Dicellophilus carniolensis, (C. L. KOCH, 1847)

Bonato, Lucio, Dányi, László & Minelli, Alessandro, 2010, Morphology and phylogeny of Dicellophilus, a centipede genus with a highly disjunct distribution (Chilopoda: Mecistocephalidae), Zoological Journal of the Linnean Society 158 (3), pp. 501-532 : 520-522

DICELLOPHILUS CARNIOLENSIS (C.L. KOCH, 1847)

Clinopodes carniolensis C.L. Koch, 1847 . Original description: Koch, 1847: 185. Type locality: ‘Herzogthum Krain’ [= Carniola ( Slovenia)]. Type material: number, sex, stage, repository unknown; possibly lost.

= Mecistocephalus hungaricus Tömösváry, 1880 . Original description: Tömösváry, 1880a: 619. Type locality: ‘Hungaria orientalis’ (see notes below). Type material: number of specimens unknown, including both sexes and different developmental stages; originally deposited in the collections of the Transylvanian Museum Society, Cluj-Napoca, Romania, but the current repository is unknown; probably lost. Synonymy by Daday, 1889: 90.

= Geophilus austriacus Meinert, 1886 (nec Geophilus longicornis austriacus Latzel, 1880 ). Original description: Meinert, 1886a: 145. Type locality: ‘Razzes’ [= Bagni di Razzes ( Italy)]. Type material: holotype female, 16.5-mm long, in the Zoological Museum, University of Copenhagen (no code number). Synonymy by Verhoeff, 1901: 459.

= Geophilus apfelbecki Verhoeff, 1898 . Original description: Verhoeff, 1898: 348. Type locality: ‘Sarajevo, [...] Plivathal bei Jaice’ [= Sarajevo and Pliva Valley near Jajce ( Bosnia-Herzegovina)]. Type material: number of specimens unknown, all female, repository unknown; probably lost. Synonymy by Verhoeff, 1901: 459.

= Geophilus apfelbecki diversiporus Verhoeff, 1898 . Original description: Verhoeff, 1898: 348. Type locality: ‘Plasa bei Jablaniza’ [= Plasa near Jablanica ( Bosnia-Herzegovina)]. Type material: holotype female, 20-mm long; repository unknown; probably lost. Synonymy by Verhoeff, 1901: 459.

Main references for morphology: Koch, 1847: 185 (original description); Meinert, 1870: 94 (redescription); Latzel, 1880: 162 (redescription); Tömösváry, 1880a: 619 (redescription as M. hungaricus ); Meinert, 1886a: 145 (redescription as G. austriacus ); Daday, 1889: 90 (redescription); Verhoeff, 1898: 348 (redescription as G. apfelbecki and G. apfelbecki diversiporus ); Verhoeff, 1902 –1925: 191 (redescription); Attems, 1929: 147 (diagnosis and key); Eason, 1964: 43 (key), 60 (diagnosis); Matic, 1972: 66 (redescription); Kaczmarek, 1979: 79 (redescription); Koren, 1986: 67 (redescription); Mikoš, 1991: 13 (redescription).

Diagnosis: A Dicellophilus species with 43 leg-bearing segments. Antennae about three times as long as the maximum width of the head, slightly tapering (width of the terminal article about half of the width of article I); terminal article less than two times as long as wide, and more than 1.5 times as long as article XIII. The longest setae on the antennae are shorter than 200 Mm ( Fig. 3E ). Sensilla on the antennal tip ( Fig. 8A ) without any evident projection at roughly mid-length. Head ( Fig. 3A ) ~1.2–1.4 times as long as wide. Lateral margins of the cephalic plate markedly convex, only evidently converging in the posterior part. Frontal line ( Fig. 3C ) uniformly rounded. Clypeus ( Figs 3D , 8B ): lateral margins uniformly curved, only slightly converging backwards; setae extending backwards, only lacking on a narrow band along the posterior margin, without a distinctly isolated posteromedian pair of setae. Non-areolate part of the buccae ( Fig. 3D ) not or only slightly extending forwards beyond the labrum. Mandible with between three and five pectinate lamellae. Coxosternum of maxillae I ( Figs 3B , 9A ) more than 3.3 times as wide as medially long. Medial projection of maxillae I about as long as wide; distal hyaline part only slightly enlarged, not subtriangular; no hyaline scales on the dorsal surface ( Fig. 3H ). Distal hyaline part of the telopodite of maxillae I about as long as the basal chitinous part. Distal article of the telopodite of maxillae II subcylindrical for most of the length, tapering abruptly in the distal part, usually more than two times as long as wide ( Fig. 3B ). Condylar processes of the forcipular coxosternum ( Fig. 3I ) virtually absent. Forcipular article I 1.2–1.3 times as long as wide, the tubercle relatively small, not or only slightly projecting beyond the distal margin of the article ( Figs 3F , 9B ). Forcipular tarsungulum virtually without basal tubercle ( Fig. 9I ), the dorso-internal margin only weakly crenulated, with uniformly spaced notches ( Fig 9G, H ). Sternum of the last leg-bearing segment ( Figs 3G , 9C ) longer than wide. Posterior end of coxopleura rounded.

The main differential characters in respect to the other Dicellophilus species are presented in Table 3.

Notes on taxonomy: Originally included in the genus Clinopodes C.L. Koch, 1847 ( Koch, 1847) , the species was assigned to the genus Mecistocephalus Newport, 1843 by Meinert (1870), followed by some subsequent authors, whereas Cook (1896a) included it in the new genus Lamnonyx Cook, 1896 , currently a synonym of Mecistocephalus ( Bonato et al., 2003) . C. carniolensis was moved to Dicellophilus by Silvestri (1919), and, even though it was still occasionally cited under Mecistocephalus , since the 1960s this generic assignment has been followed universally.

The type material of C. carniolensis is apparently not present in the Natural History Museum, London (A. Minelli, pers. observ.), which is the main repository of C.L. Koch’s myriapods, nor is it to be found in the Museum für Naturkunde, Berlin ( Moritz & Fischer, 1979), where other material from Koch’s collection is preserved .

Mecistocephalus hungaricus Tömösváry, 1880 was described by Tömösváry (1880a) from a series of specimens, up to 22-mm long, including males with 41 pairs of legs and females with 43 pairs. The exact provenance of the original material is unknown, as Tömösváry (1880a) referred generically to ‘Hungaria orientalis’, i.e. the eastern part of the Kingdom of Hungary, of that time, which is a vaguely defined region that is currently included, for the most part, in Romania (Transylvania), but is also partially included in Slovakia, Hungary, and Ukraine. No other specimens were referred to M. hungaricus . After examining the type specimens, Daday (1889) recognized them as belonging to D. carniolensis , and therefore synonymized M. hungaricus under C. carniolensis . The synonymy was not questioned by any subsequent author. As for the repository of the type material, Ö. Tömösváry deposited all the specimens in the collections of the Transylvanian Museum Society (Erdélyi Múzeum-Egylet) in Cluj Napoca (formerly, Kolozsvár), Romania ( Tömösváry, 1880b), where they were later found by Daday (1889). At present, no specimens referable to the type material are apparently preserved in the collections of the Transylvanian Museum Society, in the Babeş-Bolyai University, Cluj- Napoca (V. Ilie & B. Markò, pers. comm.), nor in those of the Institute of Biological Research of the Romanian Academy (V.V. Pop, pers. comm.), nor even in the Hungarian Natural History Museum, Budapest, where a small part of Tömösváry’s heritage can be found. Therefore, as we were unable to directly check the identity of M. hungaricus , we rely on Daday’s (1889) proposal of synonymy, even though the number of 41 leg-bearing segments given for the males of M. hungaricus is inconsistent with D. carniolensis (see below).

Geophilus austriacus Meinert, 1886 was described by Meinert (1886a) on the basis of a single specimen. No other specimens have been referred to this nominal species, which was only listed as valid by Attems (1903). Verhoeff (1898) suspected that G. austriacus could be identical to his species Geophilus apfelbecki Verhoeff, 1898 (see below), and he later synonymized both under D. carniolensis ( Verhoeff, 1901) , which was followed by all subsequent authors. Based on the original description (cephalic plate elongation, pattern of forcipular tubercles, 43 pairs of legs, mid-longitudinal sternal sulcus, sternum of the last leg-bearing segment relatively narrow, and a single pore on each coxopleuron), G. austriacus is only fully compatible with a juvenile D. carniolensis .

Geophilus apfelbecki Verhoeff, 1898 and Geophilus apfelbecki diversiporus Verhoeff, 1898 were both described by Verhoeff (1898), and no other specimens were referred to either taxon. Both taxa were recognized as synonyms of D. carniolensis by the same author ( Verhoeff, 1901), followed by all subsequent authors. The type specimens are apparently not present in the Museum für Naturkunde, Berlin ( Moritz & Fischer, 1979), where the type specimens of all other taxa described by Verhoeff in the same paper are preserved; neither are they present in the Zoologische Staatssammlung München (S. Friedrich, pers. comm.). Based on the original description (features of the maxillary complex, pattern of forcipular tubercles, 43 pairs of legs, mid-longitudinal sternal sulcus, lack of sternal pores, sternum of the last leg-bearing segment relatively narrow, macropores on coxopleura, and no claw on the legs of the last pair), the type specimens of both G. apfelbecki and G. apfelbecki diversiporus are obviously recognizable as juveniles of D. carniolensis . In particular, G. apfelbecki was described from specimens of up to 19.5-mm long, with a single pore on each coxopleuron, whereas G. apfelbecki diversiporus was described from specimens with one macropore, and a dozen small pores on each coxopleuron.

Notes on morphology: Based on extensive sampling, D. carniolensis was found to be invariant in the segment number throughout its geographical range, with both males and females having 43 leg-bearing segments. The only published record apparently challenging this evidence is that by Tömösváry (1880a), who described the type series of M. hungaricus as including females with 43 pairs and males with 41 pairs of legs (see above). However, it is possible that Tömösváry (1880a) was inaccurate in describing the segment number of those specimens, or that these included specimens of other species misidentified as D. carniolensis (it is worth noting that Pachymerium ferrugineum C.L. Koch, 1835 , occasionally misidentified with D. carniolensis in the 19 th century, is a species that also occurs in that area, with the number of leg-bearing segments ranging between 41 and 57; Matic, 1972), but this cannot be ascertained now, as Tömösváry’s specimens are apparently lost (see above). Also, Takakuwa (1938a, b) assigned 41 legbearing segments to D. carniolensis , but this was based on an obviously misidentified specimen of D. pulcher from Honshu.

The terminal antennal article was sometimes described as spoon-shaped ( Latzel, 1880; Attems, 1929), but this is recognizable as an artefact occasionally occurring in preserved specimens, caused by the collapse of the integument.

Distribution: The species occurs in a restricted area in central Europe, comprising some major mountainous regions and surrounding lands, namely the centraleastern Alps, the Dinarids, the West, East and South Carpathians, and the Dobrogea ( Fig. 10 ). It has been recorded from hundreds of localities.

Moreover, introduced specimens have been occasionally recorded from a few localities in Great Britain, either in greenhouses or in other synanthropic sites ( Bagnall, 1913a, b; Fig. 10 ).

Notes on distribution: Dicellophilus carniolensis has been cited repeatedly as also occurring, at least as an introduced species, in North America and in Honshu ( Attems, 1929; Takakuwa, 1934a, b, 1938a, b; Chamberlin & Wang, 1952; Eason, 1964; Matic 1972; Kevan, 1983; Barber, 1985; Voigtländer, Spelda & Zulka, 1994). However, no evidence has been published indicating that specimens of D. carniolensis have ever been collected in North America and Japan, and those reports most probably refer to other species of Dicellophilus .

Figures

Figure 3. Dicellophilus carniolensis. A, head, dorsal view. B, right part of the maxillary complex, ventral view (detached from head). C, anterior part of head, dorsal view. D, clypeus and labrum, ventral view (maxillae removed). E, left antennal articles III–V, dorsal view. F, internal margin of left forcipule, dorsal view. G, last leg-bearing segment of an adult female, ventral view. H, left telopodite and medial projection of maxillae I, dorsal view (detached from head). I, left condylar process of forcipular coxosternum, dorsal view (head removed). Microscopic photographs and line drawings: ♀, 50-mm long, from Val di Nos, Italy; the detailed data for the specimen are presented in the Material and methods.

Figure 8. Dicellophilus carniolensis (A–C) and Dicellophilus pulcher (D–F). A, D, apical sensillum on antennal article XIV. B, E, clypeus and labrum, ventral view (maxillae removed). C, F, medial part of labrum, ventral view (maxillae removed). Arrowheads: a, antennal apical sensillum; b, lateral margin of clypeus; c, anterior ala; d, thickened line between anterior and posterior ala; e, posterior ala; f, fold on posterior ala; g, mid-piece of labrum; h, marginal hair-like projections. SEM micrographs: A, D. carniolensis, ♀, 51-mm long, from Mara, Romania; B–C, D. carniolensis, ♀, 55-mm long, from Volovec, Ukraine; D–F, D. pulcher, ♀, 38-mm long, from Shiobara, Japan; the detailed data for the specimens are presented in the Material and methods.

Figure 9. Dicellophilus carniolensis (A–C, G–I) and Dicellophilus pulcher (D–F, J–L). A, D, maxillary complex, ventral view (detached from head). B, E, forcipules and anterior part of coxosternum, dorsal view (head removed). C, F, last leg-bearing segment and terminal segments, female, ventral view. G, J, forcipular tarsungulum, dorsal view. H, K, margin of forcipular tarsungulum, dorsal view. I, L, basal part of forcipular tarsungulum with basal tubercle, dorsal view. Arrowheads: a, telopodite of maxillae II; b, telopodite of maxillae I; c, medial projection of maxillae I; d, metameric pore of maxillae II; e, foraminal process of maxillae II; f, condylar process of forcipular coxosternum; g, cerrus; h, sternum of last leg-bearing segment; i, macropore; j, basal tubercle on forcipular tarsungulum. SEM micrographs: A, D. carniolensis, ♀, 48-mm long, from Valdobbiadene, Italy; B, D. carniolensis, ♀, 55-mm long, from Volovec, Ukraine; C, D. carniolensis, ♀, 51-mm long, from Mara, Romania; D–F, J–L, D. pulcher, ♀, 38-mm long, from Shiobara, Japan; G–I, D. carniolensis, ♀, 55-mm long, from Sighetu Marmaţiei, Romania; the detailed data for the specimens are presented in the Material and methods.

Figure 10. Geographical distribution and phylogenetic relationships of the species of Dicellophilus. The shaded areas indicate the established populations; stars indicate either occasional or dubious records. The dotted line indicates the 40 °N parallel of latitude.