Clinopodes flavidus C. L. Koch, 1847

Bonato, Lucio, Iorio, Étienne & Minelli, Alessandro, 2011, The centipede genus Clinopodes C. L. Koch, 1847 (Chilopoda, Geophilomorpha, Geophilidae): reassessment of species diversity and distribution, with a new species from the Maritime Alps (France), Zoosystema 33 (2), pp. 175-205 : 187-189

Clinopodes flavidus C. L. Koch, 1847

TYPE MATERIAL AND TYPE LOCALITY. — 1 ♀, holotype ; from “Oravitza” (= Oraviţa [ Romania]) (C. L. Koch 1847).

SYNONYMS. — Arthronomalus hopei Newport, 1845 : n. syn. (see notes below) (type material and type locality: holotype, from near Naples [ Italy]). — Poabius nitens C. L. Koch, 1847 : first synonymized by Latzel (1880) (type material and type locality: unknown number of syntypes, from Carniola [ Slovenia]). — Geophilus montanus Meinert, 1870 : first synonymized by Latzel (1880) (type material and type localities: unknown number of syntypes, from Alpe Lipanza [ Slovenia], Gennazano and Ariccia [both in Italy]). — Geophilus gorizensis Latzel, 1880 : n. syn. (see notes below) (type material and type locality: holotype, fromTarnova massif [ Slovenia]). — Geophilus pannonicus Verhoeff, 1895 : first synonymized by Verhoeff (1898) (type material and type locality: holotype, from Zagreb [ Croatia]). — Geophilus makrodontus Attems, 1907 : n. syn. (see notes below) (type material and type locality: holotype, from Alushta [ Ukraine]).

NOMINAL SUBSPECIES. — Clinopodes flavidus styriacus ( Attems, 1895) ; C. flavidus polytrichus ( Attems, 1903) ; C. flavidus setosus ( Lignau, 1903) ; C. flavidus vestitus ( Lignau, 1903) ; C. flavidus noduliger (Verhoeff, 1925 nec Verhoeff, 1928); C. flavidus noduliger ( Verhoeff, 1928 nec Verhoeff, 1925); C. flavidus apruzianus (Verhoeff, 1934) ; C. flavidus pachypus ( Verhoeff, 1942) ; C. flavidus faitanus (Verhoeff, 1943) ; C. flavidus improvisus (Verhoeff, 1943) ; C. flavidus karamani (Verhoeff, 1943) ; C. flavidus sorattinus ( Verhoeff, 1951) .

MAIN REFERENCES. — Newport 1845: 433 (original description of A. hopei ); C. L. Koch 1847: 182 (original description of P. nitens ), 184 (original description, as Clinopodes flavidus ); C. L. Koch 1863: 105 (redescription); Meinert 1870: 75 (original description of G. montanus ); Latzel 1880: 168 (in key), 175 (redescription); Haase 1881: 75 (in key), 78 (redescription); Berlese 1883: 9 (redescription); Attems 1895: 163 (original description of G. flavidus var. styriaca ); Verhoeff 1895: 350 (original description of G. pannonicus ); Verhoeff 1896: 2 (redescription); Attems 1903: 222 (in key), 233 (original description of G. flavidus polytrichus ); Attems 1907: 8 (original description of G. makrodontus ); Folkmanová 1928: 111 (in key), 112 (redescription); Verhoeff 1928: 271 (in key, and original description of G. flavidus noduliger ); Attems 1929a: 202 (in key), 203 (redescription); Verhoeff 1934a: 10 (original description of G. flavidus apruzianus ); Verhoeff 1934b: 113 (in key); Verhoeff 1942: 505 (original description of G. flavidus pachypus ); Verhoeff 1943a: 82 (original description of G. flavidus faitanus ); Verhoeff 1943b: 146 (original description of G. flavidus improvisus ); Attems 1947: 119 (in key); Verhoeff 1951: 231 (original description of G. flavidus sorattinus ); Folkmanová 1952: 182 (in key); Kanellis 1959: 38 (in key); Dărăbanţu & Matic 1969: 102 (in key); Matic 1972a: 75 (in key); 76 (redescription), 79 (redescription as C. polytrichus ); Kaczmarek 1979: 62 (redescription and in key); Stoev 2002: 89 (in key).

SPECIMENS EXAMINED. — Bulgaria. Rila, Rilski Manastir, 3.VII.1983 K. Marinčeva lg, 1 ơ (50 mm, 69 leg pairs) (coll. MB).

Croatia. Plitvička Jezera, 13.IV.1968 A. Minelli lg, 1 ♀ (49 mm, 65 leg pairs) (coll. MB).

Greece. Geraki, Thrace, 5. VI.1983 G. Etonti lg, 1 ơ (42 mm, 73 leg pairs) (coll. MB). — Kato Nevrokopi, 19.IV.1982 G. Etonti lg, 1 ♀ (58 mm, 73 leg pairs) (coll. MB). — Kavala Lekanis 4. VI.1983 G. Etonti lg, 1 ♀ (68 mm, 79 leg pairs), 1 ơ (58 mm, 67 leg pairs) (coll. MB). — Phalakron, Drama, 18.IV.1982 G. Etonti lg, 1 ♀ (43 mm, 69 leg pairs) (coll. MB).

Italy. Arenzano, near Genova, 22. II.1980 G. Gardini, C. Torti & S. Zoia lg, 1 ơ (28 mm, 61 leg pairs) (coll. MB). — Caprazoppa, near Finale Ligure, 29.X.1982 G. Gardini & R. Rizzerio lg, 1 ♀ (23 mm, 65 leg pairs) (coll. MB). — Casella, near Genova, 11.IV.1979 G. Gardini lg, 1 ♀ (48 mm, 65 leg pairs) (coll. MB). — Magliolo, near Savona, 25.IV.1985 G. Gardini lg, 1 ♀ (40 mm, 65 leg pairs) (coll. MB). — San Giacomo near Roburent, 22. V.1982 G. Gardini & R. Rizzerio lg, 1 ♀ (28 mm, 59 leg pairs) (coll. MB).

Jordan. Zubya, 30.IV.2009 M. Uliana lg, 1 ơ (45 mm, 59 leg pairs) (coll. MB).

DIAGNOSIS. — A Clinopodes species usually up to c. 6 cm long; 51-81 pairs of legs, most often 53-79 in the male, 59-81 in the female; denticles of the forcipular coxosternite from distinctly wider than long to as long as wide; chitin-lines reaching the condyles; even the largest sternal pore-fields on the posterior leg-bearing segments remaining well behind the mid-length of the metasternite; all canals of the coxal organs converging into 2-4 poorly recognizable clusters aligned along the lateral margin of the metasternite and usually covered by it; no isolated pores. See also Table 3 and key to species.

GEOGRAPHIC DISTRIBUTION. — Mainly central and south-eastern Europe, westwards to the Italian side of the Western Alps and most of the Italian Peninsula including coastal Tyrrhenian islands, eastwards to the regions around the Black Sea and to the Aegean islands, northwards inhabiting the whole Alps and Carpathians and some scattered more northern localities, southwards reaching Sicily, Malta, Crete, Cyprus and the Levant. Published records from the Caucasus and Anatolia deserve confirmation, as they were possibly based on a wider concept of the species encompassing C. caucasicus and C. escherichii .

TAXONOMIC AND NOMENCLATURAL NOTES Originally described in the genus Clinopodes by C. L. Koch (1847), C. flavidus was treated by subsequent authors under Geophilus until it was reinstated in the genus Clinopodes by Brölemann (1909). Since Attems (1929a), this generic combination has been followed by most subsequent authors, with the notable exceptions of K.W. Verhoeff and B. Folkmanová, who treated Clinopodes as a subgenus of Geophilus (see notes above, under the genus Clinopodes ). The validity of the species was never disputed.

Clinopodes flavidus and its synonym P. nitens were described simultaneously in the same publication (C. L. Koch 1847), but the former is the valid name for the species since Latzel (1880), who acted as “First Reviser” ( ICZN 1999: art. 24.2) when synonymizing P. nitens under C. flavidus .

The nomenclature of the putative subspecies is confused by the fact that the name noduliger was introduced twice by K.W. Verhoeff for two actually distinct subspecies, based on specimens from Israel ( Verhoeff 1925a) and from Italy ( Verhoeff 1928) respectively.

NOTES ON NEW SYNONYMIES

Arthronomalus hopei was described by Newport (1845) based on a single male collected “prope Neapolin” (= near Naples [ Italy]). As the original description was very vague, the true identity of this nominal species remained unknown, as explicitly acknowledged by Attems (1929a) and Manfredi (1956), and no other specimens were referred to it. Arthronomalus hopei was reported as a putatively valid species only in some old publications, either under Arthronomalus Newport, 1845 (currently a synonym of Geophilus ) or Geophilus ( Gervais 1847; Newport 1856; Fanzago 1874), whereas it was almost completely ignored by subsequent authors. It is worth noting that Meinert (1870), when describing the new species G. montanus (a synonym of C. flavidus ; see above), commented on the fact that it could be identical to A. hopei . Among the very scarce morphological information provided in the original description ( Newport 1845), “labio [...] minute bidentato” may be interpreted most probably as indicating the presence of a pair of denticles on the anterior margin of the forcipular coxosternite. This character, together with the number of trunk segments and the provenance of the holotype, allows the confident recognition of A. hopei as identical to C. flavidus , which is relatively common in the area (e.g., Verhoeff1928, 1943a; Zapparoli & Minelli 2005). Even though A. hopei is recognized as a senior synonym of C. flavidus , we think that resurrecting the former name would not serve nomenclatural stability and universality and therefore we advocate the conservation of Clinopodes flavidus C. L. Koch, 1847 over Arthronomalus hopei Newport, 1845 ( ICZN1999: Art. 23.9): at best of our knowledge, the name A. hopei was not used as valid after Fanzago (1874), even though it continued to be cited as a species of uncertain validity up to Manfredi (1956); conversely, C. flavidus has been universally and repeatedly used as the valid name for this species since the end of the 19th century and, in particular, in more than 25 publications by more than 10 authors in the last 50 years and encompassing a span of more than 10 years (e.g., Dărăbanţu & Matic 1969; Negrea et al. 1970; Matic 1972a, b; Würmli 1972; Kaczmarek 1979, 1980; Minelli 1982, 1992, 1993; Minelli & Zapparoli 1985, 1992, 1994; Kos 1992; Foddai et al. 1995; Ribarov 1996; Stoev 1997, 2001a, 2002, 2004; Wytwer 1997; Zapparoli 1999, 2002; Stašiov 2001; Bonato et al. 2005, 2008).

Geophilus gorizensis was described by Latzel (1880), based on a single female from the “südöstlichste Theil desTarnowaner-Gebirges” (= extreme south-east part of Tarnova massif [ Slovenia]).It was repeatedly cited as a valid taxon, most often at the rank of species, and it was also reported from another locality, close to the type locality ( Manfredi 1940). Since Attems (1929a), it has been cited invariantly in the genus Pleurogeophilus , according to its putative affinity to P. mediterraneus ( Meinert, 1870) . This taxonomic position was first questioned by Minelli (1992), who suspected instead that G. gorizensis could actually represent a Clinopodes species, and uncertainty as to its true identity has been repeatedly expressed by modern authors ( Foddai et al. 1995; Stoev 1997). Based on the original description ( Latzel 1880), the traditional assignment of G. gorizensis to Pleurogeophilus appears unwarranted, as G. gorizensis differs from P. mediterraneus in characters that are regarded as diagnostic for Pleurogeophilus , such as a toothless anterior margin of the forcipular coxosternite and sub-circular sternal pore-fields. Geophilus gorizensis is consistent with Clinopodes in a combination of characters diagnostic for the latter genus in respect to all other western Palaearctic geophilids, including the presence of denticles and chitin-lines on the forcipular coxosternite, the arrangement of sternal pores into posterior transversal bands on most trunk segments and larger areas on the most posterior segments, and the lack of claws on the ultimate legs. Inconsistently, however, the metasternite of the ultimate leg-bearing segment was described as relatively narrow, rather than wider than long as typical for Clinopodes , and the coxal pores were indicated as distributed on the entire surface of the coxopleura, whereas they are usually concentrated towards the metasternite in Clinopodes . Nevertheless, among the known species of Clinopodes , information provided by Latzel (1880) on body size, forcipular denticles, chitinlines, and number of trunk segments are together suggestive of C. flavidus , which is known to occur widely in the area.

Geophilus makrodontus was described by Attems (1907) based on a single female from “Aluschta, Krim ” (= Alushta, in Crimea [ Ukraine]). No other specimens were referred to this species, which was only cited in major faunistic revisionary works, even though with the incorrect subsequent spelling G. macrodontus ( Attems 1927, 1929a, 1947; Titova 1969). Attems (1907) assigned G. makrodontus to the subgenus Geophilus , and considered it most similar to Geophilus strictus Latzel, 1880 (a species of uncertain identity, possibly a Clinopodes species; see below under Discussion) and secondarily to Geophilus gracilis Meinert, 1870 (a true Geophilus species; Eason 1961) and Geophilus latro Meinert, 1870 (a synonym of Arenophilus bipuncticeps (Wood, 1862) ; Chamberlin 1912b). Subsequently, Attems (1927, 1929a, 1947) confirmed G. makrodontus in the genus Geophilus and included it in keys where it was contrasted directly with G. electricus (Linnaeus, 1758) . The taxonomic position of G. makrodontus was never reassessed. Based on the quite detailed original description and illustrations ( Attems 1907), and contrary to the taxonomic position maintained by the same C. Attems, the holotype of G. makrodontus is unambiguously recognizable as belonging to the genus Clinopodes , as it shares all major diagnostic characters of this genus, including the overall features of maxillary complex, forcipular segment (general shape, and denticles on the anterior margin of the coxosternite), trunk metasternites (carpophagus socket and shape of the pore-fields), and ultimate leg-bearing segment (shape of metasternite, pattern of coxal pores, lack of claws). In particular, the presence of complete chitin-lines, the size of the denticles on the forcipular coxosternite, the number of leg pairs, the short extent of the porefields on the most posterior metasternites, and the arrangement of the coxal pores into clusters are fully and exclusively diagnostic of C. flavidus . It is worth noting that C. flavidus has been already recorded from Crimea ( Selivanov 1881, 1884; Attems 1907).