Eurygeophilus multistiliger ( Verhoeff, 1899 )

Bonato, Lucio, Barber, Anthony & Minelli, Alessandro, 2006, The European centipedes hitherto referred to Eurygeophilus, Mesogeophilus, and Chalandea (Chilopoda, Geophilomorpha): taxonomy, distribution, and geographical variation in segment number, Journal of Natural History 40 (7 - 8), pp. 415-438 : 419-431

Eurygeophilus multistiliger ( Verhoeff, 1899)

( Figures 9–13 )

Geophilus (Eurygeophilus) multistiliger Verhoeff, 1899 .

5 Eurygeophilus multistyliger [sic] velmanyensis Brolemann, 1926 (n. syn.).

Geophilus (Eurygeophilus) multistiliger Verhoeff 1899, p 367 (original description); 1902–25, p 581, 656; 1928, p 267; Attems 1903, p 170, 241.

Eurygeophilus multistiliger: Attems 1929, p 212 (diagnosis).

Eurygeophilus multistyliger [sic]: Brolemann 1926a, p 239, 260; Machado 1952, p 84 (diagnosis), Figure 9 (1–6); Serra and Ascaso 1990, p 391 (n. loc.).

Eurygeophilus multistiligeri [sic]: Attems 1952b, p 325.

Eurygeophilus multistyliger [sic] velmanyensis Brolemann 1926a, p 235, 237, 260 (original description), Figures 28–34; 1930, p 52 (in key), 187 (diagnosis), Figures 305–311; Demange 1981, p 222 (in key, as Eurygeophilus multistylifer [sic] velmanyensis), 233; Geoffroy 2000, p 162.

Eurygeophilus multistiliger velmanyensis: Attems 1929, p 212 (diagnosis); Machado 1953, p 82 (n. loc.).

Eurygeophilus multistyliger [sic] var. velmanyensis: Machado 1952, p 86 .

Holotype

Female, 32 mm long, 51 leg-bearing segments; L. Vieira leg.; in the collections of the Museum für Naturkunde, Humboldt-Universität, Berlin, ZMB 13504. Body separated into six parts (1–6), mounted on two slides (labelled ‘‘a’’ and ‘‘b’’): (1) head, forcipular segment, leg-bearing segment I; (2) maxillary complex and left mandible; (3) right mandible; (4) leg-bearing segments II–XXV; (5) leg-bearing segments XXVI– XLVI; (6) terminal part of trunk including leg-bearing segments XLVII –LI. Parts 1–3 and 6 are included in the slide labelled ‘‘a’’, parts 4 and 5 in the slide labelled ‘‘b’’.

Type locality

‘‘Coimbra’’ ( Verhoeff 1899; also on the label).

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

(Brölemann, 1898) (see also Figures 9–18 ).

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. Brölemann 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 (1926a, 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 (1926a, 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).

The newly identified specimen from Sardinia fully agrees in morphology with the holotype of the species as well as with the specimens from the eastern Pyrenees as described and illustrated in detail by Brolemann (1926a, 1930).

Eurygeophilus pinguis (Brölemann, 1898)

( Figures 14–18 )

Geophilus pinguis Brölemann, 1898 .

5 Geophilus (Mesogeophilus) baldensis Verhoeff, 1901 (n. syn.).

5 Chalandea cottiana Verhoeff, 1938 .

5 Chalandea cottiana var. castrensis Manfredi, 1948 .

5 Chalandea scheerpeltzi Attems, 1952 (n. syn.).

Geophilus pinguis Brölemann 1898a, p 46 (original description), Figures 29–32; 1898b, p 189, 201 (n. loc.); 1909b, p 211 (in key); Léger and Duboscq 1903, p 313 (n. loc.), 354.

Geophilus (Eurygeophilus) pinguis: Attems 1903, p 170 , 241.

Chalandea pinguis: Brölemann 1909a, p 330 , 338; 1926a, p 262; 1926b, p 233, 236 (also as Geophilus pinguis on p 236); 1930, p 36, 53 (in key), 190, 191 (diagnosis), Figures 11–12 , 312–318; Verhoeff 1902 –25, p 657; 1938, p 351, 352, 353 (diagnosis); 1943, p 5, 19, 20; Attems 1929, p 211 (diagnosis), Figures 185; 1947, p 127 (in key); 1952a, p 50; Manfredi 1948, p 208; Barber 1972, 46; 1985a, p 18, 21; 1985b, p 35 (diagnosis), Figures 3–5 ; 1992a, p 348; 1992b, p 31, 33, 38 (n. loc.); 2003, p 69, 70; Blower 1972, p 1, 3 (n. loc.); Würmli 1972, p 4; Barber and Fairhrust 1974, p 614; Minelli 1978, p 158; Barace and Herrera 1980, p 4, 11 (diagnosis; n. loc.), 14, Figures 60–63; 1982, p 119; Demange 1981, p 222 (in key), 233; Minelli and Zapparoli 1985, p 380 (n. loc.), 404, 406, 409; 1992, p 213, 214, 219, 225 (n. loc.); Minelli and Iovane 1987, p 10, 24 (n. loc.); Barber and Keay 1988, p V, 5, 61 (n. loc.), 108–120, Map 25; Keay 1989, p 10; 1993, p 38; Salinas 1990, p 3, 64 (n. loc.), Map 33; Zapparoli 1994, p 103 (n. loc.); Foddai et al. 1995, p 10, 31; Foddai et al. 1996, p 361, 363; Jones and Barber 1997, p 22 (diagnosis), Figures 1–12 ; Geoffroy 2000, p 162; Mann et al. 2003, p 43.

Geophilus (Chalandea) pinguis: Verhoeff 1902 –25, p 581, 655.

Geophilus (Mesogeophilus) baldensis Verhoeff 1901b, p 682 (original description), Figures 1– 4 ; 1902–25, p 304, 580, 623; 1928, p 266; Attems 1903, p 170, 240.

Mesogeophilus baldensis: Attems 1929, p 194 (in key), 195 (diagnosis); 1947, p 63, 124; 1949, p 108; Minelli 1978, p 158; 1981, p 87 (validity doubted); Foddai et al. 1995, p 10, 31 (validity doubted).

Chalandea cottiana Verhoeff 1938, p 353 (original description), 354; Attems 1947, p 127 (in key); Würmli 1972, p 4; Minelli 1978, p 158; 1985, p 32; Minelli and Zapparoli 1985, p 380 (validity doubted); Barber 1992b, p 31, 35 (validity doubted).

Chalandea cottiana var. castrensis Manfredi 1948, p 208 (original description), 223; 1976, p 229; Scossiroli 1951, p 36; Minelli 1985, p 32; Barber 1992b, p 31, 35 (validity doubted).

Chalandea scheerpeltzi Attems 1952a, p 50 (original description), Figures 1–4 ; Strouhal 1961, p 34; Würmli 1972, p 4; Koren 1986, p 36 (diagnosis; n. loc.), Figure 13 ; Kos 1992a, p 354 (n. loc.); 1992b, p 138; 1996, p 638, 639 (n. loc.); Stoev 1997, p 103.

Syntypes

Five specimens: two males, adult, 35 leg-bearing segments; two females, adult, 37 legbearing segments; one female, 8 mm long, 47 leg-bearing segments; all in the collections of the Muséum national d’Histoire naturelle, Paris.

Type localities

Indicated generically as ‘‘Ahusquy’’ in the original description ( Brölemann 1898a), more precisely as ‘‘Bois d’Ithe´’’, ‘‘Naboleguy’’, and ‘‘Et Ustarila’’ by Brölemann (1898b).

Diagnosis

See Table I, Figures 1–3 and Figures 14–18 .

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 Brölemann, 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 are indicated by a single symbol. Contour lines of 500 m are drawn in maps b and c.

illustrated as completely absent in G. (M.) baldensis ( Verhoeff 1901b) but invariantly present and complete in G. pinguis both from the Pyrenees ( Brölemann 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 für 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 ( Minelli and Zapparoli 1985, 1992; Barber 1992b; Foddai et al. 1995). Chalandea cottiana Verhoeff, 1938 is here confirmed as a synonym of Geophilus pinguis Brölemann, 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 Brölemann (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 Brölemann, 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 1992a, 1996). Chalandea scheerpeltzi Attems, 1952 is here recognised as a synonym of Geophilus pinguis Brölemann, 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 ( Brölemann 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.

Distribution (see Appendix and Figure 19 )

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 ( Minelli and 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).

Figures

Figure 19. Geographical occurrence of Eurygeophilus multistiliger (Verhoeff, 1899) (white symbols) and Eurygeophilus Europe generally; (b) the Pyrenees; and (c) the Alps. Type localities are indicated with diamonds, other published Localities of the specimens directly examined by us are indicated with dotted symbols. All published records of E.

Figure 20. Elongation of antennal article XIV in representative specimens of E. pinguis (Brölemann, 1898). The ratio between the length of article XIV and the total length of the previous two articles is plotted against head width as a measure of the body size. A black circle marks the only specimen putativily representative of the nominal species Chalandea scheerpeltzi.

Figures 9–13. Eurygeophilus multistiliger (Verhoeff, 1899), female, 24 mm long, Campu Omo (Sardinia). (9) Head, dorsal (right antenna and setae not drawn). (10) Head, ventral (antennae not drawn). (11) Forcipular segment, ventral. (12) Sternum of leg-bearing segment V, ventral. (13) Last leg-bearing segment and terminal segments, ventral (right telopodite and setae on the relevant coxopleuron not drawn).

Figures 5–8. Eurygeophilus multistiliger (Verhoeff, 1899), female, 32 mm long, Coimbra (Iberian Peninsula), holotype, mounted on a slide. (5) Forcipular segment, ventral. (6) Calyx inside the left forcipule, ventral. (7) Sternum of leg-bearing segment XII. (8) Last leg-bearing segment and terminal segments, ventral. Images from light microscopy. Scale bars: 200 Mm (5); 100 Mm (6, 8); 50 Mm (7).

Figures 14–18. Eurygeophilus pinguis (Brölemann, 1898), female, 27 mm long, M. La Marzola (Alps). (14) Head, dorsal (right antenna and setae not drawn). (15) Head, ventral (antennae not drawn). (16) Forcipular segment, ventral. (17) Sternum of leg-bearing segment V, ventral. (18) Last leg-bearing segment and terminal segments, ventral (right telopodite and setae on the relevant coxopleuron not drawn).

Figures 1–4. (1–3) Eurygeophilus pinguis (Brölemann, 1898), female, 33 mm long, M. La Marzola (near Trento, Alps). (1) Distal articles of the left antenna, ventral. (2) Left claw of the second maxillae, ventral. (3) Right part of the forcipular coxosternum, ventral. (4) Geophilus linearis C. L. Koch, 1835, male, 34 mm long, Cologna Veneta (near Verona, Italy): right part of the forcipular coxosternum, ventral. Arrows indicate the putative position of the forcipular chitin-lines. Micrographs from ESEM Philips XL30. Scale bars: 20 Mm (1, 2); 50 Mm (3, 4).