Megaphyllum sensu

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 : 91-92

Key to males of subgenus Megaphyllum sensu stricto on the basis of gonopod characters and body colour patterns

1 Anterior process (ap) well discernible.................................................................... 2

- Anterior process (ap) reduced to a low ridge or fully absent................................................... 5

2 Posterior process (pp) shorter than solenomere............................................................. 3

- Posterior process (pp) longer or subequal to solenomere...................................................... 4

3 Anterior process (ap) perpendicular to the main opisthomere body, with toothed apex ( Fig. 5d ).............. M. silvaticum

- Anterior (ap) process parallel to the main opisthomere body, with normal blunt apex ( Figs 4d, 4f )........... M. projectum

4 Posterior solenomere process (psp) large, massive, standing high above the anterior solenomere process (asp) ( Fig. 2d ). Dark grey animals with very wide reddish longitudinal band......................................... M. erythronotum

- Posterior solenomere process (psp) subequal to the anterior solenomere process (asp) ( Fig. 3d ). Reddish-brown animals with two bright longitudinal dorsal bands............................................................ M. metsovoni

5 Apical anterior hump (ah) well developed into a more or less blunt protuberance in front of the anterior solenomere process (asp)............................................................................................... 6

- Apical anterior hump (ah) not detectable................................................................ 11

6 Promere with a unique rod-like process below its apex ( Fig. 14c ). Dark grey animals with two bright dorsal longitudinal bands; uniformly dark forms also present........................................................ M. transsylvanicum

- Promere with normal apex............................................................................. 7

7 Posterior process (pp) joining the main opisthomere body at a relatively wide angle of ca. 50˚ ( Fig. 17b ). Promere strongly tapering ( Fig. 17c )....................................................................... M. spathulatum

- Posterior process (pp) joining the main opisthomere body at a more acute angle of not more than 20˚–30˚. Promere more grad- ually narrowing..................................................................................... 8

8 Posterior process (pp) shorter than solenomere ( Fig. 16d ). Uniformly dark brown animals without longitudinal lines............................................................................................... M. rhodopinum

- Posterior process (pp) subequal to solenomere............................................................. 9

9 Anterior solenomere process (asp) more than two times longer than the posterior solenomere process (psp) ( Fig. 19d ). Plain brownish animals............................................................................ M. tauricum

- Anterior solenomere process (asp) just slightly exceeding the posterior solenomere process (psp).................... 10

10 Posterior solenomere process (psp) enlarged, thicker than the anterior solenomere process (asp) ( Fig. 17b ). Promere posteri- orly with a large lateral lobe (l on Figs 17a, 17c ). Brown animals with two bright longitudinal bands...... M. sjaelandicum

- Posterior solenomere process (psp) normal, maybe only slightly ticker than the anterior solenomere process (asp) ( Fig. 17b ). Promere without a prominent lateral lobe. Dark grey animals with one bright dorsal longitudinal line...... M. unilineatum

11 Anterior solenomere process (asp) strongly elongated, surpassing by far the posterior solenomere process (psp)......... 12

- Anterior solenomere process (asp) shorter or subequal to the posterior solenomere process (psp)..................... 13

12 Posterior process (pp) subequal to solenomere, apically serrated, as the anterior solenomere process (asp) ( Fig. 14d *). Dark grey animals with two yellowish dorsal longitudinal bands............................................ M. kievense

- Posterior process (pp) much shorter than solenomere, apically smooth like the anterior solenomere process (asp) ( Fig. 13b ). Males uniformly dark grey with reddish legs; females with two bright longitudinal dorsal bands.......... M. rosenauense

13 Posterior solenomere process (psp) much longer and wider than the anterior solenomere process (asp)................ 14

- Posterior solenomere process (psp) subequal to the anterior solenomere process (asp).............................. 17

14 Promere apically with a large toothed or serrated lobe (sl on Fig. 9a ) and a shelf-like protrusion from the distal part of its ante- rior surface ( Fig. 9c )......................................................................... M. dentatum

- Promere without special characters..................................................................... 15

15 Posterior solenomere process (psp) swan-neck shaped; posterior process (pp) higher than solenomere ( Fig. 8d ).................................................................................................... M. cygniforme

- Posterior solenomere process (psp) differently shaped; posterior process (pp) subequal to solenomere................. 16

16 Posterior solenomere process (psp) wavy ( Fig. 11d ). Uniformly dark brown animals................... M. montivagum

- Posterior solenomere process (psp) smooth ( Fig. 10d )............................................. M. glossulifer

17 Posterior process (pp) surpassing solenomere............................................................. 18

- Posterior process (pp) shorter or subequal to solenomere..................................................... 19

18 Posterior process (pp) wide and rounded, fully concealing solenomere from posterior view ( Fig. 12a ); posterior solenomere process (psp) smooth ( Fig. 12d ). Dark grey animals with yellow to reddish dorsal side, usually with a thin blackish mid-dorsal line..................................................................................... M. bosniense

- Posterior process (pp) slender, strongly tapering; posterior solenomere process (psp) wavy ( Fig. 7b ). Dark brown or grey animals with thin mid-dorsal black line........................................................... M. carniolense

19 Posterior process (pp) wide and rounded; anterior and posterior solenomere processes (asp and psp) slender, finger-like ( Fig. 15d ). Very large dark grey animals with narrow, yellow to reddish longitudinal dorsal band.................. M. hercules

- Posterior process (pp) narrow, apically pointed; anterior and posterior solenomere processes (asp and psp) rather short, weakly protruding (Fig. 6d)........................................................................ M. austriacum

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 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 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 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 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 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 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 9 a – g. Megaphyllum dentatum males and a female from Ljuboten, Serbia, Inv. No. 8018: (9 a) gonopods, posterior view; (9 b) right gonopods, mesal view; (9 c) right promere, meso-posterior view; (9 d) right opisthomere, lateral view; penis, (9 e) posterior and (9 f) left lateral views; (9 g) right vulva, anterior 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, pr: protrusion, psp: posterior process of the solenomere, sl: serrated lobe, 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.

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 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 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 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 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.