Megaphyllum transsylvanicum (Verhoeff, 1897)

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 : 80-82

Megaphyllum transsylvanicum (Verhoeff, 1897)

Figs 14a–d and 14e–g

Brachyiulus transsylvanicus Verhoeff, 1897 : Verhoeff 1897b: 109–110, Figs IV–V. Brachyiulus transsilvanicus: Verhoeff 1899b: 763 ; 1937: 109, 117; 1929: 617, Figs 16–17 . Brachyiulus (Chromatoiulus) transsilvanicus: Verhoeff 1899a: 194 ; 1940: 8.

Chromatoiulus transsilvanicus: Attems 1927: 233 –234, Figs 311–312.

Chromatoiulus transsylvanicus: Ceuca et al. 1977: 248 , 251–253, 256–257, Figs 2–4 . Chromatoiulus (Chromatoiulus) transsilvanicus: Attems 1940: 306 .

Chromatoiulus (Chromatoiulus) transsilvanicus transsilvanicus: Lokšina & Golovatch 1979: 385 . Chromatoiulus (Chromatoiulus) transsylvanicus: Attems 1959: 305 .

Chromatoiulus transsilvanicus transsilvanicus: Jawłowski 1930: 8 , 10, 12.

Brachyiulus transsilvanicus croaticus Verhoeff, 1929: 617 , Figs 18–19 .

Chromatoiulus transsilvanicus croaticus: Strasser 1971a: 41 .

Chromatoiulus transsilvanicus transdanubicus Loksa, 1962: 163 , Figs 44–48. syn. nov. Chromatoiulus transsylvanicus transdanubicus: Gebhardt 1964: 14 .

Megaphyllum transsylvanicum transdanubicus: Korsós 1994: 38 , Fig. 38.

Megaphyllum transsilvanicum transsilvanicum: Golovatch 1984: 101 –102, 112–113, 130. Megaphyllum transsylvanicum: Enghoff & Kime 2009 ; Lazányi et al. 2012: 12, 26–27, 41.

Material examined. Inv. Nr. 8135, 3♂, Ungarn, Siebenbürgen, Kroatien, [ Hungary, Romania and Croatia], 1919, don. Latzel, Ch. austriacus sensu Latzel det. Attems (NHMW); Hungary: 1♂, Bátaapáti, between Cser-erdő and Cser-dűlő, 2002. IX.25., leg. Sziráki Gy. (HNHM); 1♂, 1♀, Bátaapáti, silver lime forest, 2003. Sept.18., leg. Z. Korsós (HNHM); Republic of Moldova: 1♂, 1♀, Orhei district, Trebujeni Rent valley, 1998. June25., leg. Z. Korsós (HNHM); A20033662, 1♂, gonopods and 7th pleuroterga, slide preparation regarded as syntype, Siebenbürgen [Transylvania] (ZSM); Romania: ZMB 12977 (Nr. 1234 Coll. Verhoeff), 1♂, gonopods, 1st and 2nd leg-pair, Syntypus, slide preparation, Hermannstadt [Sibiu], Baumgarten [Bungard] (MNB); Nr. 1233 Coll. Verhoeff, 1♂, gonopods, slide preparation, Herkulesbad [Bǎile Herculane, Romania] (MNB).

Distribution. Bosnia and Hercegovina (Attems 1929); Bulgaria (Vagalinski & Stoev 2007); Croatia (Strasser 1971b); Greece (Strasser 1976; Lazányi et al. 2012); Hungary: Villányi Mts: Tenkes, Mecsek: Tubes (Loksa 1962); Misina and Tubes (Gebhardt 1964); Orosháza (Matic & Ceuca 1969); Republic of Macedonia (Makarov et al. 2004); Republic of Moldova: Bălţi: Corneşti; Chişinau: Vorniceni (Jawłowski 1930); Bǎrnova (Jawłowski 1935); Romania: Dobrogea (Dobrudzsa): Băneasa, Babadag (Tăbăcaru 1966); Judeţul Vrancea (Tăbăcaru 1976); Sibiu, Gherla, Detunata, Siria, Sîngiorz-Băi, Băişoara, Păltiniş (Ceuca et al. 1977); Podu Olt (Ceuca et al. 1983); Valea Arieşului (Crişan 1999); Munţii Metaliferi: Topliţei cave (Tăbăcaru et al. 2004); Serbia: Obedska Bara, v. Obrež (Makarov et al. 2004); Turkey (Enghoff 2006); Ukraine: Podole: Krzemieniec [today: Kremenets]; Bóbrka [Bibrka]; “Obiżowa, Dniestru” [today: Obiazd on Dnestr River]; Żeżava, near Zaleszczyki [today: Zhezhava near Zalishchiki]; Wołczków (Zaleszczyki Region) [today Vilchkov, Zalishchiki Region] (Jawłowski 1936); Lviv Region, Ternopol Region, Odessa Region (Chornyi & Golovatch 1993).

Diagnosis. Differs from all consubgeners by the distinct rod-like process (rp) below promere’s apex ( Figs 14a–c ). Opisthomere’s ( Figs 14a–b, 14d ) posterior process (pp) long, only overpassed by the anterior solenomere process (asp).

Both sexes have two bright dorsal longitudinal bands on a dark grey ground. Body length and height: males: 24.1–44.6mm, 2.2–3.2mm; females: 35.4–48.8mm, 3.2–3.9mm.

Remarks. This species has a wide distribution range: it can be found in the Balkan Peninsula and in Eastern Europe, from Greece to Ukraine; from sea level up to 1700 m a.s.l.; in diverse habitats as pastures, forests and even in caves (Tăbăcaru et al. 2004).

Some of the examined specimens had unusual body colour: one syntype male from Transylvania, and most male specimens from Moldova were uniformly dark; some other males from Bulgaria and Serbia showed faded, hardly visible dorsal bands. A subspecies called M. t. transdanubicum was described by Loksa (1962). According to literature data (Gebhardt 1964; Korsós 1994) only this subspecies lives in Hungary though Matic & Ceuca (1969) reported one male without subspecific designation from Orosháza, far from the subspecies’ known distribution. Freshly collected males from Bátaapáti fit well into the subspecies’ distribution area. Unfortunately we could not find any type material so we could only rely on the original description, according to which M. t. transdanubicum differs from the nominotypical subspecies by its slightly brighter body colour; wider promere; slightly curved and apically irregularly toothed posterior process of the opisthomere; the pad-like apex of the solenomere which is covered with small spines; the straight or apically slightly curved long, pointed anterior process of the solenomere; and basically to this latter process there can be found a small, posterior toothed process and an anterior hump. We compared Loksa’s (1962) original drawings with the individual collected from Bátaapáti, syntypes of the nominotypical subspecies from Sarajevo, Sibiu and Transylvania and non-types collected from different parts of Europe and we found no difference in the above listed characters, thus M. t. transdanubicum is not a valid subspecies.

M. t. croaticum (Verhoeff, 1929) was described on the basis of one male and four females. The gonopod preparation considered as type in the Zoologische Staatsammlung, München (ZSM) was in poor condition and not suitable for comparison.

4. The Megaphyllum unilineatum — species group Fig. 1e

Anterior ridge (ap) well-developed in all species (except for M. hercules (Verhoeff, 1900)) . Apical humps (ah and ph) more or less developed (hardly detected in M. hercules ). Solenomere with a short, finger-formed posterior process (psp) and a usually longer, also finger-formed anterior process (asp).

Vulvae (15–17g, 20g) differing from other Megaphyllum sensu stricto species in numerous features: shape not cylindrical but more or less skull-shaped, i.e. with a posterior projection (see Figs 15g and 17g ); bursae apically with a more or less distinct posterior hump (h) (see Figs 16–17g , 20g ). Central ampulla (ca) positioned obliquely inside bursa; connecting tube (ct) not spiralled, just rumpled and very thin in relation to the posterior ampulla (pa).

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 16 a – g. Megaphyllum rhodopinum. Male from near Shabran Peak, Slavyanka Mt., Bulgaria: (16 a) gonopods, posterior view; (16 b) right gonopods, mesal view; (16 c) right opisthomere, meso-posterior view; (16 d) right opisthomere, lateral view. Males from near Proastio, Kavala county, Greece: penis form (16 e) posterior and (16 f) left lateral views, (16 f *) another penis, right lateral view. Female from the first locality: (16 g) left vulva, posterior view (setae omitted). Abbreviations: ah: anterior hump; ap: anterior process, 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 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 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 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 19 a – f. Megaphyllum tauricum. Syntype male from Krim, Alupka, Inv. No. 3134: (19 a) gonopods, posterior view; (19 b) left gonopods, mesal view; (19 c) left promere, meso-posterior view; (19 d) left opisthomere, postero-lateral view. Other male from Krim, ZMB 8652: penis, (19 e) posterior and (19 f) right lateral views. Abbreviations: ah: anterior hump; ap: anterior process, asp: anterior process of the solenomere, i: indentation, 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 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 20 a – g. Megaphyllum unilineatum. Male from Sminthi, Xanthi county, Greece: (20 a) gonopods, posterior view; (20 b) right gonopods, mesal view; (20 c) left promere, meso-posterior view. Male from the shore of Lake Vistonida, Lagos, Greece: (20 d) left opisthomere, lateral view. Male from near Sminthi, Kamertsi Mts, Greece: penis, (20 e) posterior and (20 f) right lateral views. Female from Nestos Delta, Kavala county, Greece: (20 g) left vulva, latero-posterior view (setae omitted). Abbreviations: ah: anterior hump; ap: anterior process, asp: anterior process of the solenomere, ca: central ampulla, ct: connecting tube; h: hump, op: operculum, pa: posterior ampulla, ph: posterior hump, pp: posterior process, psp: posterior process of the solenomere, P: promere. Scale bars: 0.2 mm.