Megaphyllum austriacum (C. L. Koch, 1838)

Lazányi, Eszter & Vagalinski, Boyan, 2013, Redefinition of the millipede subgenus Megaphyllum sensu stricto Verhoeff, 1894 and neotype designation for Megaphyllum austriacum (Latzel, 1884) (Myriapoda: Diplopoda: Julida: Julidae), Zootaxa 3741 (1), pp. 55-100 : 68-70

Megaphyllum austriacum (C. L. Koch, 1838)

Figs 6a–q

Iulus fasciatus C. L. Koch, 1838 : unnumbered pages, Fig. 22.8.

Iulus austriacus: Latzel 1884: 296 –300, Figs 157–159.

Iulus (Chromatoiulus) austriacus: Verhoeff 1894a: 153 ; 1894b: 24. Iulus fasciatus: Attems 1895: 225 –226, Fig. 99.

Megaphyllum (Chromatoiulus) austriacus: Verhoeff 1895: 240 .

Brachyiulus austriacus: Verhoeff 1896a: 219 , Fig. 29; 1897b: 112–113; 1901b: Fig. 6; 1929: 616–617, Figs 14–15 . Brachyiulus (Chromatoiulus) austriacus: Verhoeff 1899a: 194 .

Brachyiulus (Chromatoiulus) dahli Verhoeff, 1901b: 231 –232, Fig. 7 . Chromatoiulus austriacus dahli: Attems 1927: 238 .

Chromatoiulus austriacus: Attems 1927: 237 –238, Figs 323–324; 1929: 330. Chromatoiulus dahlii: Attems 1929: 331 .

Chromatoiulus (Chromatoiulus) austriacus: Attems 1940: 306 , Fig. 27; 1959: 305. Chromatoiulus (Chromatoiulus) dahlii: Attems 1959: 305 .

Megaphyllum dahli: Enghoff & Kime 2009 .

Megaphyllum austriacum: Enghoff & Kime 2009 , Lazányi et al. 2012: 6–8, 18, 23, 41, Figs 1–8 .

Material examined. Bosnia and Hercegovina: Inv. Nr. 8140, 6♂ Preng Waldplateau, [Hercegovačko-neretvanski kanton, Prenj planina [mountains], forest plateau], leg., don. and det. Attems (NHMW); Croatia: Inv. Nr. 8144, 8♂, 20♀ and juveniles, Zengger Hain [=bosk], [Ličko-senjska županija, Senj (Zengg)], 1919. X.07., don. Dobiasch F., det. Attems (NHMW); Inv. No. 8145, 4♂, 16♀ and juveniles, Umbgebung von [environs of] Zengg, [Ličkosenjska županija, Senj (Zengg)], 1919. X.07., don. Dobiasch F., det. Attems (NHMW); Inv. Nr. 8146, 2♂, 12♀ and juveniles, Mendaričeva [probably Mundaricevac] bei Zengg, [Ličko-senjska županija, Senj (Zengg)], 3.XI.1907, don. Dobiasch F., det. Attems (NHMW); Inv. Nr. 8141, 1♂, 1♀ Jurdani, [Primorsko-Goranska zupanija, Jurdani], leg., don. and det. Attems (NHMW); Inv. Nr. 8148, 1♂, Birbital bei novi Vinodol, [Primorsko-goranska županija, Novi Vinodolski municipality], 1919. X.07, don. Dobiasch, det. Attems (NHMW); 1♂, Fiume [Rijeka], Tersato [Trsat hill], leg., don. and det. Attems (NHMW); Slovenia: Inv. No. 8138, 1♂, many ♀, Lipizzaner Wald, [Sezana municipality, Sezana city, forest of Lipica], leg., don. and det. Attems (NHMW).

Distribution. Bosnia and Hercegovina (Verhoeff 1901b); Croatia (Attems 1929; Strasser 1965; Mršić 1987); Italy: Tre Venezie, North Italy (Foddai et al. 1995); Montenegro (Attems 1929; Makarov et al. 2004); Republic of Macedonia (Makarov et al. 2004); Serbia: Kosmaj Mt., Central Serbia, Pećina in Barskom Ridu, Divna Gorica, Suva Planina Mt., Deliblatska Peščara, Fruška Gora (Makarov et al. 2004); Slovenia (Strasser 1966a).

Diagnosis. Differs from all consubgeners by having the simplest opisthomere, i.e. with short processes without any special forms, enlargements or alterations. Distinguishable from the closest species M. carniolense by the very short posterior process (pp) just reaching the height of the small posterior hump (ph) (Figs 6b, 6d, 6o–q).

Opisthomere’s (Figs 6a–b, 6d–q) posterior process (pp) shorter than solenomere; solenomere very small, mostly obscure, the size and orientation of the solenomere’s posterior process (psp) variable, usually longer than the anterior process (asp) (except for individuals with rudimentary psp). Some individuals with a rather small posterior apical hump (ph).

Body colour: all examined individuals were quite old, so body colour could have faded, but both sexes seemed to be uniformly dark or brighter brown, metazona sometimes darker, with a longitudinal black dorsal thin or wider line. Body length and height: males: 30.7–33.6mm, 2.1–2.3mm; females: 36.6–37mm, 2.7–2.8mm.

Remarks. Iulus austriacus Latzel, 1884 was a replacement name for Iulus fasciatus C.L. Koch, 1838 (preoccupied by Iulus fasciatus De Geer, 1778 ) (Latzel 1884). The type specimens of I. austriacus are thus Koch’s original specimens of his I. fasciatus (ICZN Article 72.7). The specimens that Latzel referred to as I. austriacus are not types, and it is unnecessary to invoke Article 75.6 to “de-typify” Latzel’s specimens. However, Koch’s types are inaccessible.

After the original description (C. L. Koch 1838) Latzel (1884) redescribed M. austriacum as a highly variable species living in the Austro–Hungarian Empire and gave some very small figures. The specimens that he referred to as I. austriacus represent three species: M. p. projectum , M. silvaticum and M. bosniense . There was confusion on the species identity in the following years, e.g. Attems in his “Die Myriapoden Steiermarks” (=Myriapods of Styria) (1895) gave a figure of the later-described M. silvaticum while discussing Julus fasciatus . Verhoeff (1897b, 1899a, 1901) stated that the form living in the Croatian coastline should be regarded as M. austriacum . Since then this M. austriacum sensu Verhoeff has been used.

FIGURES 6a–q. Megaphyllum austriacum . (6a) Gonopods of the neotype male individual from Fiume, ZMB4377, posterior view; (6b) same individual, right gonopods, mesal view. (6c) Left promere of a M. austriacum male, meso-posterior view, unknown locality, ZMB1769. Neotype’s (6d) right opisthomere, lateral view, penis, (6e) posterior and (6f) left lateral views. (6g) Female from Fiume, ZMB4377b, left vulva, posterior view. Neotype male: (6h) antenna; (6i) gnathochilarium; (6j) telson; (6k) subanal scale; (6l) left first leg, lateral view; (6m) left second leg, posterior view; (6n) left anterior leg, 25th body ring. Left opisthomeres, mesal view: (6o) environs of Senj, Croatia; (6p) Jurdani, Croatia; (6q) Fiume, Croatia. Abbreviations: asp: anterior process of the solenomere, ca: central ampulla, ct: connecting tube; f: femur, fl: flagellum, g: groove, ll: left lamella, op: operculum, pa: posterior ampulla, pf: postfemur, ph: posterior hump; pp: posterior process, psp: posterior process of the solenomere, r: ridge, rl: right lamella, sl: serrated lobe, t: tibia, P: promere. Scale bars: 0.2mm. Setae of antenna, gnathochilarium, walking legs and vulva omitted.

As C. L. Koch’s types cannot be found, we designate a neotype for M. austriacum to stabilize the usage of M. austriacum sensu Verhoeff. The designated neotype is the specimen deposited in the MNB, deriving from Fiume, identified as M. austriacum most probably by Verhoeff himself (according to the label and the fact that he was actively collecting in Croatia; there are other, type specimens in the MNB, collected by him from Fiume—Jason Dunlop, pers comm.). Thus, it represents perfectly M. austriacum sensu Verhoeff. Its label is as follows: “ZMB 4377 1♂, Brachyiulus austriacus Latzel & Verhoeff, Fiume ”.

Description of neotype: Length: 33.6mm, height: 2.3mm, number of body rings: 48+1+T, 10 rows of ocelli, i.e. XI. developmental stadium, number of ocelli: R41+L40. Body colour probably faded during conservation: bright yellowish, metazona slightly darker. Antennal articles and gnathochilarial parts as on Figs 6h and 6i. Metazona densely striated, posterior margin with long setae, openings of repugnatorial glands right on the suture. Anal valves densely setose. Preanal process long, pointed, setose (6j), subanal scale triangular, apically with 4 blunt teeth and 5 setae (Fig. 6k). Mandibles with protruding lobe; first leg-pair forming a simple, slightly angular hook (Fig. 6l); second leg-pair with two simple pads on postfemur (pf) and tibia (t), coxa without modifications (Fig. 6m); penis (Figs 6e–f) stout, antero-posteriorly flattened, apically with two minute lamellae (ll and rl); pleurotergum of the 7th body ring protruding like a simple shovel; walking legs with three simple pads on femur (f), postfemur (pf) and tibia (t) (Fig. 6n). Promere (P) (Figs 6a–c): higher than opisthomere, lateral and mesal margins parallel; from mesoposterior view triangular, i.e. apically tapering; on the posterior surface with an oblique ridge (r). Opisthomere (6a– b, 6d–q): posterior process (pp) pointed, shorter than solenomere; no anterior process or ridge. Solenomere: posterior process (psp) triangular, hardly discernible, anterior process (asp) longer than the posterior one. Opisthomere located into a groove (g) formed by the promere’s ridge (r).

The M. austriacum specimens (from now on used as sensu Verhoeff ) investigated in the present study depict a highly variable species: the posterior solenomere process (psp) can vary from rudimentary (Fig. 6d) to welldeveloped (Figs 6o, 6q), large (Fig. 6p); the opisthomere’s posterior process (pp) can be very tapering and pointed (Fig. 6o), but also quite broad (Figs 6p–q), or just slightly tapering (Fig. 6d), too. Though most individuals are visibly decolored, it seems that both males and females have two bright, faded longitudinal dorsal bands on a dark background.

The species M. dahli (Verhoeff, 1901) or subspecies M. austriacum dahli has recently been proven to be just a variant of M. austriacum (see Lazányi et al. 2012) on the same grounds (i.e. variability).

M. austriacum occurs mostly in the Balkan Peninsula, and is very common in Croatia; outside the Balkans it is found in North Serbia and NE Italy (Lazányi et al. 2012).

Figures

FIGURES 14 a – g. Megaphyllum transsylvanicum and Megaphyllum kievenese. M. transsylvanicum syntype male from Transylvania, Romania, ZMB 4380: (14 a) gonopods, posterior view; (14 b) left gonopods, mesal view; (14 c) left promere, meso-posterior view. M. transsylvanicum male from Dobrostan, Bulgaria: (14 d) left opisthomere, antero-lateral view. M. kievense male, after Lohmander (1928): (14 d *) opisthomere. M. transsylvanicum. male from Fruska Gora, Serbia: penis, (14 e) posterior and (14 f) right lateral views. M. transsylvanicum. female from Sinemorec, Bulgaria: (14 g) right vulva, posterior view (setae omitted). Abbreviations: ah: anterior hump, asp: anterior process of the solenomere, ca: central ampulla, ct: connecting tube; pa: posterior ampulla, ph: posterior hump, pp: posterior process, psp: posterior process of the solenomere, rp: rod-like process, P: promere. Scale bars: 0.2 mm.

FIGURES 15 a – g. Megaphyllum hercules. Male from Kechrokambos, Lekanis Mts, Greece: (15 a) gonopods, posterior view; (15 b) right gonopods, mesal view; (15 c) left promere, mesal view; (15 d) left opisthomere, mesal view. Male from Demir Kapija, Republic of Macedonia: penis, (15 e) posterior and (15 f) right lateral views. (15 g) Female from the Rodopi Mts, Greece: left vulva, antero-mesal view (setae omitted). Abbreviations: asp: anterior process of the solenomere, ca: central ampulla, ct: connecting tube; h: hump, op: operculum, pa: posterior ampulla, pp: posterior process, psp: posterior process of the solenomere, P: promere. Scale bars: 0.2 mm.

FIGURES 7 a – g. Megaphyllum carniolense males and a female from Biogradsko Jezero, Montenegro: (7 a) gonopods, posterior view; (7 b) right gonopods, mesal view; (7 c) left promere, meso-posterior view; (7 d) left opisthomere, lateral view; penis, (7 e) posterior and (7 f) right lateral views; (7 g) left vulva, posterior view (setae omitted). Abbreviations: asp: anterior process of the solenomere, ca: central ampulla, ct: connecting tube; pa: posterior ampulla, ph: posterior hump, pp: posterior process, psp: posterior process of the solenomere, P: promere. Scale bars: 0.2 mm.

FIGURES 1 a – e. Schematic opisthomere drawings (right opisthomeres, lateral view). (1 a) Generalized scheme for all Megaphyllum sensu stricto. (1 b) Scheme for the M. projectum species group. (1 c) Scheme for the M. austriacum species group. (1 d) Scheme for the M. transsylvanicum species group. (1 e) Scheme for the M. unilineatum species group. Abbreviations: ah: anterior hump, ap: anterior process, asp: anterior process of the solenomere, ph: posterior hump, pp: posterior process, psp: posterior process of the solenomere.

FIGURES 2 a – f. Megaphyllum erythronotum male from Sfânta Elena, Banat, Romania. (2 a) Gonopods, posterior view; (2 b) left gonopods, mesal view; (2 c) right promere, meso-posterior view; (2 d) right opisthomere, lateral view; penis, (2 e) posterior and (2 f) left lateral views. Abbreviations: asp: anterior process of the solenomere, pp: posterior process, psp: posterior process of the solenomere, P: promere. Scale bars: 0.2 mm.

FIGURES 3 a – f. Megaphyllum metsovoni male from Eptolofos, Parnassos Mts, Greece. (3 a) Gonopods, posterior view; (3 b) right gonopods, mesal view; (3 c) left promere, meso-posterior view; (3 d) left opisthomere, lateral view; penis, (3 e) posterior and (3 f) right lateral views. Abbreviations: ap: anterior process, asp: anterior process of the solenomere, pp: posterior process, psp: posterior process of the solenomere, P: promere. Scale bars: 0.2 mm.

FIGURES 4 a – l. Megaphyllum projectum. (4 a – d) M. p. deubeli male from Băile Borşa, Maramureş Mts, Romania: right gonopods, (4 a) posterior and (4 b) mesal views; (4 c) left promere, meso-posterior view; (4 d) left opisthomere, lateral view. (4 e and 4 g – i) Megaphyllum projectum kochi male from Bukova, Malé Karpaty, Slovakia: (4 e) gonopods, posterior view; penis, (4 g) posterior and (4 h) right lateral views. (4 i) M. p. kochi female from Sanssouci, Germany: right vulva, posterior view (setae omitted). (4 f and 4 j – l) M. p. projectum male [originally Julus austriacus syntype, ZMB 1846]: (4 f) right gonopods, lateral view; (4 j) gonopods, posterior view; (4 k) right gonopods, mesal view. (4 l) M. p. projectum female from Haragistya, Aggtelek National Park, Hungary: right vulva, posterior view (setae omitted). Abbreviations: ap: anterior process, asp: anterior process of the solenomere, ca: central ampulla, ct: connecting tube; pa: posterior ampulla, pp: posterior process, psp: posterior process of the solenomere, sl: serrated lobe, P: promere. Scale bars: 0.2 mm.

FIGURES 5 a – g. Megaphyllum silvaticum. (5 a) Gonopods, posterior view of a male from Begurje, Slovenia; (5 b) left gonopods, mesal view of a male from Staţiunea Borşa, Rodna Mts, Romania; an other male from the same locality: (5 c) left promere, meso-posterior view; (5 d) left opisthomere, lateral view. Penis of the above mentioned Slovenian male, (5 e) posterior and (5 f) right lateral views. (5 g) Left vulva of a female from the above mentioned Romanian locality, posterior view (setae omitted). Abbreviations: ap: anterior process, asp: anterior process of the solenomere, ca: central ampulla, ct: connecting tube; pa: posterior ampulla, pp: posterior process, psp: posterior process of the solenomere, P: promere. Scale bars: 0.2 mm.

FIGURES 8 a – f. Megaphyllum cygniforme. (8 a) Gonopods, caudal view, male from near Sostis, Papikio Mts, Greece. Male individual from near Kechrokambos, Lekanis Mts, Greece: (8 b) right gonopods, mesal view; (8 c) right promere, mesoposterior view; (8 d) right opisthomere, mesal view; penis, (8 e) posterior and (8 f) right lateral views. Abbreviations: asp: anterior process of the solenomere, fl: flagellum, ph: posterior hump, pp: posterior process, psp: posterior process of the solenomere, P: promere. Scale bars: 0.2 mm.