfamily present in the Iberian Peninsula and
Balearic Islands. These maps were made usingbelonging to this grouping are distributed along the
Pyrenees, the Catalonian coast, the Cantabrian biogeographical data and biological material gleaned
in various sampling surveys. The material was collectedmountain range, and the northern part of the Iberian
mountain range, which is located to the south of the between April 1987 and December 1990 in 1051
localities (Fig. 2). Of these, 476 localities wereEbro river, and can be called a relictic zone. The
Atlantic grouping also includes hygrophylic species homogenously distributed in the northern-central part
of the Iberian Peninsula (Altonaga et al., 1994). Thewhose common distribution area includes the
Moroccan coast, Portugal, Galicia, the Cantabrian other 575 localities were mainly spread over the least
known areas of the Iberian Peninsula in order to reachcornice, the French coast up to Brittany, Cornwall, the
western part of Wales and the western part of Ireland a similar level of malacological knowledge in the whole
Peninsula (Puente, 1994a, b). The number of species(Caziot, 1915; Castillejo, 1981). In the Iberian
Peninsula, species distributions of this grouping seem considered for the study area was 141; this figure
represents the present-day fauna of Helicoidea speciesto demonstrate three patterns (Puente, 1994a): the first
pattern corresponds with the area from Galicia and in that area (Puente, 1994a).
1998 Blackwell Science Ltd, Global Ecology and Biogeography Letters,7, 97–113
Biogeographical areas in the Iberian Peninsula 99
Fig. 1.
Map of the study zone, showing some regions and the main geographic features.
Fig. 2.
Map showing records of Helicoidea in the Iberian Peninsula. Each spot corresponds to one UTM square of 20×20 km.
Black spots contain localities sampled in this work; empty ones show localities cited in the literature.
For biogeographical analyses, the study area was Balearic Islands (Fig. 4B). These dimensions were
chosen because they gave a regular network with adivided into 113 squares, each of 100km×100 km,
although the area is less in the case of cells high enough number of squares, each of which was
large enought to contain meaningful data.corresponding to the edge of the area, those adjacent
to the coordinate lines and those encompassing the Species were recorded as present or absent and those
1998 Blackwell Science Ltd, Global Ecology and Biogeography Letters,7, 97–113
100 Ana I. Puente et al.
present in only one square were discarded from the The proposed biogeographical hypothesis is based
on the dendrogram obtained using the Jaccard indexmatrix to reduce distortion in the analysis. The data
matrix (113 zones and108 species) was used to generate (JI) (Fig. 4A), which produced the best correlation (r=
0.725, compared with 0.686 for Ochiai, 0.684 for Dice,Jaccard, Dice, Kulczynski (coefficient no. 2) and Ochiai
index values for each grid cell pairing (Rohlf, 1989), and 0.676 for Kulczynski). The dendrograms obtained
from the other indices did not vary significantly fromas recommended by Huba
´lek (1982) for binary data
analyses. Huba
´lek (1982) also cites the Phi index which, that based on the Jaccard index. In looking for
coherency in the definition of the groupings, andhowever, was not used as it does take into account
double absences, because, as Buser & Baroni-Urbani following similar protocols to biogeographical analyses
based on other groups (Hagmeier & Stults, 1964), we(1982) stated, double absences of species do not give
any biogeographical information. Hengeveld (1990) considered those groupings which presented a JI value
between 0.29 and 0.41 as being provinces orargues that for presence-absence data, Jaccard is one
of the coefficients that gives the fewest problems, and subprovinces, and those with a JI>0.43 as sectors (with
one exception for the southern sector of the provincethe results are easy to interpret. Dendrograms were
obtained using the UPGMA grouping procedure from of Castilla la Vieja, as described below). Thus the study
area was divided into five large zones (Fig. 4B) whichthe NTSYS v.1.5 program (Rohlf, 1989).
A simple index was used to determine which species were given the category of provinces (cf. Sainz &
Herna
´ndez, 1985; Rivas-Martı
´nez,1987). Furthermore,characterized each one of the provinces, subprovinces
or sectors (Prieto & Sevillano, 1994), by dividing the square 56 was treated separately as it only contained
two common species, Cepaea nemoralis and Helixnumber of squares occupied by each species in a specific
province, subprovince or sector by the total number aspersa, due to the scarceness of samples in that square.
The fiveprovinces defined are:the Basque-Pyrenean-of squares which lie in the study area, and multiplying
the quotient by 100. When the result of the operation Catalonian province (BPC); the province of Castilla la
Vieja (CLV); the Mediterranean province (MED);is 100, it means that the species is exclusive to this
province, subprovince or sector. If the species is absent, the province of Extremadura-La Mancha-Andalusia
(EMA); and the Lusitanian province (LUS).the index score is 0. A value higher than 50 was
considered significant and thus the corresponding
species is deemed a ‘characteristic’ species of this zone.
Basque-Pyrenean-Catalonian province
Appendix 1 indicates the number of squares occupied
by each species in each defined biogeographical unit Made up of nineteen squares (JI separation level=
0.33), the Basque-Pyrenean-Catalonian provinceas well as in the entire study area.
comprises Gascogne, Languedoc, Pyrenees, the
Catalonian coastal mountain range, the Basque
mountains and the Ebro headwaters. It harbours
RESULTS AND DISCUSSION
seventy-four species and five endemics not considered
in the analysis because they were only found in one
Analysis of biogeographical similarity
square. These are Pyrenaearia cotiellae (square 24),
Hygromia gofasi,H. tassyi and Pyrenaearia parva (25),The number of species per square (richness) oscillates
between eight and thirty-five (except for square 56, and Trochoidea montserratensis (38). As exclusive
species it contains Norelona pyrenaica,Metathebawhich has two species, and square 26, with forty-one
species) (Fig. 3), with an average of 17.6 species per atacis,Trochoidea betulonensis,Mastigophallus
rangianus,Trissexodon constrictus,Candidula arganica,square. The highest richness values correspond to the
squares located, first, in the north of the Ebro valley, C. unifasciata,Helicella orzai,Cernuella aginnica,
Ciliella ciliata,Hygromia cinctella,Pyrenaeariaand second, in Asturias, central Leo
´n, Levantine
region, Gymnesian Islands, Andalusia (except for carascalensis,P. organiaca,P. velascoi,Montserratina
bofilliana,M. martorelli,Arianta xatarti and Chilostomasouthern Almerı
´a), and the central coast and southern
Portugal. Galicia, northern Portugal, Extremadura and squammatinum; these species, except C. unifasciata,C.
aginnica,C. ciliata,H. cinctella and C. squammatinum,both plateaus show low richness values which could
be related to the type of substrate (mostly non- are Iberian endemics. Concerning the characteristic
species of the Basque-Pyrenean-Catalonian province,limestone) and/or the climate (great thermal variations
and scarce precipitation). six belong to the Septentrional malacofaunistic
1998 Blackwell Science Ltd, Global Ecology and Biogeography Letters,7, 97–113
Biogeographical areas in the Iberian Peninsula 101
Fig. 3.
Map showing the number of species per 100×100 km square (richness).
grouping (Helicodonta obvoluta,Euomphalia strigella, the remaining two sectors: Elona quimperiana,
Oestophorella buvinieri,Candidula rocandioi,HelicellaTrichia hispida,Hygromia limbata,Helicigona lapicida,
and Cepaea hortensis), three are Mediterranean species gonzalezi,H. ordunensis,H. orzai,Pyrenaearia
schaufussi,P. velascoi and Ponentina subvirescens. The(Monacha cantiana,Cernuella neglecta and Xerosecta
explanata), and seven are Iberian endemics (Trochoidea species Ashfordia granulata,Oestophora silvae and
Mengoana jeschaui are only found in the Basquepallaresica,T. penchinati,T. ripacurcica,Helicella
gonzalezi,H. iberica,Pyrenaearia schaufussi and Septentrional sector and Helicella striatitala only in
the Basque Meridional sector. The Basque sectors seemChilostoma desmoulinsi). It also includes various
elements belonging to the Lusitanian or Atlantic to be a contact zone between the Septentrional and
the Atlantic groupings, and harbour various elementsgrouping (such as Elona quimperiana,Ashfordia
granulata,Oestophora barbula and Zenobiella belonging to the Duero plateau.
Likewise, there is a series of species which appear insubrufescens), and thus this province appears as a
contact zone between the malacofaunistic groupings of the Pyrenean and Catalonian-Provenzal sectors and
not in the other two. These are Norelona pyrenaica,western Europe, which is possible because of the great
heterogeneity of habitats that it includes. Metatheba atacis,Trochoidea pallaresica,Candidula
gigaxii,Helicella conspurcata,Cernuella neglecta,The Basque-Pyrenean-Catalonian province has been
divided into four sectors (Figs 4B and 5): the Basque Pyrenaearia organiaca,Montserratina martorelli,
Arianta xatarti,Chilostoma desmoulinsi,C.Septentrional sector (BS) (JI=0.5) (thirty-seven
species); the Basque Meridional sector (BM) (JI=0.57) squammatinum and Eobania vermiculata. Furthermore,
species found only in the Catalonia-Provenzal sector(forty species); the Pyrenean sector (PY) (JI=0.53)
(forty-four species plus four endemics); and the and not in the Pyrenean are Atenia quadrasi,Cochlicella
conoidea,Trochoidea pyramidata,T. trochoides,T.Catalonian-Provenzal sector (CP) (JI=0.51) (fifty-
three species plus one endemic). betulonensis,Caracollina lenticula,Mastigophallus
rangianus,Suboestophora tarraconensis,HelicellaThe Basque Septentrional and Basque Meridional
sectors share various species which are not present in huidobroi,H. madritensis,Microxeromagna armillata,
1998 Blackwell Science Ltd, Global Ecology and Biogeography Letters,7, 97–113
102 Ana I. Puente et al.
Fig. 4.
Dendrogram obtained (summarized) (A) and its geographical projection (B). Figures on the map indicate the 113 squares
of 100×100 km used for biogeographical analyses. BPC: Basque-Pyrenean-Catalonian province; CLV: province of Castilla la
Vieja; EMA: province of Extremadura-La Mancha-Andalusia; LUS: Lusitanian province; MED: Mediterranian province.
Xerosecta explanata and Montserratina bofilliana; with (Candidula sp., square 32). Only one species is exclusive,
Helicella silosensis, and another one is characteristic,the exception of M. rangianus, the rest of these species
belong to the Mediterranean grouping. Candidula rocandioi. The province has been divided
into two sectors (Figs 4B and 6): the Northern sector
(CS) (JI=0.53) (twenty-four species plus one endemic);
Province of Castilla la Vieja
and the Southern sector (CM) (JI=0.39) (twenty-four
species).The province of Castilla la Vieja is composed of twelve
squares (JI=0.37) and includes the eastern part of The Northern sector harbours species characteristic
of the Basque-Pyrenean-Catalonian and Lusitanianthe Duero plateau, part of the Central and Iberian
mountain ranges and the northeastern part of the provinces (see below), such as Elona quimperiana,
Helicodonta obvoluta,Helicella gonzalezi,HygromiaSouthern plateau. It harbours only thirty-five species,
and one endemic which was not included in the analysis limbata,Mengoana jeschaui,Pyrenaearia cantabrica,
1998 Blackwell Science Ltd, Global Ecology and Biogeography Letters,7, 97–113
Biogeographical areas in the Iberian Peninsula 103
Fig. 5.
Partial dendrogram for the Basque-Pyrenean-Catalonian Province, and the sectors considered: BM, Basque Meridional
sector; BS, Basque Septentrional sector; CP, Catalonian-Provenzal sector; PY, Pyrenean sector. Figures correspond to squares
used for biogeographical analyses. See Fig. 4B.
Fig. 6.
Partial dendrogram for the province of Castilla la Vieja, and the sectors considered: CM, Southern sector; CS, Northern
sector. Figures correspond to squares used for biogeographical analyses. See Fig. 4B.
Portugala inchoata,Ponentina subvirescens and In any case, according to the minimum spanning tree
obtained, this square shows maximum affinity with theHelicigona lapicida, which do not appear in the
Southern sector. adjacent square 58, and thus was included in the
Southern sector. The relatively isolated position ofThe squares of the Southern sector are related to
the north with a low affinity value (JI=0.39). However, square 59 within the sector can be explained by the
fact that it encompasses the southwestern edges of theif square 59 is eliminated, the value increases to 0.51.
1998 Blackwell Science Ltd, Global Ecology and Biogeography Letters,7, 97–113
104 Ana I. Puente et al.
Iberian mountain range, and so it contains exclusively species are Helicodonta obvoluta,Trichia hispida,
Hygromia limbata and Cepaea hortensis; also,or almost exclusively various species endemic in the
Peninsula or belonging to the Septentrional Euomphalia strigella and Helicigona lapicida are
relatively frequent (in five and four squares,malacofaunistic grouping (Euomphalia strigella,
Candidula camporroblensis,Cepaea hortensis; respectively). The presence of Septentrional species in
this sector is due to the fact that it includes part of theTrochoidea geyeri and Candidula rocandioi are also in
square 46; Candidula gigaxii is also in 58). Otherwise, Iberian mountain range which seems to be a relictic
area for many species. Among the Iberian endemicthe Southern sector includes species which mostly
belong to the Mediterranean grouping, such as elements, Trochoidea turolensis (exclusive to this sector),
T. ripacurcica,Candidula najerensis,C. rocandioi,Caracollina lenticula,Cernuella neglecta,
Microxeromagna armillata,Xerosecta promissa,Iberus Helicella iberica and H. ordunensis appear in the Ebro
Valley sector and not in the other two sectors whichgualtieranus and Theba pisana, as well as other typically
Septentrional species (e.g. Euomphalia strigella), or make up the province.
The Levantine sector, characterized by species of theother Atlantic species (e.g. Candidula intersecta) which
do not appear in the Northern sector. Mediterranean grouping, harbours Atenia quadrasi,
Monacha cantiana,Trochoideabarceloi (exclusive to this
sector), T. derogata (exclusive), Suboestophora boscae
Mediterranean province
(exclusive), Helicella huidobroi,Cernuella neglecta and
Xerosecta explanata, which are (with the exception ofComposed of twenty-three squares (JI=0.34), the
Mediterranean province includes the middle and lower the exclusive ones) species also found in theCatalonian
sector of the Basque-Pyrenean-Catalonian province.Ebro valley, the central and southeastern Iberian
mountain range, the eastern part of the Subbetica These species do not appear in the other two sectors.
Within this sector, square 50 is in a relatively isolatedmountain range, the Levantine region, and the Balearic
Islands. It is home to a total of fifty-six species, and position although it still has its maximum affinity with
square 62 according to the minimum spanning tree.also eleven endemics not included in this analysis,
which are: Pyrenaearia navasi (square 34), P. molae We believe this position is due to the fact that it is
a transition zone between the Ebro Valley and the(50), Trochoidea sp. (60), Suboestophora hispanica,S.
jeresae and S. kuiperi (83), Trochoidea caroli and T. Catalonian-Provenzal sectors.
Finally, the Baleary sector has three exclusive species:ebusitana (111), and Allognathus graellsianus,
Trochoidea boissyi and T. claudinae (112). Its exclusive Trochoidea nyeli,Ganula lanuginosa and Iberellus
minoricensis. Another species, Marmorana muralis,isspecies are: Trochoidea barceloi,T. derogata,T. grata,
T. nyeli,T. turolensis,Suboestophora boscae,Ganula only found here and in the Lusitanian province. Other
than the twelve species present in the three sectors oflanuginosa and Iberellus minoricensis, all of which
except G. lanuginosa are Ibero-Balearian endemics. The this province, the Baleary sector does not have any
species in common with the Ebro Valley sector, but itcharacteristic species of the Mediterranean province
are: Trochoidea elegans,T. pyramidata,T. trochoides, has six species in common with the Levantine sector.
The Ebro Valley and Levantine sectors have anotherT. murcica,T. geyeri,Suboestophora tarraconensis,
Candidula camporroblensis,C. najerensis,Helicella thirteen species in common.
huidobroi,Eobania vermiculata,Otala punctata and
Pseudotachea splendida, all of which belong to the
Province of Extremadura-La Mancha-Andalusia
Mediterranean grouping. The Mediterranean province
has been divided into three sectors (Figs 4B and 7): The province of Extremadura-La Mancha-Andalusia
consists of twenty-eight squares (JI=0.29), andthe Ebro Valley sector (EV) (JI=0.56) (thirty-seven
species plus two endemics); the Levantine sector (LE) includes the Toledo mountains, Sierra Morena, most
of the Southern plateau, the Guadalquivir basin, the(JI=0.47) (forty species plus four endemics); and the
Baleary sector (BA) (JI=0.65) (twenty-two species plus western part of the Subbetic mountain range and the
Penibetic mountain chain. It was divided into twofive endemics).
The Ebro Valley sector mainly contains subprovinces, Almerı
´a (square 109) and Extremadura-
La Mancha-Andalusia (the rest of the squares; JI=Mediterranean elements, and also Iberian endemics
and Septentrional species which do not appear in the 0.41) (Fig. 8). The Almerı
´a subprovince houses only
nine species plus two endemics not considered in theother two sectors. Some examples of Septentrional
1998 Blackwell Science Ltd, Global Ecology and Biogeography Letters,7, 97–113
Biogeographical areas in the Iberian Peninsula 105
Fig. 7.
Partial dendrogram for the Mediterranean province, and the sectors considered: BA, Baleary sector; EV, Ebro Valley
sector; LE, Levantine sector. Figures correspond to squares used for biogeographical analyses. See Fig. 4B.
Fig. 8.
Partial dendrogram for the province of Extremadura-La Mancha-Andalusia, and the sectors considered: ALM, subprovince
of Almerı
´a; AM, sector of Andalusia-La Mancha; EM, sector of Extremadura-La Mancha. Figures correspond to squares used
for biogeographical analyses. See Fig. 4B.
analysis, Helicella mariae and Xerosecta adolfi. The low number of species that it contains, perhaps because
of undersampling. In any case, according to thespecies that it contains are mostly widely distributed
Mediterranean ones (Caracollina lenticula,Cernuella minimum spanning tree, this square has its maximum
affinity with the adjacent square 108.virgata,Microxeromagna armillata,Iberus gualtieranus,
Otala lactea, and Theba pisana), but it has one species The Extremadura-La Mancha-Andalusia sub-
province contains thirty-nine species and five endemicscharacteristic of the Lusitanian province (Oestophora
barbula) and also Trochoidea cobosi (a Betic endemic) not considered in the analysis: Helicella gasulli (square
106), Theba sp. (108), and Oestophora calpeana,O.and Helicella stiparum (an Almeriense endemic), both
shared by the Manchego-Andalusian sector of the dorotheae and O. tarnieri (110). It only harbours four
exclusive species, Oestophora ortizi,Helicella zujarensis,Extremadura-La Mancha-Andalusia subprovince. The
relatively isolated position of square 109 is due to the Pseudotachea litturata and Theba andalusica, and its
1998 Blackwell Science Ltd, Global Ecology and Biogeography Letters,7, 97–113
106 Ana I. Puente et al.
characteristic species are Trochoidea cobosi,Gasulliella Guadiana rivers. It harbours forty-eight species, along
with eight endemics not considered in the analysis.simplicula,Suboestrophora gasulli,Candidula gigaxii,
Helicella cistorum,H. conspurcata,H. vatonniana, These are Cryptosaccus asturiensis (square 4),
Pyrenaearia daanidentata,P. oberthueri,P. poncebensisXerosecta promissa,X. reboudiana and Otala lactea. The
subprovince of Extremadura-La Mancha-Andalusia is and Plentuisa vendia (5), Helicella bierzona (16),
Candidula setubalensis (84) and C. codia (103). Thedivided into two sectors (Figs 4B and 8): the sector
of Extremadura-La Mancha (EM) (JI=0.49) (twenty- exclusive species found here are Gittenbergeria
turriplana,Oestophora lusitanica,Candidula belemensis,four species); and the sector of Andalusia-La Mancha
(AM) (JI=0.43) (thirty-eight species plus five C. olisippensis,Helicella corderoi,H. jamuzensis and H.
zaratei. The characteristic species are Elonaendemics).
The Extremadura-La Mancha sector is a fairly poor quimperiana,Ashfordia granulata,Oestophora barbula,
O. silvae,Candidula intersecta,Mengoana jeschaui,malacofaunistic zone as it contains only a few species,
none of which are exclusive and only two of which Pyrenaearia cantabrica,Portugala inchoata and
Ponentina subvirescens. Another effectivelyare characteristic: Helicella cistorum and Xerosecta
reboudiana. The soil in this sector is mostly non- characteristic species is Marmorana muralis, due to the
fact that of the five squares in which it was present inlimestone and the climate is severe which makes it
rather unfavourable for the flourishing of the helicoids. the whole study area, four belong to the Lusitanian
province. The province was divided into three sectorsAlso, its geographical location is rather far away from
the principal expansion zones of Helicoidea species. (Figs 4B and 9): the Galician-Asturian sector (GA)
(JI=0.43) (twenty-eight species plus five endemics); theThe species present in this sector, apart from the
characteristic ones cited above, belong to the Portuguese sector (PO) (JI=0.57) (thirty-one species
plus two endemics); and the Duero River sector (DU)Mediterranean grouping and are widely distributed
and able to tolerate ample variations of environmental (JI=0.43) (twenty-eight species plus one endemic).
The Galician-Asturian and Portuguese sectors sharefactors. These species are Monacha cartusiana,
Cochlicella acuta,C. barbara,Caracollina lenticula, various widely distributed Mediterranean species, such
as Cochlicella acuta,C. conoidea and Helicella apicina,Candidula gigaxii,Helicella apicina,H. conspurcata,H.
madritensis,Cernuella virgata,Microxeromagna which in northern Portugal and Galicia are restricted
to the coast. The Galician-Asturian and Duero Riverarmillata,Xerosecta cespitum,Helix aspersa,Otala
lactea and Theba pisana. It also houses species sectors share a Septentrional species (Helicella itala),
two Lusitanian ones (Elona quimperiana and Ashfordiacharacteristic of the Lusitanian province such as
Oestophora barbula,Candidula intersecta,Portugala granulata) and various northern peninsular endemics
such as Oestophora silvae,Oestophorella buvinieri,inchoata and Ponentina subvirescens, which, with the
exception of C. intersecta, are also to be found in the Mengoana jeschaui,Pyrenaearia cantabrica,Helicella
corderoi and H. ordunensis. Some of these are typicallysector of Andalusia-La Mancha. The Andalusia-La
Mancha sector presents the largest number of species Lusitanian while others are from the Cantabrian
mountain range or the edge of the Duero plateau.and harbours theexclusive elementsof thesubprovince.
Characteristic species are Trochoidea cobosi, The exclusive species of the Galician-Asturian sector
are Candidula rocandioi,Helicella gonzalezi,Suboestophora gasulli,Helicella vatonniana and
Xerosecta promissa. The majority of species are Pyrenaearia schaufussi and Zenobiella subrufescens. The
relatively isolated position of square 5 within the sectorMediterranean, although some are Atlantic
(Oestophora barbula,Portugala inchoata and Ponentina could be due to the fact that it is a transition zone
with the Basque-Pyrenean-Catalonian province. Thesubvirescens) and one is European (Cepaea nemoralis).
Portuguese sector has the following exclusive species:
Euomphalia strigella,Caracollina lenticula,Gasulliella
Lusitanian province
simplicula,Gittenbergeria turriplana,Suboestophora
gasulli,Candidula belemensis,C. gigaxii,C. olisippensis,The Lusitanian province is made up of thirty squares
(JI=0.35). In the north it includes the western part of Helicella vatonniana,Xerosecta promissa,X. reboudiana,
Marmorana muralis and Otala lactea, of which E.the Cantabrian mountain range, the western part of
the Duero plateau, the mountains of Galicia and the strigella is typically Septentrional and M. muralis is a
Tyrrhenian species; the rest belong to thewestern edges of the Central mountain range. In the
south it includes the lower valleys of the Tajo and Mediterranean or Atlantic groupings. Finally, the
1998 Blackwell Science Ltd, Global Ecology and Biogeography Letters,7, 97–113
Biogeographical areas in the Iberian Peninsula 107
Fig. 9.
Partial dendrogram of the Lusitanian province, and the sectors considered: DU, Duero River sector; GA, Galician-
Asturian sector; PO, Portuguese sector. Figures correspond to squares used for biogeographical analyses. See Fig. 4B.
Duero River sector has the following exclusive species: affinities with Catalonia and Tyrrhenian Islands.
Majorca harbours a distinctive and varied fauna dueHelicella jamuzensis,H. madritensis,H. zaratei and
Xerosecta cespitum, which belong to the Mediterranean to the heterogeneity of its habitats. (7) Plateau: zone
very poor in land molluscs because of the lack ofgrouping.
limestone soils, it presents biogeographic characteristics
similar to those of the Ebro basin because its climate
Comparison of the results obtained with
is continental and severe. Sacchi also distinguished the
previous analyses
region formed by Galicia and Portugal, which has two
types of fauna, first, that associated with forests withSacchi (1957, 1964) divided and subjectively
characterized the Mediterranean region of the Iberian non-limestone soil, and second, fauna associated with
dunes, where there are typically Mediterranean species.Peninsula based on its malacological fauna, and
established several parallels with distributions shown Our results confirm the hypothesis of Sacchi (1957,
1964) that Catalonia constitutes a separate region,by other groups of animals and plants. His divisions
were as follows. (1) Catalonia: characterized by its associated with the northern Peninsula but set apart
from the rest of the Mediterranean zones. Likewise,Septentrional species, which are also partly found in
the north and northwestern Iberian Peninsula, and Sacchi’s ‘Levante’ coincides with what we defined here
as the Levantine sector, except that in our analyses thewhose southern limit is the Ebro valley. (2) Levante
(Castello
´n, Valencia, Alicante and Murcia provinces), Pityusan Islands show more affinity with the eastern
Balearic Islands. However, our analyses do not confirmincluding the Pityusan and Columbretes Islands: it is
faunistically poor because the Septentrional species the relationship alluded to by Sacchi between his
‘Levante’ and the provinces of Cuenca, Teruel andare disappearing towards the south and the elements
endemic in southern Spain have not yet appeared. The Albacete. Aparicio (1986) also took issue with this
point. Moreover, Sacchi’s hypothesis that there is aprovinces of Cuenca, Teruel and Albacete were, in his
opinion, related to this region. (3) Eastern Andalusia similarity between the Plateau and the Ebro Valley
seems to be false in the light of our results. However,(Almerı
´a, Granada, Jae
´n and part of Ma
´laga provinces)
or ‘dry Andalusia’. (4) Western Andalusia or ‘humid our analysis does support the relationship between
Galicia and Portugal established by Sacchi. TheAndalusia’ (Ca
´diz and part of Ma
´laga provinces): with
affinities with the North African fauna. (5) differences found are probably a consequence of the
improved distributional data on which our analysesGuadalquivir basin (Huelva, part of Ca
´diz, Sevilla and
Co
´rdoba provinces): arid zone, subcontinental, and are based. In fact, Sacchi himself stressed that the
boundaries were only approximate, particularly in theextensively cultivated, its fauna is not very
characteristic but coastal species are widespread. (6) central part of Spain, the gastropod fauna of which
was little-known at the time.Eastern Balearic Islands (Gymnesian Islands): rich in
endemic elements but quantitatively poor on the whole Andre
´(1984) carried out a similar biogeographical
analysis of 829 localities in the Mediterranean region,with respect to all the faunistic groups, it presents
1998 Blackwell Science Ltd, Global Ecology and Biogeography Letters,7, 97–113
108 Ana I. Puente et al.
Fig. 10.
Sectorization of the Mediterranean region of the Iberian Peninsula according to Andre
´(1984).
by means of a computer assisted analysis based on a similarity with the Levantine region, both of which can
be grouped subsequently with the Eastern part of bothmeasure of the entropy between species and variables.
There were a total of 125 land mollusc species found, Castillas. In our case, the Ebro Valley and Levantine
sectors are also defined, but Teruel has been includedand fifty-one variables were gathered on these. Three
variables (provinces or departments, corological or in the Ebro Valley sector. The western Peninsula is
divided by Andre
´(1984) into nine regions whichecological sectors and vegetation series) were positively
correlated with the distribution areas of the species broadly coincide with our sectorization. First, Palencia
and Burgos form a region all by themselves in theand thus were selected for the analysis. In this manner,
fifteen groups were established, and of these, the analysis of Andre
´(1984), whereas we include these in
the province of Castilla la Vieja. Second, the fourthirteen which correspond with our study zone are
shown in Fig. 10. Andre
´divided the peninsular regions of Andre
´(1984) comprised in Portugal
correspond to the southern Lusitanian province in ourMediterranean region into two parts, western and
eastern. After successive divisions, the eastern case, but do not include Huelva. Third, Almerı
´ais
individualized both in our results (at least thePeninsula was parcelled into: (1) Languedoc-
Roussillon-Catalonia, (2) Arago
´n-Catalonia, (3) southeastern part) and those of Andre
´(1984). The
northern border of Andre
´’s Andalusian region is moreEastern part of both Castillas and, (4) Levante; and
the western Peninsula into: (5) Palencia-Burgos, (6) meridional than in our analysis, because our
Extremadura-La Mancha-Andalusia provinceWestern part of both Castillas, (7) Evora and Portalegre
(=Upper Alentejo), (8) Badajoz, (9) Setu
´bal (=W comprises Ca
´ceres, the western part of Toledo and
Ciudad Real, which in Andre
´’s analysis were placed inLower Alentejo), (10) Beja (=E Lower Alentejo) and
Huelva, (11) Algarve, (12) Andalusia and, (13) Almerı
´a. the zone named the Western part of both Castillas.
The sectorization obtained in the present paperAt first glance, there is a certain similarity between
the results of Andre
´(1984) and those reported here. corresponds well with our field work experience. Other
analytical methods would probably yield similar results,Region 1 of Andre
´(1984), which is separated from the
rest of the regions of the eastern Peninsula, corresponds although there could be a modification of the boundary
squares. On the other hand, more exhaustive samplingto the Catalonian-Provenzal sector which in our
analysis is joined with the Basque and the Pyrenean in certain regions, which are not very well known,
would provide a greater precision in the sectorizationsectors. His analysis also defined the zone
corresponding to the Ebro Valley, which shows profiled herein. For instance, Hermida et al. (1994)
1998 Blackwell Science Ltd, Global Ecology and Biogeography Letters,7, 97–113
Biogeographical areas in the Iberian Peninsula 109
fauna del a
´rea lusita
´nica: Moluscos terrestres de Galicia
carried out a biogeographical study of the terrestrial
(Familias Arionidae, Zonitidae y Helicidae). V Reunio
´n
gastropods of north-western Spain and found a small
Bienal RSEHN, Oviedo.
isolated Eurosiberian area in the south of Salamanca
Caziot, E. (1915) La faune terrestre lusitanienne. Soc. Linn.
province, more or less corresponding to square 55 in
Lyon,62, 43–65.
the present paper. Furthermore, dividing the area into
Doadrio, I. (1988) Delimitation of areas in the Iberian
smaller squares, whenever possible, or using other types
Peninsula on the basis of freshwater fishes. Bonn. zool.
of units (e.g. natural regions) would help refine the
Beitr. 39, 113–128.
Faci, G. (1991) Contribucio
´n al conocimiento de diversos
analysis.
moluscos terrestres y su distribucio
´n en la Comunidad
As a final comment we would like to quote Sainz &
Auto
´noma Aragonesa. Ph. Thesis (unpublished),
Herna
´ndez (1985), ‘the sectorization attained will not
University of Zaragoza.
necessarily have to be considered as more important
Germain, L. (1930) Mollusques terrestres et fluviatiles
than those proceeding from other points of view and
(Premie
`re partie). Faune de France 21, 477 pp. Kraus
methods, but, at least, it will question the exclusive
Reprint, Nendels.
acceptance of some of them and make it possible to
Hagmeier, E.M. & Stults, C.D. (1964) A numerical analysis
of the distribution patterns of North American
search for even more synthetic criteria in the future,
mammals. Syst. Zool. 13, 125–155.
which will well combine and converge the different
Hengeveld, R. (1990, repr. 1992) Dynamic biogeography,
approximations’.
249 pp. Cambridge University Press, Cambridge.
Hermida, J., Outeiro, A. & Rodrı
´guez, T. (1994)
Biogeography of terrestrial gastropods of north-west
Spain. J Biogeogr. 21, 207–217.
ACKNOWLEDGMENTS
Hidalgo, J.G. (1875) Cata
´logo iconogra
´fico y descriptivo
de los moluscos terrestres de Espan
˜a, Portugal y las
This research was supported by a predoctoral research
Baleares. Parte 1, 224 pp.+la
´m. 1–24. Imprenta Segundo
grant awarded to Ana I. Puente by the Department of
Martı
´nez, Madrid.
Education, Universities and Research of the Basque
Huba
´lek, Z. (1982) Coefficients of association and
Government, and by projects X-86.044 to the same
similarity, based on binary (presence-absence) data: an
evaluation. Biol. Rev. 57, 669–689.
Department and PB88-0397, PB89-0081 and PB92-
Mun
˜oz, B. (1992) Gastero
´podos terrestres (Mollusca,
0121 (‘Fauna Ibe
´rica’) of the Spanish General
Gastropoda, Pulmonata) de Ca
´ceres, Badajoz y Huelva.
Directorate for Scientific and Technical Research
Ph. Thesis (unpublished), University Complutense of
(DGICYT).
Madrid.
Prieto, C.E. (1986) Estudio sistema
´tico y biogeogra
´fico de los
Helicidae sensu Zilch, 1959–60 (Gastropoda: Pulmonata:
Stylommatophora) del Paı
´s Vasco y regiones adyacentes.
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APPENDIX 1.
Number of squares per defined sector and subprovince in which each species occurs (even those which appear
in only one square are included), and the total number of squares occupied by each of the zones.
BPC CLV MED EMA LUS Total
BS BM PY CA CS CM EV LE BA EM AM ALM GA PO DU squares
Elona quimperiana (Fe
´russac, 1821) 4 2 – – 2 ––––––– 7– 1 16
Norelona pyrenaica (Draparnaud, 1805) – – 1 2 –––––––– ––– 3
Helicodonta obvoluta (O.F. Mu
¨ller, 1774) 22551– 1––––– ––– 16
Atenia quadrasi (Hidalgo, 1885) – – – 2 – – – 2 –––– ––– 4
Euomphalia strigella (Draparnaud, 1801) 1144–– 51–––– – 1– 17
Ashfordia granulata (Alder, 1830) 2 ––––––––––– 6– 1 9
Metatheba atacis (Gittenberger & De – – 2 1 –––––––– ––– 3
Winter, 1985)
Monacha cartusiana (O.F. Mu
¨ller, 1774) 43664798– 35– 245 66
Monacha cantiana (Montagu, 1803) 2 – 2 4 – – – 2 –––– ––– 10
Cochlicella acuta (O.F. Mu
¨ller, 1774) 4 – 2 6 – – 6 10 3 1 10 – 6 12 – 60
Cochlicella conoidea (Draparnaud, 1801) – – – 5 – – – 4 3 – 8 – 2 8 – 30
Cochlicella barbara (Linnaeus, 1758) 431534 91031115– 698 91
Trochoidea elegans (Gmelin, 1791) – 3 3 6 – – 7 7 3 – 2 – – – – 31
Trochoidea pyramidata (Draparnaud, – – – 4 – – – 5 3 – 1 – – – – 13
1805)
Trochoidea trochoides (Poiret, 1789) – – – 6 – – – 5 3 – – – – – – 14
Trochoidea barceloi (Hidalgo, 1878) ––––––– 4–––– ––– 4
Trochoidea betulonensis (Bofill, 1879) – – – 2 –––––––– ––– 2
Trochoidea boissyi (Terver, 1839) –––––––– 1––– ––– 1
Trochoidea caroli (Dohrn & Heynemann, –––––––– 1––– ––– 1
1862)
Trochoidea claudinae (Gasull, 1963) –––––––– 1––– ––– 1
Trochoidea cobosi (Ortiz de Za
´rate, 1962) –––––––––– 41––– 5
Trochoidea derogata (Rossma
¨ssler, 1854) ––––––– 5–––– ––– 5
Trochoidea ebusitana (Hidalgo, 1869) –––––––– 1––– ––– 1
Trochoidea geyeri (Soo
´s, 1926) – 1 3 – – 2 8 ––––– ––– 14
Trochoidea grata (Haas, 1924) –––––– 11–––– ––– 2
Trochoidea montserratensis (Hidalgo, – – – 1 –––––––– ––– 1
1870)
Trochoidea murcica (Guirao in –––––– 311–– 2– ––– 16
Rossma
¨ssler, 1854)
Trochoidea nyeli (Mittre, 1834) –––––––– 2––– ––– 2
Trochoidea pallaresica (Fagot, 1886) – – 3 2 – – 1 2 –––– ––– 8
Trochoidea penchinati (Bourguignat, – 3 3 4 – – 7 1 –––– ––– 18
1868)
Trochoidea ripacurcica (Bofill, 1886) – 1 2 – – – 2 ––––– ––– 5
Trochoidea turolensis (Ortiz de Za
´rate, –––––– 3––––– ––– 3
1963)
Trochoidea sp. –––––– 1––––– ––– 1
1998 Blackwell Science Ltd, Global Ecology and Biogeography Letters,7, 97–113
Biogeographical areas in the Iberian Peninsula 111
APPENDIX 1.
Continued
BPC CLV MED EMA LUS Total
BS BM PY CA CS CM EV LE BA EM AM ALM GA PO DU squares
Caracollina lenticula (Michaud, 1831) – – – 5 – 1 3 11 3 11 15 1 – 9 – 59
Gasulliella simplicula (Morelet, 1845) ––––––––– 13– – 3– 7
Gittenbergeria turriplana (Morelet, 1845) –––––––––––– – 3– 3
Mastigophallus rangianus (Fe
´russac in ––– 2–––––––– ––– 2
Deshayes, 1830)
Oestophora barbula (Rossma
¨ssler, 1838) – 1 – 1 – – – 1 – 11 4 1 6 12 11 48
Oestophora calpeana (Morelet, 1854) –––––––––– 1– ––– 1
Oestophora dorotheae Hesse, 1930 –––––––––– 1– ––– 1
Oestophora lusitanica (Pfeiffer, 1841) –––––––––––– 544 13
Oestophora ortizi De Winter & Ripken, –––––––––– 2– ––– 2
1991
Oestophora silvae Ortiz de Za
´rate, 1962 2 ––––––––––– 7– 1 10
Oestophora tarnieri (Morelet, 1854) –––––––––– 1– ––– 1
Oestophorella buvinieri (Michaud, 1841) 2 1 – – 2 ––––––– 3– 1 9
Suboestophora boscae (Hidalgo, 1869) ––––––– 4–––– ––– 4
Suboestophora gasulli (O. Za
´rate&O. ––––––––– 12– – 1– 4
Za
´rate, 1961)
Suboestophora hispanica (Gude, 1910) ––––––– 1–––– ––– 1
Suboestophora jeresae (Ortiz de Za
´rate, ––––––– 1–––– ––– 1
1962)
Suboestophora kuiperi (Gasull, 1966) ––––––– 1–––– ––– 1
Suboestophora tarraconensis (Aguilar- – – – 1 – – 2 1 –––– ––– 4
Amat, 1935)
Trissexodon constrictus (Boube
´e, 1836) 3 1 2 ––––––––– ––– 6
Candidula arganica (Servain, 1880) 3 3 1 ––––––––– ––– 7
Candidula belemensis (Servain, 1880) –––––––––––– – 5– 5
Candidula camporroblensis (Fez, 1944) ––––– 141–––– ––– 6
Candidula codia (Bourguignat, 1859) –––––––––––– – 1– 1
Candidula gigaxii (Pfeiffer, 1848) – – 5 3 – 2 – – – 4 11 – – 2 – 27
Candidula intersecta (Poiret, 1801) 4243– 2––– 4–– 5129 45
Candidula najerensis (Ortiz de Za
´rate, – 3 1 – – 1 5 ––––– ––– 10
1950)
Candidula olisippensis (Servain, 1880) –––––––––––– – 5– 5
Candidula rocandioi (Ortiz de Za
´rate, 1 2 – – 4 2 1 ––––– 1–– 11
1950)
Candidula setubalensis (Pfeiffer, 1850) –––––––––––– – 1– 1
Candidula unifasciata (Poiret, 1801) 1 – 5 3 –––––––– ––– 9
Candidula sp. –––– 1––––––– ––– 1
Helicella apicina (Lamarck, 1822) 2 – – 5 – – – 1249– 411– 38
Helicella bierzona Gittenberger & –––––––––––– –– 1 1
Manga, 1977
Helicella cistorum (Morelet, 1845) –––––––––123– – 21 18
Helicella conspurcata (Draparnaud, 1801) – – 1 6 – – – 1 2 12 14 – – 7 1 44
Helicella corderoi Gittenberger & Manga, –––––––––––– 2– 1 3
1977
Helicella gasulli Ortiz de Za
´rate, 1950 –––––––––– 1– ––– 1
Helicella gonzalezi (Azpeitia, 1925) 1 3 – – 3 ––––––– 1–– 8
Helicella huidobroi (Azpeitia, 1925) – – – 1 – – – 5 – – 3 – – – – 9
Helicella iberica (Rambur, 1869) 1 3 1 – – – 2 ––––– ––– 7
Helicella itala (Linnaeus, 1758) 4364475––––– 4– 3 40
Helicella jamuzensis Gittenberger & –––––––––––– –– 5 5
Manga, 1977
Helicella madritensis (Rambur, 1868) – – – 23767– 34– –– 5 37
Helicella mariae Gasull, 1972 ––––––––––– 1––– 1
1998 Blackwell Science Ltd, Global Ecology and Biogeography Letters,7, 97–113
112 Ana I. Puente et al.
APPENDIX 1.
Continued
BPC CLV MED EMA LUS Total
BS BM PY CA CS CM EV LE BA EM AM ALM GA PO DU squares
Helicella ordunensis (Kobelt, 1883) 2 3 – – 4 2 1 ––––– 2– 2 16
Helicella orzai Gittenberger & Manga, 2 1 –––––––––– ––– 3
1981
Helicella silosensis Ortiz de Za
´rate, 1950 –––– 31–––––– ––– 4
Helicella stiparum (Rossma
¨ssler, 1854) –––––––––– 11––– 2
Helicella striatitala Prieto, 1985 – 1 – – 1 ––––––– ––– 2
Helicella vatonniana (Bourguignat, 1867) –––––––––– 3– – 1– 4
Helicella zaratei Gittenberger & Manga, –––––––––––– –– 2 2
1977
Helicella zujarensis Ortiz de Za
´rate, 1950 –––––––––– 3– ––– 3
Plentuisa vendia Puente & Prieto, 1992 –––––––––––– 1–– 1
Trichia hispida (Linnaeus, 1758) 4364423––––– ––– 26
Cernuella aginnica (Locard, 1882) 4 – 5 3 –––––––– ––– 12
Cernuella neglecta (Draparnaud, 1805) – – 4 3 – 1 – 1 –––– ––– 9
Cernuella virgata (Da Costa, 1778) 436626911310151 4123 95
Ciliella ciliata (Studer, 1820) – 1 3 1 –––––––– ––– 5
Cryptosaccus asturiensis Prieto & Puente, –––––––––––– 1–– 1
1994
Ganula lanuginosa (Boissy in Guerin- –––––––– 2––– ––– 2
Meneville, 1835)
Hygromia cinctella (Draparnaud, 1801) 2 – – 1 –––––––– ––– 3
Hygromia gofasi Prieto & Puente, 1992 – – 1 ––––––––– ––– 1
Hygromia limbata (Draparnaud, 1805) 43641– 2––––– ––– 20
Hygromia tassyi (Bourguignat, 1884) – – 1 ––––––––– ––– 1
Mengoana jeschaui (Ortiz de Za
´rate, 2 – – – 1 ––––––– 3– 2 8
1949)
Microxeromagna armillata (Lowe, 1852) – – – 4 – 7 8 11 1 12 13 1 – 4 7 68
Pyrenaearia cantabrica (Hidalgo, 1873) –––– 1––––––– 2– 1 4
Pyrenaearia carascalensis (Fe
´russac, 1 1 3 ––––––––– ––– 5
1821)
Pyrenaearia cotiellae (Fagot, 1906) – – 1 ––––––––– ––– 1
Pyrenaearia daanidentata Raven, 1988 –––––––––––– 1–– 1
Pyrenaearia molae Haas, 1924 ––––––– 1–––– ––– 1
Pyrenaearia navasi (Fagot, 1907) –––––– 1––––– ––– 1
Pyrenaearia oberthueri (Ancey, 1884) –––––––––––– 1–– 1
Pyrenaearia organiaca (Fagot, 1905) – – 2 1 –––––––– ––– 3
Pyrenaearia parva Ortiz de Za
´rate, 1956 – – 1 ––––––––– ––– 1
Pyrenaearia poncebensis Ortiz de Za
´rate, –––––––––––– 1–– 1
1956
Pyrenaearia schaufussi (Kobelt in 12–––––––––– 1–– 4
Rossma
¨ssler, 1876)
Pyrenaearia velascoi (Hidalgo, 1867) 2 1 –––––––––– ––– 3
Xerosecta adolfi (Pfeiffer, 1854) ––––––––––– 1––– 1
Xerosecta cespitum (Draparnaud, 1801) 4346489111910– –– 5 74
Xerosecta explanata (O.F. Mu
¨ller, 1774) – – – 4 – – – 3 –––– ––– 7
Xerosecta promissa (Westerlund, 1893) ––––– 1––– 414– – 4– 23
Xerosecta reboudiana (Bourguignat, 1863) ––––––––– 75– – 3– 15
Zenobiella subrufescens (Miller, 1822) 4 1 1 ––––––––– 6–– 12
Portugala inchoata (Morelet, 1845) –––– 1–––– 31– 71210 34
Ponentina subvirescens (Bellamy, 1839) 1 1 – – 1 –––– 56– 61211 43
Montserratina bofilliana (Fagot, 1884) – – – 2 –––––––– ––– 2
Montserratina martorelli (Bourguignat, – – 1 4 –––––––– ––– 5
1870)
Arianta xatarti (Farines, 1834) – – 1 1 –––––––– ––– 2
1998 Blackwell Science Ltd, Global Ecology and Biogeography Letters,7, 97–113
Biogeographical areas in the Iberian Peninsula 113
APPENDIX 1.
Continued
BPC CLV MED EMA LUS Total
BS BM PY CA CS CM EV LE BA EM AM ALM GA PO DU squares
Chilostoma desmoulinsi (Farines, 1834) – – 4 2 – – 2 1 –––– ––– 9
Chilostoma squammatinum (Moquin- – – 3 5 –––––––– ––– 8
Tandon, 1855)
Helicigona lapicida (Linnaeus, 1758) 13654– 41–––– 122 29
Marmorana muralis (O.F. Mu
¨ller, 1774) –––––––– 1––– – 4– 5
Allognathus graellsianus (Pfeiffer, 1853) –––––––– 1––– ––– 1
Cepaea hortensis (O.F. Mu
¨ller, 1774) – 365115––––– ––– 21
Cepaea nemoralis (Linnaeus, 1758) 43664893– 15– 71211 79
Eobania vermiculata (O.F. Mu
¨ller, 1774) – – 1 6 – – 9 8 3 – 1 – – – – 28
Helix aspersa O.F. Mu
¨ller, 1774 43564891131215– 71211 110
Iberellus minoricensis (Mittre, 1834) –––––––– 3––– ––– 3
Iberus gualtieranus (Linnaeus, 1758) – 2 – 1 – 5 9 11 – – 13 1 – – – 42
Otala lactea (O.F. Mu
¨ller, 1774) ––––––– 326121– 8– 32
Otala punctata (O.F. Mu
¨ller, 1774) – 3 2 4 – – 9 10 3 – 7 – – – – 38
Pseudotachea litturata (Pfeiffer, 1851) –––––––––– 2– ––– 2
Pseudotachea splendida (Draparnaud, – 3 3 6 – – 9 11 1 – – – – – – 33
1801)
Theba andalusica Gittenberger & Ripken, –––––––––– 2– ––– 2
1987
Theba pisana (O.F. Mu
¨ller, 1774) 4216– 171137141 6122 77
Theba sp. –––––––––– 1– ––– 1
Total species/sector or subprovince 37 40 48 54 25 24 39 44 27 24 43 11 33 33 29
Total species/province 79 36 67 46 56
1998 Blackwell Science Ltd, Global Ecology and Biogeography Letters,7, 97–113
