The European centipedes hitherto referred to Eurygeophilus, Mesogeophilus, and Chalandea (Chilopoda, Geophilomorpha): taxonomy, distribution, and geographical variation in segment number

We have carried out a comprehensive revision of the European centipede species currently assigned to Eurygeophilus, Mesogeophilus, and Chalandea. This was based on comparative morphological study of specimens from throughout the range of the group and representative of the different nominal taxa, as well as a critical evaluation of all relevant literature. We consider this group to be represented by a single genus Eurygeophilus Verhoeff, 1899 [ = Geophilus (Mesogeophilus) Verhoeff, 1901, n. syn.; = Chalandea Brölemann, 1909, n. syn.] including two morphologically clearly distinguished species, Eurygeophilus multistiliger (Verhoeff, 1899) [ = Eurygeophilus multistiliger velmanyensis Brolemann, 1926, n. syn.] and Eurygeophilus pinguis (Brölemann, 1898) [ = Geophilus (Mesogeophilus) baldensis Verhoeff, 1901, n. syn.; = Chalandea cottiana Verhoeff, 1938; = Chalandea cottiana var. castrensis Manfredi, 1948; = Chalandea scheerpeltzi Attems, 1952, n. syn.]. On the basis of analysis of both published and new records these two species appear to be geographically vicariant: E. multistiliger occurs mainly in Mediterranean woodlands of southern regions (the western part of the Iberian Peninsula, eastern Pyrenees and Sardinia), whereas E. pinguis occurs mainly in temperate deciduous woodlands in montane and more northern regions (a limited area in Great Britain, the Cantabrian Mountains, most of the western and central Pyrenees, Corsica and the entire Alps). Despite morphological uniformity throughout its range, E. pinguis shows a consistent geographical pattern in variation of the number of segments, the modal values being different between the three major areas: (1) Pyrenees, Cantabrian Mountains and Great Britain, (2) western and central Alps, and (3) eastern Alps.

known but most fascinating groups among the geophilomorphs of the western Palaearctic region. Their quite stout, ''swollen'' body and their delicate, slender poison claws are very distinct compared with those of all other geophilids, suggesting specialised, but as yet unknown, ecological and behavioural traits. In addition, they have hitherto been recorded from scattered localities in western Europe and their distribution appears puzzling from a biogeographical perspective.
Although the geophilid fauna of many of these areas, including Great Britain and the Alps, has been investigated in some detail over the last century, only very few records are currently available for this particular group. However, it is still unclear as to whether this is a consequence of genuine rarity, secretive habits, or both. In addition, the group is in need of a thorough taxonomic revision: three genus-group and seven species-group taxa have been described to date, but the validity of most of them has awaited critical evaluation.
For this reason we have carried out a comprehensive revision of the species of Eurygeophilus, Mesogeophilus, and Chalandea based on morphological examination of representative specimens from throughout the range of the group, together with a critical evaluation of the whole of the relevant literature. We have also updated and analysed the known geographic distribution of the species and investigated geographical variation in the number of trunk segments as possible evidence of phylogeographic patterns.

Materials and methods
We have examined directly a total of 22 specimens from 14 localities (see Appendix). Our material included the holotype of Geophilus (Eurygeophilus) multistiliger Verhoeff, 1899, which has never been adequately described before, as well as specimens recognisable as representative of almost all the nominal taxa of the group, based on morphology and provenance, namely Geophilus pinguis Brö lemann, 1898, Geophilus (Mesogeophilus) baldensis Verhoeff, 1901, Chalandea cottiana Verhoeff, 1938, Chalandea cottiana var. castrensis Manfredi, 1948, andChalandea scheerpeltzi Attems, 1952. The only nominal taxon for which we did not examine directly any specimens was Eurygeophilus multistiliger velmanyensis Brolemann, 1926 for which, however, we could rely on the detailed description and accurate illustrations published by Brolemann (1926aBrolemann ( , 1930. Our material also included representative specimens from all the disjunct areas of the geographical range of the group with the exception of Corsica, and particularly from different localities throughout the Alps. We were not able to locate the only specimen recorded from Corsica by Léger and Duboscq (1903), which has probably been lost, and field collection performed by one of us (L.B.) in that island failed to secure new material.
All specimens were studied under light microscopy after having been cleared in ethylene glycol with the exception of the holotype of G. (E.) multistiliger which was already mounted on a slide. Because of the scarcity of available material, detachment of the head and dissection of the mouth parts were performed on a few representative specimens only, leaving the others undamaged. One specimen was also studied through ''environmental'' scanning electron microscopy (Philips TM XL30).
Specimens were checked on a comparative basis for all morphological characters traditionally considered in the taxonomic literature, particularly those previously reported as diagnostic among the nominal taxa of the group. Many other characters were investigated for possible interspecific differences, sexual dimorphism, age-dependent changes and inter-individual variation. We also carried out an exhaustive bibliographic search to retrieve all published accounts of the relevant taxa.
For each valid taxon, we present here a full list of citations, including synonymies, as well as an emended diagnosis. Standard drawings of a representative specimen for each species were produced using a camera lucida.
The geographical distribution of the species and the variation in the number of leg-bearing segments have been analysed on the basis of all published records and on new data either collected by ourselves or provided by colleagues (see under Acknowledgements).

Taxonomy
Critical evaluation of the literature as well as the comparative study of representative specimens have allowed us to recognise two morphologically clearly distinct species, which we consider as belonging to a single genus.
Each of these species is very homogeneous throughout its geographical range. A geographically consistent variation was however found in the number of trunk segments, at least in one species (see below under Geographical variation). Despite the fact that segment number is only in part genetically determined in geophilid centipedes (Kettle and Arthur 2000), this geographic pattern suggests a phylogeographic differentiation, possibly deserving taxonomic recognition but in need of being investigated adequately. In any case, the diagnostic value of segment number will be very limited because of interindividual variation within each population and the overlap of the range of variation between different population clusters. We believe that further taxonomic division of either species lacks an adequate factual basis at present; accordingly, we consider many of the nominal taxa described so far as invalid.

Diagnosis
Head shield convex, slightly wider than long; lateral margins distinctly curved, convergent anteriorly; posterior margin truncate, slightly concave; no frontal line. Antennae 2.2-3.6 times as long as the head width. Clypeus with a pair of non-areolate semicircular areas along the posterior margin. Labrum mid-part with a few round-tipped teeth; side-pieces well delimited, each bearing a fringe of slender projections. Mandibles and maxillae as typical of geophilids; two pairs of lateral lappets on the first maxillae; claw of the second maxillae quite long but round-tipped. Forcipular segment stout; coxosternum two to three times wider than long, with lateral margins sinuous, without distinct chitin-lines; forcipular tergum with lateral margins convergent anteriorly, only slightly narrower than the following tergum; no teeth on the internal side of forcipules; basal article of forcipules shorter than wide; tarsungulum strongly narrowed close to the base, slender and flattened. Sternal pores in a narrow transverse band on the posterior part of each sternum from the first to the penultimate leg-bearing segment; no median sockets on the anterior margin of sterna. Praetergum of last leg-bearing segment not separated from pleurites. Last pair of legs distinctly swollen and bearing additional dense setae on the ventral side in the male, slender and without additional setae in the females; claw present. Gonopods as typical of geophilids.

Taxonomic and nomenclatural remarks
Mesogeophilus was described by Verhoeff (1901b) as a subgenus of Geophilus to include the only species Geophilus (Mesogeophilus) baldensis Verhoeff, 1901, which is its type species by monotypy. Mesogeophilus was then elevated to genus rank by Attems (1929), followed by most other authors, but the diagnosis of this nominal genus remained very vague indeed (Minelli 1992;Foddai et al. 1995). Mesogeophilus Verhoeff, 1901 is here synonymised with Eurygeophilus Verhoeff, 1899 since we recognize the type species of the former as a junior synonym of Geophilus pinguis Brö lemann, 1898 (see remarks under E. pinguis) and we include this latter species in the genus Eurygeophilus (see below).
Chalandea was described by Brö lemann (1909a) to include only the species Geophilus pinguis Brö lemann, 1898, which is its type species by monotypy. Chalandea was regarded as a genus by most authors; only Verhoeff treated it as a subgenus of Geophilus in one of his major works (Verhoeff 1902-25). Chalandea Brö lemann, 1909 is here synonymised with Eurygeophilus Verhoeff, 1899 because the type species of both nominal taxa share some major characters which differentiate them from all other geophilids such as to justify their inclusion in the same genus. Among these characters are the general body shape and the unusual shape of the forcipular segment. Despite the fact that the phylogenetic relationships within the geophilids are still unresolved Minelli 1999, 2000;Edgecombe and Giribet 2004) and thus the current taxonomic arrangement of this group remains in need of a thorough revision, some of the peculiar characters shared by these two species are very probably synapomorphic. All previous authors had indeed acknowledged that Eurygeophilus and Chalandea are closely related: Attems assigned G. pinguis to the subgenus Eurygeophilus in one of his earliest papers (Attems 1903) andBrolemann (1930) introduced the tribe Eurygeophilini to include Eurygeophilus and Chalandea.
Published diagnoses of Chalandea and Mesogeophilus are widely congruent, only differing in respect to the chitin-lines, which have been described as present and complete in the type species of Chalandea (Brö lemann 1909a(Brö lemann , 1930, absent in Mesogeophilus (Verhoeff 1901b(Verhoeff , 1902. This putative difference has been reported uncritically by subsequent authors (e.g. Attems 1929) even though some inconsistency has been highlighted (Verhoeff 1938;Koren 1986). Attems (1929Attems ( , 1947 indeed introduced another putative difference in relation to the trunk sterna, which he described as provided with tubercles in Chalandea but lacking any tubercle in Mesogeophilus. However, our examination of specimens from throughout the geographic range of E. pinguis, including some representing G. (M.) baldensis, revealed that chitin-lines are invariantly weakly evident and the sternal tubercles invariantly present (see also remarks under E. pinguis). Attems (1929Attems ( , 1947 also considered Chalandea as having anterior sternal sockets, but we did not find such sockets in any specimen. According to the diagnoses of Eurygeophilus previously published (Verhoeff 1899(Verhoeff , 1902Attems 1903Attems , 1929Brolemann 1930), the poison calyx is characteristically composed of an anterior swollen part and a posterior slender part and the forcipular coxosternum is provided with shortened chitin-lines. However, direct examination of the holotype of G. (E.) multistiliger has revealed that these diagnoses were incorrect in respect of both points (see remarks under E. multistiliger). Chitin-lines, in particular, are only weakly marked and their appearance is therefore similar to what is observed in E. pinguis ( Figure 3).

Diagnosis
See Table I and Figures 9-13.

Taxonomic and nomenclatural remarks
The correct spelling of the specific epithet is ''multistiliger'', which is the name originally used in the description of the species (Verhoeff 1899) and also applied in the original label of the holotype. The name is from the Latin word ''stilus'', referring to the specialised setae on the trunk sterna. The alternative spelling ''multistyliger'' was invariably used by H.-W. Brö lemann and by some other authors but has to be considered an unjustified emendation.
Eurygeophilus multistiliger velmanyensis was described by Brolemann (1926a) based on two specimens from ''Velmanya'' in the eastern Pyrenees. Relying only on the brief original description of G. (E.) multistiliger published by Verhoeff (1899), Brolemann (1926a, 1930 established the new subspecies on a few putative differences compared with the nominotypical subspecies concerning the shape of the sternal pore groups and the pattern of coxal pores. However, our direct examination of the holotype of G. (E.) multistiliger reveals that its original description was incorrect in certain respects. Looking at the detailed description and careful illustrations of one of the two syntypes of E. multistiliger velmanyensis provided by Brolemann (1926aBrolemann ( , 1930, we found that these latter specimens agreed completely in their morphology with the holotype of G. (E.) multistiliger, even in the case of the putative differential characters mentioned above apart from some obvious sexually dimorphic traits. The only real difference is in the segment number, but we do not regard this character alone as taxonomically significant since inter-individual and geographical variation is commonly found within individual geophilid species and even within the related E. pinguis (see below). Consequently, Eurygeophilus multistiliger velmanyensis Brolemann, 1926 is here synonymised with Geophilus (Eurygeophilus) multistiliger Verhoeff, 1899. The original description of the holotype of E. multistiliger by Verhoeff (1899) was inaccurate or incorrect in relation to the following points: the head was described as being as long as wide, but it is actually slightly wider than long; the ''Endgliedorgane'' (probably the apical sensilla of the antennae) were described as absent but are actually present and visible; the chitin-lines were described as shortened but are actually weakly visible and uniform along the entire length of the exposed part of the coxosternum (see also remarks under Eurygeophilus); the poison calyx was described as composed of an anterior swollen part and a posterior slender part but the left calyx (the only visible one) is only apparently shaped as described (Figures 5, 6) because of a probable secondary displacement, while it is actually quite uniform in width and slightly S-shaped and thus similar to that known in E. pinguis; the sternal pore fields were described as ''Haufen'' and ''Häuflein'' by Verhoeff (1899) and these terms were interpreted by Brolemann (1926aBrolemann ( , 1930 as indicating quite broad groups of pores, but the pore fields are actually transverse narrow bands like those described in E. multistiliger velmanyensis (Figure 7); the most anterior coxal pores were described as grouped together, but they actually open independently of each other and are scattered along an arc from the dorsal to the ventral side, resembling those described in E. multistiliger velmanyensis (Figure 8); the female gonopods are described as absent but are actually present in the shape typical of female geophilids ( Figure 8).

Distribution (see Appendix and Figure 19)
Western part of the Iberian Peninsula (one locality), eastern Pyrenees including the neighbouring Sierra de Montseny (three localities), south-eastern part of Sardinia (one locality).

Diagnosis
See Table I

Taxonomic and nomenclatural remarks
Geophilus (Mesogeophilus) baldensis was described by Verhoeff (1901b) based on one specimen from ''Mori'', near Trento, in the Italian Pre-Alps. No other specimen was ever referred to this nominal taxon and its identity with Geophilus pinguis has indeed been suspected even though not formally recognised before (Minelli 1981;Foddai et al. 1995). Geophilus (Mesogeophilus) baldensis Verhoeff, 1901 is here recognised as a synonym of Geophilus pinguis Brö lemann, 1898 based on critical evaluation of its original description as well as on the examination of representative specimens of E. pinguis from throughout its range including four specimens of both sexes from La Marzola, no more than 30 km from the type locality of G. (M.) baldensis. The only putative difference between the two species recognized in the literature was in the forcipular chitin-lines which were described and illustrated as completely absent in G. (M.) baldensis (Verhoeff 1901b) but invariantly present and complete in G. pinguis both from the Pyrenees (Brö lemann 1898a, 1930 and from Great Britain (Jones and Barber 1997). However, our direct examination of representative specimens revealed that chitin-lines are actually poorly evident in all specimens of E. pinguis throughout its range including the Pyrenees and Great Britain (Figure 3 versus Figure 4). Differences described in the literature are thus very probably only due to different subjective interpretations by authors or to the different microscopic techniques used by them. Another minor putative difference between G. (M.) baldensis and G. pinguis was described by Verhoeff (1938) in the position of some coxal pores, but this is not significant in the light of inter-individual variability. Worth notice is the fact that the holotype of G. (M.) baldensis was considered lost by Foddai et al. (1995) but is actually still in existence in the collections of the Museum fü r Naturkunde, Humboldt-Universität, Berlin (Moritz and Fischer 1979). Chalandea cottiana was described by Verhoeff (1938) based on two females from ''Crissolo'', in the Cottian Alps. No other specimens were ever identified under this name and subsequent authors did not accept the validity of this taxon, considering it as a synonym of G. pinguis Zapparoli 1985, 1992;Barber 1992b;Foddai et al. 1995). Chalandea cottiana Verhoeff, 1938 is here confirmed as a synonym of Geophilus pinguis Brö lemann, 1898 on the basis of critical evaluation of its original description and direct examination of three topotypical specimens of both sexes. Putative differential characters of C. cottiana compared with G. pinguis were described by Verhoeff (1938) relying only on the description of the latter provided by Brö lemann (1898a, 1930). Labral teeth, setae on the second maxillae and coxal pores were described as more numerous in C. cottiana than in G. pinguis, but we found that the numbers of all these elements are sizerelated in E. pinguis as in other geophilid species; thus the different values were due to the different body size of the representative specimens, i.e. up to 28 mm long for C. cottiana versus up to 20 mm for G. pinguis (Brolemann 1930;Verhoeff 1938). Other minor differences were described in the shape of the intermediate article of the telopodite of the second maxillae, in the shape of the poison calyx and in the shape of the sternum of the last leg-bearing segment (Verhoeff 1938), but we found that these differences are not significant in comparison with the inter-individual variation in E. pinguis.
Chalandea cottiana var. castrensis was described by Manfredi (1948) based on one female from ''Gana di Sclés de Sota'' in the Orobian Pre-Alps. Subsequent authors did not accept the validity of this form and often considered it as a synonym of G. pinguis (Barber 1992b;Foddai et al. 1995). Chalandea cottiana var. castrensis Manfredi, 1948 is here confirmed as a synonym of Geophilus pinguis Brö lemann, 1898 on the basis of critical evaluation of its original description and direct examination of representative specimens of E. pinguis from throughout its range, including one specimen from Gorno, no more than 20 km from the type locality of C. cottiana var. castrensis. The only diagnostic characters described by Manfredi (1948) in respect to the typical C. cottiana were minor differences in the number and shape of the labral teeth, in the shape of the sternum of the last leg-bearing segment and in the position of the coxal pores relative to the sternum of the last leg-bearing segment. All these characters, however, are affected by slight inter-individual variation and developmental changes within E. pinguis (see above); thus the described differences lack taxonomic value.
Chalandea scheerpeltzi was described by Attems (1952a) based on two specimens from ''am Fusse des Jovanberges, im Obirstock'', in the Caravanche Alps. All the other few specimens collected in the eastern Alps have been so far referred to this nominal species (Koren 1986;Kos 1992aKos , 1996. Chalandea scheerpeltzi Attems, 1952 is here recognised as a synonym of Geophilus pinguis Brö lemann, 1898, on the basis of critical evaluation of its published descriptions (Attems 1952a;Koren 1986) and examination of specimens of E. pinguis from throughout its range, including one specimen from the Julian Alps which should be considered representative of C. scheerpeltzi because of its provenance and the number of segments. We found C. scheerpeltzi to be fully congruent in morphology with all other examined specimens of E. pinguis. Putative diagnostic characters of C. scheerpeltzi in respect to G. pinguis were described by Attems (1952a) relying only on the descriptions of the latter species provided by previous authors (Brolemann 1930;Verhoeff 1938). The antennal article XIV was said to be relatively longer in C. scheerpeltzi than in G. pinguis (Attems 1952a;Koren 1986), but we found that the elongation of this article in representative specimens of the former is well within the range of variation estimated for the latter (Figure 20). The forcipular pretergum was described as completely covered by the head shield in C. scheerpeltzi but partially exposed in G. pinguis (Attems 1952a), but this sclerite is actually visible from above to different degrees in different specimens of E. pinguis depending on the degree of contraction of the body articulations. Chitin-lines were described as absent in C. scheerpeltzi but present and complete in G. pinguis (Brö lemann 1898a, 1930), but they are actually indistinct in all the examined specimens of E. pinguis from throughout its range (see above; Figure 3). Anal pores were described as absent in C. scheerpeltzi but present in G. pinguis, but they are actually present in all specimens of E. pinguis examined by us (including the representative specimen of C. scheerpeltzi) even though sometimes covered by the gonopods. In considering the detailed description and illustration of C. scheerpeltzi provided by Koren (1986), other possible differential characters with respect to G. pinguis may be suspected in the number of projections of the labral side-pieces, in the elongation of the lappets of the first maxillae and in the shape of the sternum of the last leg-bearing segment; however, we found that all these characters are within the inter-individual variation of E. pinguis and thus lack taxonomic value. North Devon (12 localities), middle part of the Cantabrian chain (one locality), most of the Pyrenees with the exception of the easternmost part (23 localities), Corsica (one locality), Alps from Ligurian to Julian Alps (26 localities).

Phyletic relationships
Eurygeophilus represents a well-differentiated lineage within the diverse group of geophilomorphs currently recognised as the family Geophilidae s.l. (including Linoteniidae and Dignathodontidae; Attems 1929). It shares with other geophilids some major diagnostic characters involving the mandibles (each bearing only one pectinate lamella and no dentate lamella), the maxillary complex (with a typical shape and pattern of appendages) and the female gonopods (reduced to a short undivided lamina).
The phyletic position of Eurygeophilus within the geophilids is however hard to assess since neither morphological nor molecular analyses adequately resolve the internal phylogeny of this group (Foddai and Minelli 2000;Edgecombe and Giribet 2004). A close relationship to some lineages traditionally included in the Geophilinae (Geophilus and allies) is suggested by morphological characters such as the structure of the labrum and the pattern of the sternal pores. Indeed Eurygeophilus has been traditionally classified under this subfamily and was usually considered very close to Geophilus. However, other characters, such as the peculiar shape of the forcipular segment, resemble strongly those of Henia, a lineage traditionally classified in a different subfamily, Dignathodontinae (or family Dignathodontidae).

Geographical distribution
On the basis of both published and new records, Eurygeophilus appears to be limited to several disjunct regions of western and southern Europe (see Appendix and Figure 19).
Eurygeophilus multistiliger was known previously from only five specimens from four localities in the western part of the Iberian Peninsula and in the eastern Pyrenees, including the neighbouring Sierra de Montseny. The identification of a new specimen from south-eastern Sardinia is relevant in extending significantly the geographical range of the species.
Eurygeophilus pinguis has been known up to the present from some tens of specimens from a limited region of Great Britain and some localities in the Pyrenees, Corsica and the Alps. Our new records extend the known distribution of the species within the Pyrenees and the Alps and include the first record from the Cantabrian chain. In Great Britain E. pinguis appears to be restricted to an area in North Devon of less than 900 km 2 (see Barber 1992b); this limited occurrence is particularly striking in view of the extensive existing knowledge of the British centipede fauna and the indigenous status of the species in this region has been questioned Zapparoli 1985, 1992;Barber 1992a). In the Pyrenees, E. pinguis appears quite uniformly distributed along the main axis of the mountain chain from its western margin to the central part, the easternmost record being from the Ariège. The occurrence of the species in the Cantabrian chain is established by a new record from the Picos de Europa. Its occurrence in Corsica is based on an old, but reliable, record (Léger and Duboscq 1903). Within the Alps, E. pinguis occurs throughout the chain from the Ligurian and Maritime Alps in the west, to the Caravanche and the southernmost Julian Alps in the east; almost all records are from the southern marginal part of the chain.
The two species are apparently both geographically and ecologically vicariant: E. multistiliger occurs in more southern, Mediterranean regions, mainly in arid soils of sclerophyllous woods, whereas E. pinguis occurs in more northern, temperate regions, mainly in fresh and moist soils under beech and other broadleaf woodland. In Great Britain E. pinguis has been found in lowland sites less than 200 m a.s.l. whilst in the Cantabrians, the Pyrenees, and the Alps it occurs in the altitudinal range from 650 to 1650 m. The vicariant pattern is particularly evident in a continuous mountainous area such as the Pyrenees where E. multistiliger appears limited to the south-eastern relief whilst E. pinguis inhabits most of the strictly montane chain. Further, while the former species is found in Sardinia, the latter is found in Corsica.

Geographic variation in segment number
As in most geophilomorphs, in both Eurygeophilus species the number of trunk segments was found to be variable among individuals of the same population. The data collected (see Appendix) show that within each population females usually have two more segments than males. We have also recorded variation in segment number between populations and have been able to recognise some geographical patterns.
In E. pinguis segment number is quite uniform within large areas of the species' range but different between such regions. In North Devon (data on specimens from a dozen localities) all known males have 35 leg-bearing segments while all females have 37. In the Pyrenees (data on 19 males and 17 females from 21 localities) 35 is again the most frequent value for the males and 37 for the females; rare exceptions were one male with 37 segments and one female with 35 segments, but there was also one female from Et Ustarila with 47 segments. The only specimen from the Cantabrian chain is a male with 35 segments, which is consistent with the modal values in Devon and the Pyrenees. In the western and central Alps from the Ligurian and Maritime Alps to the Venetian Pre-Alps (eight males and 19 females from 15 localities), the most frequent values are 41 for the males and 43 for the females; other values are found less frequently, the range for females being from 41 to 47. Within this same region most of the populations are similar in segment numbers but comparatively higher values are typical of some (e.g. Casteldelfino and M. La Marzola). In the eastern Alps (nine males and eight females from seven localities), all known males have 33 leg-bearing segments while all females have 35. The only record from Corsica is for one female with 45 leg-bearing segments. From this, three main regions may be recognised within the geographical range of E. pinguis on the basis of the prevailing number of segments: (1) the Pyrenees and Devon (and most probably also the Cantabrian chain), (2) the western-central Alps (and most probably also Corsica), and (3) the eastern Alps. Whether this geographical pattern in the segment number is consistent with a phylogeographic structure remains to be evaluated.
By comparison with E. pinguis, too few data are available for E. multistiliger to recognize any geographical pattern of variation in the segment number. Worth noticing, however, is that similarly high values were found in the three specimens from the eastern Pyrenees (55-59 leg-bearing segments) whereas different, lower values were found from other regions (49 and 51 in two females from Sardinia and western Iberian Peninsula, respectively).