Eupholidoptera megastyla ( Ramme, 1939 )

Ciplak, Battal, Heller, Klaus-Gerhard & Willemse, Fer, 2009, 2156, Zootaxa 2156, pp. 1-75 : 37-42

Eupholidoptera megastyla ( Ramme, 1939)

( Figures 21, 66, 112, 158, 202, 234, 238, Appendix)

Pholidoptera megastyla Ramme, 1939: 101 .

Eupholidoptera megastyla (Ramme) ; Ramme, 1951: 198, 203, 205–207.

Eupholidoptera kinzelbachi Harz, 1981: 174 .

Eupholidoptera danconai La Greca, 1959: 65 .

Eupholidoptera megastyla kinzelbachi Harz, 1981 ; syn. n.

Eupholidoptera karatolosi Mofidi-Neyestanak & Quicke, 2007 syn. n.

Type information: Holotype M, GREECE, Island of Zakinthos (=” Zante ”), 10.6.1937, (W. Ramme); paratypes 4M and 1F, same data as holotype ( ZMB).

Material examined: Specified down to department /district and locality (all CW except otherwise stated). GREECE: Holotype and paratypes; Peloponnese: Akhaia: 4M, 10F, Mt.Erimanthos above Kalentzion 1700–2000m, 4.8.1970 (Willemse & Scherpbier); 1M, A.Pandon-Alepochori, 700m, 17.7.1987 (F. Willemse); 1M, Mt.Panakhaikon, Sella-Pititsa, 450m, 20.7.1987 (F. Willemse); 5M, 2F, above Zastova, 1000–1650m, 29.7.1975 (F. & L. Willemse); 2 M, Mt. Skollis, Portes-Kharavyi, 300m, 20.7.1987 (F. Willemse); 1M, Kato Klitoria, 600m, 17.7.1991 (F. & L.Willeme); 1M, surr. Khalandritsa, 500m, 8.7.1986 (L. Willemse); 1M, Paos E of Tripotamia, 500m, 28.7.1975, F. & (L. Willemse); 2M, Achaia, Kernitsa, 600m, 22.6.1987 (F. Willemse); 2M, 1F, Kalavrita, Mt.Chelmos, 1700–2000m, 1.8.1970 (Willemse & Scherpbier); 1M, 2F, Vlasia, W of Kalavrita, 500m 3.8.1970 (Willemse & Scherpbier); 5M, 2F, Aroania village, W of Klitoria, 1000m, 2.8.1970 (Willemse & Scherpbier); 1M, 1F, K a lavryta, 18– 22.7.1982 (J. Paul.)( NHM); 1M, Kalentzi am Erymanthos (37°57'N, 21°46'E), 900 m, 4.7.1986 (K.-G. Heller)( CH); 1M, Panachaikon Geb. (38°12'N, 21°51'E), 1700 m, 3.7.1986 (K.-G. Heller)( CH); 1F, Panachaikon (38°12'N, 21°51'E), 1500 m, 3.7.1986 (K.-G. Heller)( CH); Ilia: 1F, Killini town, 5m, 2.7.1986 (L. Willemse); 3M, 1F, Lechena, N of Andravida, 50m, 14.7.1974 (F.Willeme); 1M, 1F, Platanos-Pelopion (Pyrgos), 150m, 15.7.1987 (F. Willemse); 6M, 3F, Kallidea, N of Andritsena, 400m, 14.7.1974 (F. Willemse); 1F, Andritsena, 1000m, 20.7.1978 (L. Willemse & J. Tilmans); 1F, Rovia, E of Andritsena, 500m, 13.7.1974 (F. Willemse); 1F, 1.5 km W of Khania Panopoulo, 650m, 14.7.1991 (F. & L. Willemse); 4M, 1F, surr. Bassae, 1100m, 13.7.1974 & 272.7.1970 (Willemse & Scherpbier); Messinia: 1M, Kalamata, Proastion, 10m, 13.7.1987 (F. Willemse); 1M, Kalamata, Mikra Mantinea, 25m, 20.7.1971 (F. Willemse); 7M, 9F, surr. Kambos, 400–700m, 21.7.1971 (F. Willemse); 1M, Karitena (Megalopolis), 500m, 26.7.1970 (Willemse & Scherpbier); 1M, 3F, Pila (Pilos), 125m, 13.7.1987 (F. Willemse); 1M, Umg. Phoinikous (36°48'N, 21°49'E), 20 m, 9.4.1979 (K.-G. Heller & M. Volleth)( CH); 1M, s Ag. Dimitrios (Ag. Nikolaos) (36°48'N, 22°18'E), 5 m, 8.6.1993, (O. v. Helversen)( CH). Lakonia: 1M, Tripi, 2.8.1988 (F. Willemse); 2M, Mistra, W of Sparti, 500m, 25.7.1971 (F. Willemse); 1M, 7 km SE Gythion, orange orchard, 1.10.1992 ( R. Kleukers) (F. Willemse); 3M, 3F, W of Mavrovouni (Gythion), 10m, 12.7.1987 (F. Willemse); 1M, 1F, Karavostasi (Gythion), 5m, 13.7.1987 (F. Willemse); 1M, 1F, Skala (Eyrotas river), 10m, 25.7.1971 (F. Willemse); 4M, 9F, Tayietos Mts., Toritsa [=Topitsa] above Anoyeia, 1200m, 22.7.1971 (F. Willemse); 3M, 4F, below summit Prof. A. Ilias (above Anoyiea), 1700–2200m, 23.7.1971 (F. Willemse); 1F, Mt. Parnon, Vamvakou, 950m, 27.7.1978 (F. Willemse); Arkadia: 1M, 1F, Mt. Mainalon above Kapsia, 750m, 10.7.1974 (F. Willemse); 8M, 2F, Chrissovitsion, 12 km W, 1100m, 12.7.1974 (F. Willemse); 1M, Chrissovitsi, 4 km W, 1100m, 28.7.1978 (L. Willemse & J. Tilmans); 5M, 5F, Ipsous (Dhimitsana), 800m, 27.7.1970 (Willemse & Scherpbier); 1M, 1F, Dhimitsana, 14.7.1977 (M. & J. Duffels) ( ITZ); 1F, Mt. Parnon, A.Vasileos, 1000–1200m, 26.7.1978 (F. Willemse); 1M, above Kastanitsa, 1500–1800 m, 26.7.1978 (F. Willemse); Argolis: 2M, Ligourion, 4 km W, 200m, 6.7.1974 (F. Willemse); 1M, 4F, Ag. Eleni, SE of Epidavros, 450m, 8– 9.7.1974 (F. Willemse); 3M, Nauplion, 12 km E, 7.7.1974 (F. Willemse); 11M, 7F, Mt. Ktenias, Achladokambos, dry habitat, 500 m, 9.7.1974 (F. Willemse); 1M, Mykene (37°44'N, 22°54'E), 12 iv 1982 (E. Bluemm)( CH); Central Greece: Zakinthos Isl.: 3M, 1F, surr. Alikanas, 10m (topotypes) 29.6- 1.8.1986 (L. Willemse); 1M, 3F, surr. A. Nikolaos, 200m, 30.6.1986 Cape Skinari, 50m, 29.6.1986 (L. Willemse); 1F, Kato Gerakari, 50m, 29.6.1986 (L. Willemse); Evia Isl.: 2M, 2F, Steni-Kathenoe, 300m, 22.7.1974 (F. & L. Willemse); 1M, Kimi, 30.6.1971 (A. Kaltenbach)( NMW); Voiotia: 2M, Mt. Parnassos above Arachova, 1800m, 26.7.1971 (F. Willemse); 1M, Palaiokhori, 800m, 20.7.1978 (F. Willemse); 3M, 1F, Mt. Elikonas, SW slopes below summit Paliovouna (1748m), 1500–1650m, WGS84 N 38º17’39.4–49.1’’ E 022º52’44.2–21.3’’, along forest road, dry stony habitat, poor vegetation, mainly above timberline, 25.7.2002 (D. & L.& F. Willemse); Fthiotis: 6M, 10F, Melidoni, W of Arkitsa, moist habitat, seashore, 0–2m, 4.7.1974 (F. Willemse); 1F, Brallos, 600m, 13.6.1986 (F. Willemse); 1F, Mt. Oiti, above Pavliani, 10 km NW, 1750m, 5.8.1972 (F. Willemse); 1M, Pavliani village (Mt. Oiti), Hotel, at light, 24.7.2004 (F. Willemse); 1M, Mt. Oiti, northern slopes, Kastania, 1000m, entrance of village near church & cementary, humid green slope, 22.7.2002 (L. Willemse); 1M, Tymfristos village, 1100m, 5.8.1975 (F. & L. Willemse); 6M, 10F, Melidoni, W of Arkitsa, moist habitat, seashore, 0–2m, 4.7.1974 (F. Willemse); 2M, 2 km n Vitoli, westl. Makrakomi (38°58'N, 22°1'E), 320 m, 1– 30.6.1998 (K.-G. Heller & M. Volleth)( CH) (sound record); 1M, Kenourgio (Camping Venezuela) (38°48'N, 22°44'E), 5 m, 28– 29.7.2003 (K. G. Heller)( CH); 5M, 2 km nördl. Vitoli, westl. Makrakomi (38°58'N, 22°1'E), 330 m, 1– 5.6.1985 (O. v. Helversen)( CH) (sound record); 1M, Pelasyia, 6.7.1976 (J. P. Duffels)( ITZ); Fokis: 1M, Mt. Vardoussia ab. Mousounitsa, 1200m, 6.8.1975 (F. & L. Willemse); 2M, 3F, Kokkinochori, 350m, 17.7.1974 (F. Willemse); 1M, 1F, Mt. Giona, 1750m, above Kaloskopi, Lakkes Lyritsa, WGS84 N 38º38’42.9’’ E 022º19’40.0’’, above timberline, grazed, 23.7.2002 (L.& D. Willemse); Evritania: 1F, Mt. Timfristos above Karpenission, 1500m 4.8.1975 (F. & L. Willemse); 1M, E Agrafa Mts. (Kamaria village) pass about 10km E of Agrafa village, below summit Kornopi, 1650–1800m, WGS84 N 39º08’36.4 – 100’’ E 021º41’32.6–850’’, above timberline, grassy slopes with Juniperus, thistles, stony ground, 6.8.2003 (D. & L. & F. Willemse & T. Blauw); 1M, Mt. Kaliakouda, Megalo Chori-Stournara (=Doliana), 1600–1700m, WGS84 N 39º49’38.7’’ E 021º46’00.6’’, near ass, steep slopes, prickly shrubs, 20.7.2002 (D.& L.& F. Willemse); 1M, 6 km N of A.Theodoros, S of Karpenisson, 1500m, WGS84 N 39º50’56.3’’ E 021º50’35.8’’, small clearing in Abies forest, 19.7.2002 (D.& L.& F. Willemse); 1M, Mt. Helidona above Palaio Mikrochori, SW Karpenisi, 1420–1650m, WGS84 N 39º30’12.6’’ E 021º41’22.0’’, stony ground, grazed, 21.7.2002 (D.& L.& F. Willemse); 1M, between Trovato-Agrafa, S of Tridendro, 690m, WGS84 N 39º09’49.1’’ E 021º37’56.0’’, on car while driving, 16.7.2002 (D. & L. & F. Willemse); 1M, S of Proussos, from Kastania to Aghiro Pighadi, below summit Triandafilia (1817m), 1380–1460m, WGS84 N 38º42’36.2-34.1’’ E 021º41’35.3-19.5’’, scattered coniferous trees, sparse vegetation, stones, 9.8.2004 (D.F.& J.& L. Willemse & T. Blauw); 1M, Kaliakouda--Geb. (38°48'N, 21°47'E), 29.7.2001 (K.-G. & M. Heller)( CH) (sound record). Aitolia/Akarnania: 2M, 1F, Mt. Tsakalakis, S of Ano Chora & Kendriki, 1200–1560m, WGS84 N 38º33’36.7’’ E 021º53’26.9’’, steep stony slopes, trees, herbs, grass, 7.8.2004 (D.F.& J. & L. Willemse & T. Blauw); 1M, nr Eleftheriani, 1060m, pine forest with Juniperus, WGS84 N 38º32’07.2’’ E 021º50’30.9’’, 6.8.2004 (D. & F. & J. & L. Willemse & T. Blauw); 1M, 1F, Terpsithea, 580m, WGS84 N 38º33’34.4’’ E 021º58’11.7’’, Rubus , wayside, 5.8.2004 (D.F.& J.L. Willemse & T. Blauw); 1M, 1F, between Podos & Chomoni along river Kotsala, 400m, WGS84 N 38º35’51.0’’ E 021º51’09.2’’, near bridge, mixed vegetation, 8.8.2004 (D. & F. & J. & L. Willemse & T. Blauw); 1M, NE of Platanos, between Mts. Ardinis & Xerovouni, 1080m, WGS84 N 38º35’30.4’’ E 021º49’12.6’’, pass, 8.8.2004 (D. & F. & J. & L. Willemse & T. Blauw) ( CW). 9M, 4F, Sklavaina, 100m, 1.7.1986 (F. Willemse); 2F, Katafourkon (Makrinoros), 0–100m, 24– 25.6.1972 (J.J.J.Smid & F.H.J.Smid-Elbers); 1M, Katafourka, N of Amfilochia, 0–5m, WGS84 N 38º59’28.7’’ E 021º09’13.8’’, behind beach, Juncus , Phragmites , 13.8.2004 (F. & L. Willemse); 2M, 2F, Anoixiatikon, N of Amfilochia, 30.6.1986 (F. Willemse); 3M, 3F, Acheloos Delta near hill Koutsilaris, 15 km S of Neochori, 5m, 9 & 11.7.1991 (F. & L. Willemse); 3M, 25.6.2000 (F. Willemse); 2M, Umg. Trifos westl. Amfilochia (38°50'N, 21°3'E), 29.6.1999 (K.-G. Heller & M. Volleth)( CH); Ipiros: Arta: 4M, 6F, Ramia-Livadion, 500m, 25.7.1976 (F. & L. Willemse & J.Tilmans) and 1M, 1F ( NHM); 3M, 1F, Mt. Tzoumerka, SW Theodhoriana, below summit Spilia, 1400–1600m, 17– 18.7.1988 (F. Willemse); 1M, 1F, 24.7.1987 (F. Willemse); 1M, 1F, Mt. Tzoumerka above Vourgareli, 1550m, WGS84 N 39º23’32.3’’–38.6’’ E 021º10’57.1’’–58.1’’, above pine forest, rocky ground, road to summits Sklava & Tria Sinora, 28– 29.7.2003 (D. & E. & J. & L. & F. Willemse & T. Blauw); 1M, 1F, Mt. Gavrogo below summit Gerabi above village of Megalochari, 1500–1750m, WGS84 N 39º11-12’46.4-1’’ E 021º16-17’16.4-51.8’’, 5–7 km after turn-of branching of road near Kanalia, stony open slopes, 3– 4.8.2003 (D.& L.& F. Willemse & T. Blauw); 4M 1F, Mt. Tzoumerka above Katarraktis, 1550m, 10.8.2005 (L. Willemse); 2M, 2F, N of Aghnada along road to Konakia, 11.8.2005 (L. Willemse); 2M, Tzoumerka-Geb. östl. Arta (39°24'N, 21°9'E), 5.8.1978 (K.-G. Heller)( CH) (sound record); 5M, 1F, Mt. Tzoumerka above Theodoriana (39°25'N, 21°10'E), 5– 6.8.2003 (K.-G. & M. Heller & M. Volleth)( CH) (sound record). Ioannina: 4M, 2F, Mt. Kakarditsa (connecting Mt. Lakmos and Mt. Tzoumerka), 1800–1850m, WGS84 N 39º35’13.0’ E 021º10’37.5’’ – N 32’42.4’’ E 11’13.2’’, above Matsouki, N of Pramanda, slops below summit Katarachias (2280m), 27.7.2003 (D. & E. & J. & L. & F. Willemse & T. Blauw); 1F, Mt. Lakmos above Mikro Peristeri- Rahoula, 1300m, WGS84 N 39º43’16.3’’ E 021º06’02.7’’, grassy basin, grazed, few shrubs, 13.7.2002 (D. & L. & F. Willemse); 1M, 2F, bridge over Metsoyitiko river at Mikro Peristeri village, 600m, 13.7.2002 (D. & L. & F. Willemse); 1M, Katara Pass, 1750m, 14.7.2002 (D. & L. & F. Willemse); 1M, Metsobon, Valia Kalda, 30– 31.8.2002 (O. v. Helversen); 6M, 2F, above Matsouki, 1150m, N 39º34’ E 21º11’, 9.8.2005 (L. Willemse); 1F, Mt. Peristeri, Sirako – Ag.Athanasios, 1550m, 12.8.2005 (L. Willemse); 2M, Smolikas-Geb. bei Samarina/Distraton (40°4'N, 20°59'E), 25.7.1978 (K.-G. Heller) (sound record). Thessalia: Trikala: 1F, Mt. Chatzi above Sitia, SE of Mesochora, WGS84 N 39º26’09.8’’ E 021º19’10.2’’, open slopes, stony ground, 31.7.2003 (D. & L.& F. Willemse & T. Blauw); 1M, Mesochora, Hotel, at night, 1.8.2003, L. Willemse (1 male); Mt. Avgo above Nea Pefki, NE of Mesochora, 1550–2000m, WGS84 N 39º28’39.1’’ E 021º22’45.0’’ – N 39º29’19.2’’ E 021º22’40.5’’, grassy meadows, rocky slopes, 30.7.2003 (D. & E. & J. & L. & F. Willemse & T. Blauw); 4M, 1F, Mt. Neraidha above Neraidochori, W of Pertouli, 1650m, below summit Boudoura (2010m), WGS84 N 39º33’08.3’’ E 021º26’32.0’’, open pine forest, undergrowth of grasses, Sambucus, Urtica, ferns, verge of road, 1.8.2003 (D. & L. & F. Willemse & T. Blauw); 1M, plateau between Elati (6 km N) and Pertouli, 1250– 80m, WGS84 N 39º32’35.8’’ E 02130’38.0’’, open pine forest, shrubs, wet places, tall grasses, ferns, near asphalt road, 1.8.2003 (D.& L.& F. Willemse & T. Blauw) ( CW); Larisa: 1M, Elasson, 2 km N, 19.7.1978 (L. Willemse & J. Tilmans); 1M, 20 km NW, 22.7.1974 (F. Willemse); 2M, 2F, 4 km W, 7.7.1976 (F. & L. Willemse & J. Tilmans); 1M, Vrisia near Farsala, 200m, 10.7.1987 (F. Willemse); 4M, 20F, Mt. Ossa, Dimitra-Anatoli, 400m, 2– 3.7.1974 (F. Willemse) ( CW); 1M, 4F, Tirnavos, 17.8.1965 (Blommers et al.) ( ITZ); 2 M, 10 km östl. Larissa (39°40'N, 22°24'E), 21.6.1980 (K.-G. Heller) ( CH) (sound record). Karditsa: 1F, 3 km sso Ano Ktimeni (39°5'N, 22°6'E), 700 m, 1– 30.6.1998 (K.-G.K.-G. Heller & M. Volleth)( CH); 1M, 1F (paratypes of E. karatolosi ) Ellinoprigos, 39° 23' 59'' N, 21° 44' 8'' E, 600 m, 15.6.2005 (N. Karatolos) ( NHM). Makedonia: Pieria: 1F, Olymp, oberh. Ag. Dionysios (40°5'N, 22°23'E), 13.8.1979 (K.-G.K.-G. Heller & M. Volleth)( CH); 3M, 1F, Nea Agathopulos, 0–10m, 22.6.1986 (L. Willemse); 14M, 18F, Litochoron, 0–20m, Hotel & beach and surroundings, between 1971 and 1978 (F. & L. Willemse & J.Tilmans); 4M, 1F, Mt. Olimbos, Leptokaria-Karia, 500–1000m, 24.7.1973 & 1.7.1978 (F. Willemse & L. Willemse & J.Tilmans); 1M, Refuge A, 1000–2100m, 16– 18.7.1978 (L. Willemse & J. Tilmans); 1M, 8M, 20F, Mt. Olimbos, Refuge B, 1800–1900m, 14.8.1966 (F. Willemse & J. Scherpbier); 1F, 1900m, 15– 17.7.1969 (F. Willemse); 3F, Mt. Olimbos, W. of Litokhoron, 1450& 1750 m, 13.7.& 2.8.1965; 10km W of Kolindros, 24– 26.7.1965 (both Blommers et al.)( ITZ); 1M, Kolindros, 10 km W, 24– 26.7.1965 (Blommers et al.) ( ITZ); Kastoria: 1M, 5.8.1961 ( R.A. Farrow)( NHM); 1M, Eptakhorion, 190m, 5.8.1978 (L. Willemse & J.Tilmans); Kozani: 5M, 2F, Neapolis, 800m, 31.7.1976 (F. & L. Willemse & J. Tilmans); 1M, Velvendos (40°15'N, 22°4'E), 400 m, 28.6. 1986 (K.-G. Heller) ( CH) (sound record); 9M, 5F, Mt. Pieria, Velvendos, 250m, 23.7.1974 (F. Willemse); Pella: 1M, 4F, Edhessa, 23.7.1965 (Bloommers et al.) ( ITZ); Grevena: 2M, Paraskevi near Deskati, 600m, 8.7.1976 (F. & L. Willemse & J. Tilmans); Magnisia: 1M, Mt. Pilion above Portaria, 700–1200m, 20.7.1974 (F. Willemse); 1M, Lekhonia, 7– 11.8.1965 (Blommers et al.) ( ITZ); ITALY: 5M, 5F, Irsina (Matera-Lucania), 21.7.1957 (M. La Greca) (holotype M and paratypes of E. danconai ) ( MSNM); 1M, 1F, Colloreto (Mt. Pollino, Calabria), 16.7.1950 (paratypes of E. danconai ) ( MSNM); 2M, 1F, Basilicata, E. Irsina (Matera), 500 m, Co latoi, 23.6.2000, ( P. Fontana ) ( CW).

Other records and data on the species: Willemse (1980, 1984), La Greca (1959), Ruffo & Stoch (2005), Nadig (1986), Papapavlou & Katsouni (2008). Additional sound records by Schmidt are in DORSA/ SYSTAX.

Distribution: E. megastyla is one of the most widespread species of the genus, its range covering continental Greece including Peloponnesus, some Ionian and western Aegean coastal islands (see maps in Willemse 1984, Willemse & Willemse 2004, Figure 239) as well as Central Italy on the western side of the Adriatic Sea.

Remarks: The species shows great variations especially in male characteristics such as emargination of last abdominal tergite, structure of cerci and its teeth, lobes of subgenital plate, length of styli, size and asymmetry of titillators, colouration and length of female ovipositor (also see Willemse, 1980). These variations have gradual tendencies and none of them corresponds to any geographic fragment. Consequently, one of such populations which was described as a separate taxon, Eupholidoptera kinzelbachi Harz, 1981 (for details see Nadig, 1986), is now placed in synonymy with E. megastyla .

Recently Mofidi-Neyestanak & Quicke (2007) described two new Eupholidoptera species , E. karatolosi from Northern Greece and E. mirzayani from Khuzestan, Iran. E. karatolosi was described on the base of type material that all came from Ellinoprigos (probably Ellinopirgos), 39° 23' 59'' N, 21° 44' 8'' E, 600 m, situated within the range of E. megastyla . E. karatolosi also shows the typical characteristics of E. megastyla in all structures of male and female genital morphology, especially the titillators with non-divergent unfused parts and indistinct lateral expansions of apical arms and slightly raised seventh sternum of female. From both the description by Mofidi-Neyestanak & Quicke (2007) and examination of the paratypes (1M, 1F) we decided that E. karatolosi Mofidi-Neyestanak & Quicke, 2007 is an objective junior synonym of Eupholidoptera megastyla ( Ramme, 1939) .

The other species Eupholidoptera mirzayani Mofidi-Neyestanak & Quicke (2007) does not belong to the genus Eupholidoptera since it does not show the basic autapomorphic characters of the genus (see the section “Definition of Eupholidoptera in Discussion). Mofidi-Neyestanak & Quicke (2007) did not discuss the relationship of their new species in respect to other Pholidopterini . However, after examination of two paratypes (1M, 1F) we clearly identified them as Apholidoptera kurda (Uvarov, 1916) which is distributed in a range around the borders between Turkey, Iraq and Iran. Since there are no clear differences between Eupholidoptera mirzayani Mofidi-Neyestanak & Quicke, 2007 and Apholidoptera kurda Uvarov , the first has to be accepted as an objective junior synonym of the second.