Megaphyllum projectum projectum Verhoeff, 1894

Lazányi, Eszter & Korsós, Zoltán, 2011, Revision of the Megaphyllum projectum Verhoeff species complex (Myriapoda: Diplopoda: Julida: Julidae), Zootaxa 2864, pp. 43-56 : 45-49

Megaphyllum projectum projectum Verhoeff, 1894

Figs. 1–8 .

Megaphyllum projectum: Verhoeff 1894 a (pp. 323–324, no figures) Brachyiulus projectus var. alticolus (Verhoeff, 1897) (Fig. III) Brachyiulus austriacus Latz. ssp. projectus: Verhoeff 1896 a , Figs. 27–28 Brachyiulus projectus alticolus: Verhoeff 1907 ( Figs. 10–12 ) Brachyiulus projectus dioritanus ( Verhoeff, 1907) ( Figs. 5–8 ) Chromatoiulus projectus dioritanus: Szalay 1940

Megaphyllum (Chromatoiulus) projectum dioritanum: Gulička 1985 Brachyiulus projectus kochi ( Verhoeff, 1907) ( Figs. 3 , 9 ) partim. Chromatoiulus projectus: Attems 1940

Megaphyllum projectum Verhoeff, 1894 : Hoffman 1980 (p. 104)

Material examined. Brachyiulus projectus alticolus : MNB: ZMB 3313: 13, 1Ƥ, syntypes, ( Romania) Bucsecs (Bucegi Mts.), Verhoeff leg.; ZMB 12971a: 13, gonopods, 1st and 2nd legpairs, syntype, slide preparation (Nr. 1268 Coll. Verhoeff), ( Romania) Bucsecs (Bucegi Mts.); ZMB 12971bc: 233, gonopods, syntypes, slide preparation (Nr. 1269 and 1270 Coll. Verhoeff), ( Romania) Bucsecs (Bucegi Mts.); Nr. 1271 (Coll. Verhoeff): 13, gonopods, slide preparation, ( Romania) Schülerspitze (Mt. Postavarul); Nr. 1272 (Coll. Verhoeff): 233, gonopods, slide preparation, ( Romania) Krähenstein (Ciucaş Mts.); ZMB 4382: 13, 1Ƥ, Siebenbürgen (Transylvania); ZSM: A20033649: 13, gonopod, slide preparation, regarded as type, ( Romania) Siebenbürgen, Schuler (Transylvania, Mt. Postavarul); A20060716 (Etk. Nr.: 648): 13, ( Romania) Siebenbürgen (Transylvania); A20060716 (Etk. Nr.: 648): 13, ( Romania) Schulerhöhe (Mt. Postavarul);

Brachyiulus projectus dioritanus : MNB: ZMB 4285: 233, 2 juv. 33, 1Ƥ, syntypes, S.W.-Siebenbürgen (SW Transylvania), Verhoeff leg.; ZMB 12973: 233, gonopods, 2 legpairs, syntypes, slide preparations (Nr. 3175 Coll. Verhoeff), Broos; ZSM: A20033650: 13, gonopod, slide preparation, regarded as type, ( Slovakia) Kremnitz (Kremnica); A20033651: 13, gonopod, slide preparation, regarded as type, ( Austria) Feldbach; A20033652: 13, gonopod, slide preparation, regarded as type, ( Austria) Oberaudorf; A20033653: 13, gonopod, slide preparation, regarded as type, ( Austria) Inntal, Oberaudorf; A20033654: 13, gonopod, slide preparation, regarded as type, ( Italy) Sterzig (Vipiteno); A20033655: ( var. medius ) 13, gonopod, slide preparation, regarded as type, ( Germany) Grünkopf, Oberbayern; Reg. Nr. 20060718: 13, Österreich ( Austria), Steiermark (Styria);

Megaphyllum projectum : MNB: Nr. 1260 (Coll. Verhoeff): 13, gonopods, slide preparation, ( Romania) Kronstadt (Braşov); Nr. 1261 (Coll. Verhoeff): 13, gonopods, 1st (modified) legpair, 1 legpair, slide preparation, ( Hungary) Oedenburg (Sopron); Nr. 1262 (Coll. Verhoeff): 13, gonopods, slide preparation, ( Romania) Kronstadt (Braşov); Nr. 1263 (Coll. Verhoeff): 13, gonopods, slide preparation, ( Romania) Kronstadt (Braşov); Nr. 1265 (Coll. Verhoeff): 13, gonopods, slide preparation, ( Romania) Bucsecs (Bucegi Mts.); Nr. 1266 (Coll. Verhoeff): 13, gonopods, slide preparation, ( Austria) Gleisdorf; Nr. 1267 (Coll. Verhoeff): 13, gonopods, slide preparation, ( Romania) Hermannstadt (Sibiu); Nr. 1756 (Coll. Verhoeff): 13, gonopods, slide preparation, ( Germany) Partenkirchen; Nr. 1757 (Coll. Verhoeff): 13, gonopods, slide preparation, ( Romania) Herkulesbad (Bäile Herculane); Nr. 1758 (Coll. Verhoeff): 13, gonopods, slide preparation, ( Hungary) Schwabenberg (Svábhegy); Nr. 2546 (Coll. Verhoeff): ( var. hungaricus ) 1Ƥ, ( Hungary) Schwabenberg (Svábhegy); ZMB 1809: 13, Steiermark (Styria); ZMB 3198: 13, Ungarn ( Hungary), ZMB 4381: 23, ( Croatia) Istrien (Istria); HNHM: 13, Hungary, Balaton-felvidéki NP, Fekete Mt., Querco petrae -Carpinetum, 27. June 2009, leg. Farkas & Lazányi; 13, Hungary, Balaton-felvidéki NP, Fekete Mt., Quercetum petreae - cerris , 27. June 2009, leg. Farkas & Lazányi; 13, 1Ƥ, Hungary, Budakeszi, CT46, safari park, 1988. Oct. 13., leg. I. Fürjes; 13, Hungary, Mátra Mt., Mátraszentimre, Büdös well, DU10, “No. 74”, 690m, 1975. Apr. 0 9., leg. A. Varga; 13, 1Ƥ, Hungary, Nógrád county, Ipolytarnóc, paleontological park, 1997. Apr. 12., leg. Z. Korsós; 13, 1Ƥ, Hungary, East-Mecsek, Bátaapáti, Tilio argenteae- Fraxinetum, 2004. Sept. 30, leg. Z. Korsós & A. Horváth; 13, 1Ƥ, Hungary, Bakony, Bánd, YN11, oak, 1991. VI. 11., leg. Z. Korsós; 13, Hungary, Nyírség, Bátorliget Nature Reserve, next to the research house, ET99, 1990. June 29. leg Z. Korsós; 13, Hungary, Vértes Mts., Csákvár, Haraszt Mt., Querco petrae -Carpinetum, 1985. Oct. 12., leg. Z. Korsós; 13, Hungary, Börzsöny Mts., “2686/1935”, 1921, leg. Pongrácz Sándor; 13, Hungary, Budai Mts., Nagykovácsi, Kutya Mt., Quercetum petreae - cerris, 1988 . March 24., leg. Z. Korsós; 13, 1Ƥ, Hungary, Budapest, Széchenyi Mt., TV-station, CT46, 1989. March 25., leg. Molczer; 13, 1Ƥ, Hungary, Tömörd, soil trap, 2001; 13, Hungary, Baranya county, Palkonya, Palkonya forest, 2003. Oct. 24., leg. O. Merkl; 13, Hungary, Börzsöny Mts., Verőcemaros, Morgó and Testvér spring, yellow sign, CU50, 1991. March 31., leg. B. Farkas; 13, Hungary, Somogy county, Boronka-melléki Nature Reserve, Hosszúvíz, XM85, 1990. Oct. 25., leg L. Ábrahám; 13, Hungary, Nógrád county, Garáb, Macska Mt., oak forest, 1988. Apr. 24., leg. Korsós & Réka; 13, Hungary, Bükk NP, Nagyvisnyó, Bánkút, beech forest, DU62, 1988. Oct. 0 6., leg. Z. Korsós; 13, Hungary, Mecsek Mts., Orfű, Beech forest, BS71, 1991. Sept. 12., leg. Z. Korsós & H. Read; 13, Hungary, Várvölgy, Keszthelyi Mts., 1972. Apr. 22.–Jun. 26., leg. Dr. L. Tóth; 13, Hungary, Zala county, Zala hills, Lovászi, Koldustető hill, 2007. Sept. 18., leg. L. Dányi, J. Kontschán, D. Murányi & Zs. Ujvári; 13, Hungary, Zala county, Zala hills, Szentmargitfalva, Bikucsaerdő, N46°30’ E16°39’, 183m, 2007. Sept. 18., leg. L. Dányi, J. Kontschán, D. Murányi & Zs. Ujvári; 433, Hungary, Aggteleki NP, Szelcepuszta, 1989. III. 21–V. 11., leg. O. Merkl; 13, Hungary, Aggteleki NP, Őzescrag, DU77, 1988. July 30., leg. Z. Korsós; 13, Hungary, Aggteleki NP, Ménes-valley, 1988. Nov. 16., soil trap, leg. Z. Korsós & O. Merkl; 13, Hungary, Aggteleki NP, Patkós-side, 1987. May 13., leg. Z. Korsós; 13, Hungary, Aggteleki NP, Szögliget, Ménes-valley, 1988. May 0 2., leg. Z. Korsós; 13, Hungary, Aggteleki NP, Jósvafő, Nagyside, 1988. Aug. 0 3., leg. Z. Korsós; 13, Hungary, Aggteleki NP, Szin, Háló-valley, 1990. Apr. 24., leg. O Merkl; 233, 1Ƥ, Hungary, Aggteleki NP, Szögliget, Patkós-valley, soil trap, Aegopodio -Alnetum, 350m, 1987. May– Aug., leg. Szél & Ádám; 13, Hungary, Aggteleki NP, Jósvafő, hillside, under stones, 1990. Apr. 25., leg. O. Merkl; 333, Hungary, Aggteleki NP, Szin, Patkós-valley, Aegopodio -Alnetum, soil trap, 1987. May. 0 2., leg. Ádám; 1133, Hungary, Aggteleki NP, Aggtelek, Ménes-valley, soil trap, Querco petrae -Carpinetum, 250m, 1988. May 02.–Sept. 26., leg. O. Merkl; ( Megaphyllum projectum det. Z. Korsós 1990) 233, Hungary, Aggteleki NP, Szögliget, Szádvár, 1990. Oct. 19., leg. O. Merkl; 13, Hungary, Aggteleki NP, Szin, Patkós-valley, Puska Pálspring, DU77, 1989. Apr. 27., leg. I. Fürjes; 433, Hungary, Aggteleki NP, Aggtelek, Patkós-side, Querco petrae - Carpinetum, soil trap, 1987. June–1988. May, leg. Z. Korsós & O. Merkl; 13, Hungary, Aggteleki NP, Aggtelek, Querco petrae -Carpinetum, soil trap, 1989. March. 21.–May 0 9., leg. O. Merkl; 233, Hungary, Aggteleki NP, Szin, Szelcepuszta, Querco petrae -Carpinetum, soil trap, 1989. May 07.–June 29., leg. O. Merkl; 1Ƥ, Hungary, Aggteleki NP, Szin, Szelcepuszta, 1989. March 20., leg. O. Merkl; 1Ƥ, Hungary, Aggteleki NP, Ménes-valley, Querco petrae -Carpinetum, 1989. March 21., leg. O. Merkl; 1Ƥ, Hungary, Aggteleki NP, Ménes-valley, Ménesstream, DU77, 1989. Apr. 27., leg. I. Fürjes; 1Ƥ, Hungary, Aggteleki NP, Szögliget, Patkós-valley, 1988. May 0 2., leg. Z. Korsós; 1Ƥ, Hungary, Aggteleki NP, Jósvafő, Tohonya-valley, DU67, 1999. May 24., leg. R. D. Kime; 1Ƥ, Hungary, Aggteleki NP, Haragistya, 2009. June; 1Ƥ, Hungary, Keszthelyi Mts., Vállus, XM78, 1994. May 23., Leg. Z. Korsós; 3ƤƤ, Romania, Maramureş Mts., Mţii. Rodnei, Säcel, Pietrii Pass, spruce forest, 1200m, singled, 2006. Sept. 20., leg. A. Grabant, Z. György, O. Merkl & A. Podlussány; 1Ψ, Romania, Maramureş county, Rodna Mts., Säcel, Iza spring, pine forest, 900m, 2006. Sept. 19., leg. A. Grabant, Z. György, O. Merkl & A. Podlussány; 1Ƥ, Romania, Maramureş county, “2006/9”, Ţibleş Mts., Dragomireşti, Poienii valley, stream in a beech forest, 901m, N47°32’ 58 E24°16’ 29, leg. L. Dányi, J. Kontschán & D. Murányi; 1Ƥ, Romania, Läpuş, Mtii. Väleni, Secätura, 754m, 2006. May 23., L. Dányi, M. Földvári, J. Kontschán & D. Murányi; 1Ƥ, Romania, Maramureşului basin, Rona de Sus, Héra, 2007. June 28., leg. A. Grabant, Z. György, O. Merkl & A. Podlussány.

Diagnosis. Males 25.9–53.9 mm long, 1.8–3.2 mm high, (44+2+T)–(52+1+T) segments, adult males varied between the developmental stadia X–XII. Overall body colour greyish black with a longitudinal black line dorsally. Ozopores on the suture, form of preanal process variable, sometimes straight but mostly curved dorsally. Male genitalia ( Fig. 1 ): promeres ( Fig. 2 ) with special riffle for holding opisthomere; apical end of promere with a serrated lobe; opisthomere ( Figs. 3–7 ) with two well-pronounced processes, both well-developed, serrated and directed ventrally (parallel with the telopodite). Telopodite with two processes, a side lobe and a dermal lobe ending in a flagellal hook.

Females 30–51.9 mm long, 2.6–4.2 mm high, with (42+1+T)–(49+1+T) segments, adult females varied between the developmental stadia XI–XIII. Overall body colour variable, mainly brownish with a black longitudinal line dorsally, surrounded by two yellowish lines. Female genitalia: two big ampullas, the dorsal wide, bulbous one joining the elongated, well-developed ventral one by a long spiral apodemic tube ( Fig. 8 a–c). Vulval chaetotaxy not clear, all vulval parts bear randomly arranged setae.

Remarks. Following Verhoeff (e.g. 1907) the anterior process of the opisthomere was frequently called “Schutzblattfortsatz” while the posterior one “Mesomeritfortsatz” but its position clearly shows that it is not a part of the mesomere ( Attems 1927). In this genus, according to Attems’ (1927) nomenclature, the mesomere is a paracoxite or coxal process ( Attems 1940) and it is similar to the “Schutzblattfortsatz”. There is a telopodite situated apically on the opisthomere, with two processes: anterior called “Nebenlappen” i.e. side lobe, and posterior “Hautlappen, welches sich an das Flagellumhäckchen anschließt” ( Verhoeff 1907) i.e. “dermal lobe”, ending in a flagellal hook.

Different levels of within-population variability of male gonopods were found: (1) differences of the same opisthomere due to changing its orientation under microscope ( Figs. 5–7 ), (2) differences between right and left opisthomeres ( Figs. 8 d–e, g–h), (3) differences among individuals from the same vial ( Figs. 8 d–f), collecting locality or region ( Figs. 8 d–i). E.g. if we have a caudo-lateral view of the opisthomere ( Fig. 5 ), the anterior process seems to be thinner, the posterior and postero-apical processes wider. However, if we change to antero-lateral view ( Fig. 7 ), the opisthomere seems to have a wide anterior and a thin posterior process and the postero-apical process seems to be very thin. The first character combination describes B. p. projectus , the second fits for B. p. dioritanus . Therefore we concluded that B. p. dioritanus is not a valid subspecies. This problem of positioning has already been observed at the Craspedosoma rawlinsii -group, where 195 nominal taxa had been reduced to only 6 ( Hauser 2004).

Individuals previously identified as B. p. alticolus showed the following characters: one intact male type specimen about 26 mm long, in stadium X. All other older males, in stadium XI, therefore longer, out of the proposed range of B. p. alticolus (which would be 25–28 mm for males (Verhoeff 1897)). Shape of promere variable, sometimes like a B. p. projectus , or even tapering-like. The relative length of the two apical processes on the opisthomere was also variable, sometimes fitting the description of B. p. alticolus but long postero-apical processes were found, too. In addition, the within-population study provided individuals with even-length apical processes, while individuals in the same vials had long + short apical processes ( Figs. 8 d–e). Therefore we concluded that B. p. alticolus is not a valid subspecies, only an infrasubspecific category which has no standing in nomenclature ( ICZN 2000).

The male specimen from Berchtesgaden, Germany, identified as B. p. kochi by Verhoeff seems to be a true M. p. projectum .

The external morphology of M. projectum vulvae was already represented in an early, little-known pioneer work by Sziráki (1966), while the inner structure was illustrated by Lazányi & Korsós (see Fig. 7 in Lazányi & Korsós 2010).

We found a M. p. projectum female with elongated dorsal ampulla from the Aggtelek NP which lies on the distributional border of the two valid subspecies. An atypical male with intermediate-shaped promeres was also found in the ANP. The existence of intermediate forms in the overlapping distribution areas confirms the subspecies status of M. p. projectum and M. p. kochi .

Figures

FIGURES 1 – 7. (1) In situ gonopods of a M. p. projectum male, from ANP, Hungary, caudal view. (2) Right promere of a M. p. projectum individual, from ANP, Hungary, mesal view. (3) Right opisthomere of the same individual, lateral view (scale bars: 0.5 mm). (4) Same individual, apical end of left opisthomere, mesal view (scale bar: 0.1 mm). Right opishomere of the same individual (5) caudo-lateral view; (6) lateral view; (7) antero-lateral view (scale bars: 0.2 mm).

FIGURE 8. M. p. projectum genitalia variation: (a) – (c) female vulvae, (d) – (i) males, right and left opisthomeres from mesal view. (a) ANP, Szin, Szelcepuszta, left vulva caudal, right vulva mesal view; (b) ANP, Jósvafő, Tohonya Valley, left vulva caudal, right vulva mesal view; (c) Romania, Maramureşului Basin, Rona de Sus, left vulva caudal, right vulva anterior view. (d) – (h) five males from the same vial, ANP, Aggtelek, Ménes valley. (i) ANP, Aggtelek, Patkós Valley (scale bars: 0.2 mm).

FIGURES 9 – 13. (9) Left promere of the M. p. deubeli specimen from Bǎile Borşa, Maramureş Mts., Romania, mesal view. (10) Left opisthomere of the same individual, lateral view (scale bars: 0.2 mm). The same opisthomere from (11) caudo-lateral view; (12) lateral view; (13) antero-lateral view (scale bars: 0.1 mm).