Megaphyllum silvaticum (Verhoeff, 1898) Verhoeff, 1898

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 : 65-67

Megaphyllum silvaticum (Verhoeff, 1898) syn. nov., nomen protectum

Figs 5a–g

Julus austriacus var. nigrescens Latzel, 1884: 300 . nomen oblitum

Julus austriacus: Latzel 1884: 296 –300, partim.

Iulus fasciatus: Attems 1895: 225 –226, Fig. 99.

Brachyiulus austriacus var. nigrescens: Verhoeff 1897b: 113 .

Brachyiulus silvaticus Verhoeff, 1898: 157 , Fig. 26.

Brachyiulus silvaticus: Verhoeff 1907: 306 –308, Fig. 13 .

Brachyiulus silvaticus discolor Verhoeff, 1907: 307 , Fig. 14 , 17–18 .

Chromatoiulus silvaticus: Attems 1927: 226 , Figs 294–296.

Chromatoiulus silvaticus silvaticus: Jawłowski 1936: 272 –273, 276, 278, 288, 293–295, 297, Figs 10–13 . Chromatoiulus silvaticus discolor: Attems 1927: 226 –227, Figs 297–298. Chromatoiulus (Chromatoiulus) silvaticus: Attems 1940: 306 .

Chromatoiulus (Chromatoiulus) silvaticus silvaticus: Attems 1959: 305 . Chromatoiulus (Chromatoiulus) silvaticus discolor: Attems 1959: 305 . Megaphyllum silvaticum silvaticum: Lazányi & Korsós 2010: 39 –41, 43. Megaphyllum silvaticum discolor: Lazányi & Korsós 2010: 39 –43, Fig. 3 . Megaphyllum silvaticum: Enghoff & Kime 2009 ; Lazányi & Korsós 2010: 39–43, 45–46, Figs 2 , 4 –6, 8; 2012: 24, 40.

Material examined. Inv. No. 3090, 2♂, “ Julus austriacus Latzel ( J. fasciatus Koch )” Syntypen, Kärnten (Satnitz), Krain (Adelsberg), Mahren (Adamsthal), Schlesien (Freudenthal), leg., don. Latzel, 1884 I., “107” (NHMW); Inv. No. 3097, 3♂, 2♀, Julus austriacus var. nigrescens Syntypus Nr. 109, Galizien (Przemysl) (NHMW); Märzen (Schönberg), leg., don. Latzel, 1884. I., “109’ Syntypen 5” (NHMW); Inv. Nr. 8134, 1♂, Ungarn, Siebenbürgen, Kroatien, [ Hungary, Romania and Croatia], 1919, don. Latzel, Ch. austriacus sensu Latzel det. Attems (NHMW); Austria: A 20041645, 1♂, gonopods, 7th pleurotergum and one walking leg, Br. silvaticus discolor slide preparation, Spittal [probably Spittal an der Drau, Austria] (ZSM); Hungary: 1♂, 3♀, 1juv., Jósvafő, Tohonya spring, 1999. Oct.5., leg. Z. Korsós (HNHM); Italy: A20033659, 2♂, gonopods, 7th pleuroterga and one walking leg, regarded as Br. silvaticus discolor syntype slide preparation, Pontafel [Pontebba] (ZSM); Romania: Nr. 1253 Coll. Verhoeff, 1♂, gonopods, slide preparation, Kuhhorn [Ineu] (MNB); Slovakia: A20033658, 2♂, gonopods, 7th pleuroterga and one walking leg, regarded as syntype slide preparation (ZSM); Skalka, N. Ungarn [North Hungary, today: Slovakia] (ZSM); Slovenia: 1♂, 21(DL77) Slovenia, above Begurje, 788m, 46°22’41.2” 14°12’26.5”, 2006.04.14., leg. L. Dányi & J. Kontschán (HNHM); Inv. Nr. 8137, 3♂, many ♀ and juveniles, Marburg [Maribor], B. austriacus det. Attems (NHMW).

Distribution. Austria: Gurktaler Alpen: Lieserschlucht near Spittal; West Carinthian Mts: Eberstein; Villach- Klagenfurt Basin: Wörthersee; Gailtaler and Karnische Alpen (Carniolan Alps): Goldeck (Strasser 1959); Eckberg: Kroisenbachgraben (Voigtländer et al. 1997); Marburg, Assling (Attems 1929); Croatia: Hrvatsko Zagorje: Klanjec: Cesargradska-hg., Vukovo Selo, Rozga (Strasser 1965); Hungary: Aggteleki National Park (Lazányi & Korsós 2010); Italy: Tre Venezie (Strasser & Minelli 1984); Romania: Judeţul Vrancea (Tăbăcaru 1976); Poland: Tatry Zachodnia: Dolina Huczawa, Kobylego Wierchu (Jawłowski 1938); Bieszczadów: Roztocza, Pienin, Tatry (Jedryczkowski 1979); Roztocze (Stojałowska & Bielak–Oleksy 1970); Republic of Moldova: (Enghoff & Kime 2009); Slovakia: Skalka (Verhoeff 1941); Kremnica (Dudich 1958); Ondavskej vtchoviny, Nízkych Tatier, Juhoslovenského krasu (Ložek & Gulička 1962); Slovenia: Kočevje (type locality of discolor); “Ainöd”, Sleme Mt. (Attems 1929); Lower Carniolan (Dolenjska) mountains and hillside, Julian Alps (Strasser 1966a); Ukraine: Podole (Jawłowski 1936); Kárpát, Lviv Region, Ivano-Frankivsk Region, Ternopol Region (Chornyi & Golovatch 1993).

Diagnosis. Differs from all consubgeners by the perpendicular anterior process of the opisthomere (ap), i.e. being in a right angle to the opisthomere’s main body, ending with toothed apex ( Figs 5b, 5d ).

Opisthomere’s ( Figs 5a–b and 5d ) posterior process (pp) with three size-variants: rarely rudimentary; sometimes not exceeding the anterior process; sometimes reaching high above the anterior process. Solenomere’s posterior process (psp) mostly higher than the anterior one (asp), depending on individual variation and point of view.

The species shows sexual dimorphism in body colour: males are plain dark grey, while females have two bright longitudinal dorsal bands. Body length and height: males: 26–40.9mm, 2–3mm; females 25–38mm (females according to Verhoeff 1907).

Remarks. Investigation of Julus austriacus var. nigrescens Latzel, 1884 syntypes showed that it is a subjective senior synonym of the later described M. silvaticum . However, here we propose reversal of precedence under Article 23.9. (ICZN 2000). The younger name is valid because the senior name has not been used after 1899 (meeting condition 23.9.1.1), while the junior synonym was used as its presumed valid name in the last 50 years by at least 10 different authors in at least 25 publications encompassing a span of not less than 10 years (e.g. Čaputa 1988; Ceuca 1992; Chornyi & Golovatch 1993; Enghoff & Kime 2009; Foddai et al. 1995; Golovatch 1992; Gulička 1985b; Jedryczkowski 1979; Lác 1991; Lazányi & Korsós 2008; Lazányi & Korsós 2010; Lazányi et al. 2012; Lokšina & Golovatch 1979; Mock 2001; Mock 2002; Piróg et al. 1990; Schmölzer–Falkenberg 1975; Stojałowska & Bielak–Oleksy 1970; Stojałowska & Starega 1974; Strasser 1965; Strasser 1966a; Strasser 1971a; Strasser 1971b; Strasser & Minelli 1984; Tăbăcaru 1976; Voigtländer et al. 1997; Wytwer 2000) meeting condition 23.9.1.2.

M. silvaticum is a widespread, Central European Megaphyllum species which could be easily mistaken for M. projectum , especially because of their overlapping distribution areas. Apart from gonopod details, the two species are indistinguishable by other characters.. Although they have differently shaped anterior processes of the opisthomere (ap), this is noticeable only in lateral and anterior views. Though Lazányi & Korsós (2010) wrote about vulval differences it turned out that there is no difference; the M. silvaticum vulva figure given there shows only a juvenile Megaphyllum sp. female with incompletely developed vulvae. In addition, the species shows similar sexual dimorphism in body colour as M. projectum (except for one case when M. silvaticum females were so dark that the bands were hardly visible). This sexual dimorphism was not part of the original description because Verhoeff (1898) described the species on the basis of a single male.

Verhoeff also described a subspecies called M. s. discolor (Verhoeff, 1907) which was shown to be a variant (see Lazányi & Korsós 2010).

2. The Megaphyllum austriacum — species group Fig. 1c

Anterior process present only in some species as an anterior ridge (ap) (e.g. M. montivagum ). Some species with a small posterior apical hump (ph) (e.g. M. austriacum and M. cygniforme (Lazányi & Korsós 2012)) . Solenomere with a variable, more or less elongated, sometimes wavy posterior process (psp) usually larger than the anterior solenomere process (asp)—sometimes psp looking shorter than/subequal to asp due to individual variation or due to the bent form of the posterior process (e.g. Figs 6b, 7d).

Figures

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 13 a – f. Megaphyllum rosenauense. Syntype male from Transylvania, Romania, ZMB 2622: (13 a) gonopods, posterior view; (13 b) right gonopods, mesal view; (13 c) right promere, posterior view; (13 d) right opisthomere, lateral view. Male from Miercurea Ciuc, Romania: penis, (13 e) posterior and (13 f) right lateral views. Abbreviations: asp: anterior process of the solenomere, ph: posterior hump, pp: posterior process, psp: posterior process of the solenomere, P: promere. Scale bars: 0.2 mm.

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 17 a – g. Megaphyllum sjaelandicum male and female from Dorpat, Estonia, ZMB 8734: (17 a) gonopods, posterior view; (17 b) left gonopods, mesal view; (17 c) right promere, meso-posterior view; (17 d) right opisthomere, mesal view; penis, (17 e) posterior and right lateral views; (17 g) right vulva, antero-mesal view (setae omitted). Abbreviations: ah: anterior hump; ap: anterior process, asp: anterior process of the solenomere, ca: central ampulla, ct: connecting tube; h: hump, l: lobe, op: operculum, pa: posterior ampulla, pp: posterior process, psp: posterior process of the solenomere, P: promere. Scale bars: 0.2 mm.

FIGURES 18 b – d. Megaphyllum spathulatum after Lohmander (1936): (18 b) right gonopods, mesal view; (18 c) left promere, meso-posterior view; (18 d) probably left opisthomere, lateral view. 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, sh: serrated hump, P: promere.

FIGURES 10 a – g. Megaphyllum glossulifer males and a female from Rhodopi Mts, Bulgaria: (10 a) gonopods, posterior view; (10 b) left gonopods, mesal view; (10 c) left promere, meso-posterior view; (10 d) left opisthomere, lateral view; penis, (10 e) posterior and (10 f) left lateral views; (10 g) left vulva, meso-posterior view (setae omitted). Abbreviations: 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 11 a – g. Megaphyllum monticola. Males from Korakas Mts, Greece, ZMB 1300: (11 a) gonopods, posterior view and (11 b) right gonopods, mesal view and (11 c) left promere, meso-posterior view. Males from Ostrovo, Greece, Inv. No. 8019: (11 d) Right opisthomere, lateral view, penis, (11 e) posterior and (11 f) right lateral views, (11 f *) another, bent penis, left lateral view. (11 g) Left vulva, posterior view, female from Korakas Mts, Greece (setae omitted). Abbreviations: asp: anterior process of the solenomere, ca: central ampulla, ct: connecting tube; op: operculum, pa: posterior ampulla, ph: posterior hump, pp: posterior process, psp: posterior process of the solenomere, P: promere. Scale bars: 0.2 mm.

FIGURES 12 a – g. Megaphyllum bosniense. Syntype male from Bosnia, ZMB 2625: (12 a) gonopods, posterior view; (12 b) left gonopods, mesal view; (12 c) right promere, meso-posterior view; (12 d) left opisthomere, meso-posterior view. Male from Treskavac Mts, Montenegro: penis, (12 e) posterior and (12 f) right lateral views. Female from Maljen Mts, Serbia: (12 g) left vulva, meso-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 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 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 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 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.