Thamnotrizon smyrnensis Brunner von Wattenwyl, 1882: 336 Olynthoscelis smyrnensis ( Brunner von Wattenwyl, 1882 ) Bolivar , 1899: 601 Pholidoptera smyrnensis ( Brunner von Wattenwyl, 1882 ) Ebner, 1919: 157 Eupholidoptera smyrnensis ( Brunner von Wattenwyl, 1882 ) Ramme, 1951: 198 2156 Ciplak, Battal Heller, Klaus-Gerhard Willemse, Fer Zootaxa 2009 2009-07-13 2156 1 75 3CML9 (Brunner von Wattenwyl, 1882) Brunner von Wattenwyl 1882 [151,884,1482,1508] Insecta Tettigoniidae Eupholidoptera Animalia Orthoptera 26 27 Arthropoda species smyrnensis  ( Figures 19, 64, 110, 156, 200, 234, 237, Appendix)      Thamnotrizon smyrnensis Brunner von Wattenwyl, 1882: 336.    Olynthoscelis smyrnensis( Brunner von Wattenwyl, 1882);   Bolivar, 1899: 601.    Pholidoptera smyrnensis( Brunner von Wattenwyl, 1882);  Ebner, 1919: 157.    Eupholidoptera smyrnensis( Brunner von Wattenwyl, 1882);  Ramme, 1951: 198, 200, 204.   Type information: Unknown number of syntypesM and F, Makedonia, Bosphorus, Smyrna, Beirut( NMW, ZMB).  Material examined: TURKEY: Izmir: Syntypes1M, 1F, Efes (Ephesus) (Brunner) ( ZMB); 1F, Sevdikoi b. Izmir(Seydikoy-Izmir) (Labaume, S.G.) ( ZMB); 6M, 10F, Bergama-kinik, 12 kmW, 50m, 18.7.1979(L. Willemse & G. Ramaekers) ( CW); 3M, 3F, Ephesus ruins, 50m, 21.7.1979(L. Willemse & G. Ramaekers) ( CW); 1M, Smyrna( Izmir) ( NHM); 5F, Smyrna( Izmir), Menemen, 1930 (Sureya Bey) ( NHM); 4M, 3F, Selçuk, Efes, 200 m, 10.8.1988(D.Sirin & M.S. Taylan) ( AUZM); 1M, 1F, Kemalpasa, Nif Dagi, 784 m, 29.6.2006, N 38°23.904, E 027°24.966 (D.Sirin & M.S. Taylan) (in alcohol) ( AUZM); 1M, Teos (ca. 5 kmwsw Seferihisar) ( 38°10'N, 26°46'E), 20 m, 12.6.2000(K.-G. Heller) ( CH) (sound record); Mugla: 1M, Güllük ( 37°14'N, 27°36'E), 6.4.1987(M. Gebhardt) ( CH); 2M, Hisarönü (ca. 12 kmwsw Marmaris) ( 36°47'N, 28°8'E), 2 m, 8.6.2000(K.-G. Heller) ( CH) (sound record); 2M, Patara ( 36°15'N, 29°20'E), 20 m, 5.6.2000(K.-G. Heller) ( CH); Aydin: 2M, 3F, Miletus ruins, 100m, 25.7.1979(L. Willemse & G. Ramaekers) ( CW); 4F, Paşayayla, 1100 m, 17.7.1988( AUZM); Manisa: 1F, Manisa dag b. Manisa, 10– 11.8.1937(W. Ramme) ( ZMB); Balikesir: 1M, 2F, Edremit Yolu, Dereören, Eǧmir Köyü Kavşaǧı, 150 m, 13.7.1996; Savaştepe, Soma Yolu 5.km, 350 m, 14.7.1996, 1F (B. Çıplak) ( AUZM); Bursa: N.W. Anatolien, Bandirma b. Brussa, 8.1937 (W. Ramme) ( ZMB); 1F, Karacabey, 1932; (D. Wichgraf) ( ZMB); 1M, Uludag, 500 m, 8.7.1962(K.M. Guichard & D.H. Harvey) ( NHM); İstanbul: 1F, Bebek, 5.10.1953(M. Burr) ( NHM); 1M, Kilyos Yöresi, 19.7.1977, (S. Salman) ( AUZM); 1F, Bosphorus, Rumelihisari, 21.7.1951( NHM); 2M and 1F, Rumelihisar, 15.9.1940(M. Burr); 1F, Bosphorus, Bebek (Istanbul), 21.9.1951( NHM); 2M, Bosphorus, Buyukdere, 100 ft, 18.9.1959(K.M. Guichard) ( NHM); Izmit: 1F, Gebze, Darıca, 20.8.1969, (A. Koçak) ( NHM); Çanakkale: 2M, Gallipoli (Gelibolu), 1.6– 10.8.1923(M.J. Martin) ( NHM); Edirne: 1F, 4–6kmESE of Edirne, 14.7.1979(L. Willemse & G. Ramaekers) ( CW); 2M, 1F, 3 kmwestl. Havsa ( 41°34'N, 26°47'E), 20– 21.7.1983(K.-G. Heller) ( CH) (sound record); GREECE: Specimens from Greek mainland see Willemse (1984), Çıplak et al. (2007). Central Greece: Evia: 2M, 1F, Kiavisi, (probably Kriavrisi) 25.7– 5.9.1959(Imp. Coll. Exp.) ( NHM); 1M, Karistos, 27.7.1996(C. Monnerat) (see Monnerat et al. 1999)( CW); AegeanIslands: Kikladhes: 4M, 1F, Tzia I., Korissia-Ioulis, 2 kmS of Korissia, 60m, 12.6.2005, N 37º48’01.6’’ E024º50’54.9’’ (L. Willemse) ( CW); 6M, Tzia I., Stavroudaki-Krthea, 5–250m, along footpath, 12.6.2005, N 37º38’04.6’’ E024º19’28.2’’(L. Willemse) ( CW). Island of Lemnos: 1M, Lemnos, Kastro, 10.7.1936(F. Werner) ( ZMB); Southern Sporades: 1M, Südl. Sporaden, Nysiros (v. Oertzen) ( ZMB); Island of Samos: 5M, 2F, Koutsi, 25–27.6. & Iraion, 13.6. & Pedhias Valmaris, 29.6.1977(M.C. & G. Kruseman) ( ITZ); Island of Rhodes: 1M, b. Stadt Rhodos (v. Oertzen) ( ZMB); 1M, b. Stadt Rhodos (v. Oertzen) (labelled as  E. chabrierifestae-from Coll. Zeuner) ( NHM); 1F, Marasho, Kampa, 29.6.36 (F. Werner) ( ZMB); 1M, 1F, Kremasti, 1– 29.6.1928(G.A. Mavromoustakisi) ( NHM); 3M (one nymph) Lindos ( 36°5'N, 28°4'E), 30 m, 18– 19.4.1983(K.-G. Heller) ( CH) (sound record); 1M (nymph), 1F, Prof. Ilias (road km 46.7) ( 36°15'N, 27°55'E), 600–700 m, 21.5.2005(K. G. & M. Heller ( CH); 1M, 1 kmNW Lindos, 19.5.1983( R. Danielsson) ( CW); 1F, Lardos, maquis, 3.5.1981(M.C. & G.Kruseman) ( CW); 1M, Lindos beach, 8.6.1979(M.C. & G. Kruseman) ( CW); 1M, Kremasti, 9.6.1979(M.C. & G. Kruseman) ( CW); 1M, Paradssi, 3.6.1979(M.C. & G.Kruseman) ( CW).  Other published localities: Salman (1983), Willemse (1984), Naskrecki (1999), Ünal (2006).   Distribution:  E. smyrnensisis one of the most widespread species of the genus, its range covers western Anatolia, the southern Balkan (southern Bulgariaand north-western Greece) and the northern and eastern Aegeanislands (Thasos, Lemnos, Samos, Nysiros and Rhodes). In the Balkans its range is bordered by the range of  E. chabrieriin the north and northwest and by that of  E. megastylain the west and south-west ( Ciplak et al. 2007). However, interestingly this species has recently also been recorded from some western Aegeanislands such as Evia and Tzia, belonging to the Cyclades archipelago ( Monnerat et al. 1999). Karabag (1958)reported  E. smyrnensisfrom Maras (east Mediterranean Anatolia) by refering to Bolivar (1899). However, this species has not been recorded from east Mediterranean Anatoliain subsequent studies ( Salman 1983) and this record may refer to  E. marashensiswhich was later described from this area.   Remarks on the classification of the continental European species of the  E. chabrierigroup.When Ramme (1951)established the genus, he listed five  Eupholidopteraspeciesfrom mainland Europe ( Greeceexcept the Aegeanislands, Italy, Bulgaria, former Yugoslavia, Switzerland, Franceand Italy) or the western Mediterranean islands:  Eupholidoptera chabrieri,  E. bimucronata,  E. epirotica,  E. megastylaand  E. smyrnensis. Later, La Greca (1959)reported four species,  E. chabrieri,  E. hesperica,  E. danconaiand  E. garganica, from Italywith the first species including four subspecies (  E. chabrieri chabrieri,  E. chabrieri schmidti,  E. chabrieri magnifica,  E. chabrieri bimucronata)the other three species were described as new. Peschev (1962)described two additional species,  Eupholidoptera beibienkoiand  E. maranifrom Bulgaria. In the review of  Eupholidopterafrom former Yugoslavia, Adamovic (1972)presented descriptions of three subspecies of  E. chabrieri;  E. chabrieri galvagnii,  E. chabrieri kaltenbachiand  E. chabrieri usi. Relatively recently described taxa are  E. leucasi( Willemse 1980),  E. kinzelbachi( Harz, 1981)and  E. cephalonica( Willemse & Willemse, 2004). In addition to the new taxa described since Ramme (1951)there are some taxonomical rearrangements in  Eupholidopterafrom this area.  E. magnifica( Costa, 1863)(or  Thamnotrizon magnificum Costa, 1863) was reduced to subspecies level under  E. chabrieriby La Greca (1959)and later it was synonymized with  E. chabrieri schmidtiby Willemse (1980). Another subspecies considered to be a synonym of  E. chabrieri schmidtiby Willemse (1980)was  E. chabrieri kaltenbachi. Similarly,  E. garganica,which was later recorded from the eastern side of the Adriatic Sea (western Greece), was reduced to a subspecies under  E. chabrieriby Willemse (1980). Moreover, Willemse (1980)reported both  E. chabrieri galvagniiand  E. chabrieri usito be synonyms of either  E. chabrieri schmidtior  E. chabrieri garganica. However, Massa (1999)considered each of the taxa present in Italy(  E. chabrieri,  E. schmidti,  E. garganicaand  E. bimucronata) to be distinct species. In a more recent study by Çıplak et al. (2007)three species were listed from the Balkans (  E. smyrnensis,  E. megastylaand  E. chabrieri). Other forms reported from this area (  E. chabrieri schmidti, E. chabrieri galvagnii,  E. chabrieri kaltenbachi, E. chabrieri usi, E. garganicaor  E. chabrieri garganica, E. maraniand  E. beybienkoi) were considered as belonging to  E. chabrieri. All the species/forms occurring in mainland Europe (including the western Mediterranean islands), except for  E. smyrnensis, constitute three lineages; 1-  E. epirotica+  E. cephalonica, 2-  E. hesperica+  E. leucasiand 3-  E. megastyla+  E. chabrieri, with all remaining forms in  E. chabrieri. Of these the first pair contains two sister species,  E. epirotica(endemic to Aitolia Peninsula, on the western coast of Greeceon Ionian Sea) and  E. cephalonica(endemic to Greek island Kefallinia in the eastern Ionian Sea). They can be easily distinguished from other species by the apical arms of the titillators lacking a wing-like lateral expansion and with a median furrow dorsally. Similarly, the second species pair,  E. leucasi(endemic to Greek island Lefkas in the western Ionian Sea) and  E. hesperica(restricted to the Calabrian part of Italy) is distinguishable from all other species by their apical arms of titillators with robust fused basal part and long unfused apical parts, and presence of very narrow wing-like lateral expansions. Further, the species in each pairs have unique characters to be diagnosed from each other ( Willemse 1980, Willemse & Willemse 2004) and each is vicariant in distribution and isolated by distinct geographic barriers. Of the last pair,  E. megastyla(including  E. kinzelbachiand  E. danconai) is also diagnosable from the other by apical arms of the titillators with long and touching unfused part and narrow lateral expansions. However their intragroup-taxonomy needs some consideration. Diagnoses of three species given in  E. megastylaare based on length of styli, asymmetry of apical arms of titillators and presence of protuberance on inner margin of apical lobes of male subgenital plate all of which are variable especially on Greek specimens. For  E. kinzelbachithe suggestion by Nadig (1986)that  E. kinzelbachidoes not deserve species status is followed in this text. It is worth noting that there are other geographical distinct populations of  E. megastylain Greece, but, there is no clear cut of these variations in accordance with geography, thus, we consider these to be intraspecific variations. The very recently described  E. karatolosiby Mofidi-Neyestanak & Quicke (2007)also shows the typical characteristics of  E. megastylain all structures of male and female genital morphology and exhibits no autapomorphy to describe it as a distinct species. The Italian population of this lineage was described as  E. danconai.However, since there is no unique feature to diagnose it from the other  E. megastyla-like populations Willemse (1980)put it in synonymy with  E. megastylaand this is the agreed classification in this text too. The case for the remaining forms, which were named  E. chabriericomplex, is much more complicated. In previous studies ( Willemse, 1980; Massa, 1999; Çıplak et al., 2007) the following species/subspecies/forms were reported to be in this complex.   E. chabrieri chabrieri( Charpentier, 1825)from SW Franceup to the Central Balkans.   E. chabrieri schmidti( Fieber, 1861)from NE of Italy(Trieste), NW of former Jugoslavia(Krain; Ljubljana), island of Kerkira ( Greece).   E. chabrieri magnifica( Costa, 1863)from S. Italy( Calabria).   E. chabrieri bimucronata( Ramme, 1927)from S. Italy( Sicily).   E. chabrieri brunneri( Targioni-Tozetti, 1881)from central Italy(Abruzzi).   E. garganica La Greca, 1959from C. Italy(Gargano peninsula) (or  E. chabrieri garganica).   E. marani Peschev, 1960from SW Bulgaria.   E. beybienkoi Peschev, 1962from N. Bulgaria.   E. chabrieri usi Adamovic, 1972from former Yugoslavia(N. Adriatic Isl.).   E. chabrieri galvagnii Adamovic, 1972from former Yugoslavia(Hercegovina, Montenegro).   E. chabrieri kaltenbachi Adamovic, 1972from former Yugoslavia( Serbia, N. Macedonia). The characters used to identify each of these forms were reported to be variable in the Balkan forms ( Ciplak et al., 2007). However, this is the case not only for the area in question (Balkan), but it is also true for the remaining part of the range. Thus they do not provide useful differences to distinguish any of them. In the previous studies the characters used to distinguish the forms of the  E. chabriericomplex were defined from the male subgenital plate (number and length of spine(s) under the styli) and the titillators (length, curvature, divergence of unfused parts and largeness of the lateral expansions along the fused parts of the apical arms). Of these, structure and number of the spines may vary between right and left sides in a single male. The characteristics of the titillators need a further consideration. The long unfused parts of the apical arms of the titillators are typical for the Sicilian population (  E. chabrieri bimucronata), but it is not unique and was observed from the specimens collected from other parts of the range in mainland or Sardinian/Sicilian populations too. Another character is the large and frontward curved unfused parts of the apical arms of the titillators, which is typical for two populations; one in the Gargano peninsula of Italyand the other at the opposite eastern side of the Adriatic Sea (north-western mainland and Kerkira island of Greece) (  E. garganica La Greca, 1959or  E. chabrieri garganicaLa Greca; Willemse, 1980). This character is invariable for the Italian population, but the degree of the curvature is variable and there is no clear limit between curved and non-curved typefor the population on the eastern side of the Adriatic Sea. There are other forms of the  E. chabriericomplex defined to have non- or slightly curved unfused parts and enlarged apical arms of titillators such as  E. chabrieri usi,  E. chabrieri galvagniiand  E. chabrieri kaltenbachi.Since these three forms show intermediate states in respect to these characters, they are not valid to diagnose any of these populations as separate distinct forms. Another character which was assumed to be an autapomorphy of some taxa is related to the position of unfused parts of apical arms of titillators. The apical arms with parallel or converging unfused parts were defined as a character state of  E. chabrieri chabrieriwhile the divergent one as that of  E. chabrieri schmidti. As in the previous forms, it seems that there is a clinal tendency in respect to this character, the divergent typebeing common (not unique) in the east and the parallel/convergent typebeing common in the west of the total range. Since there is no clear gap between the two states we consider it to be an intraspecific variation. Thus, there is no sign allowing us to assume them as reproductively isolated units which could be regarded as distinct species. Consequently, we suggest that  E. chabrieri kaltenbachi, E. chabrieri galvagniiand  E. chabrieri usiare synonyms of  E. chabrieriin agreement with Willemse (1980). Similarly, we agree with Çıplak et al. (2007)that both of the Bulgarian forms,  E. maraniand  E. beibienkoi,are  schmidti-like and are synonyms of  E. chabrieri. Thus, the above listed taxa were suggested to be populations belonging to one highly variable single species,  E. chabrieri( Charpentier, 1825). The two characters used to diagnose the forms in the  E. chabriericomplex (the frontward curvature of the apical arms that is typical for  E. chabrieri garganicaand the reverse state for others; the divergence of the unfused parts of the apical arms, the diverged state for  E. chabrieri s.str.and parallel state for  E. chabrieri schmidti) exhibit a special case with the presence of mixture of intermediate forms in the northern basin of the Adriatic Sea, namely former Yugoslavia, Albaniaor close districts. A paper published during the preparation of this text reported valuable supports to the statements presented above. Lemonnier-Darcemont (2007)experimentally made cross-mating of  E. chabrieri chabrieri( France)and  E. chabrieri schmidti( Greece)and was able to produce hybrids that are viable and fertile. These results are consistent with our assumption that these slightly diverged forms are not reproductively isolated and should be considered as regional forms of a single species, of  E. chabrieri.   FIGURES 117–128.Male subgenital plate (and spines under the styli) of  Eupholidopteraspecies(L— lateral view, V— ventral view) (Scale = 2 mm). 117—  E. giuliae,118—  E. latens,119—  E. mariannae,120—  E. rammei,121—  E. astyla,122—  E. annamariae,123—  E. tasheliensis,124—  E. cretica,125—  E. anatolica,126—  E. tahtalica,127—  E. forcipata,128—  E. jacquelinae(Of the figures given above: 117–122, 124, 127 from Willemse & Heller, 2001; 128 from Tilmans, 2002).   FIGURES 129–142.Male subgenital plate (and spines under the styli) of  Eupholidopteraspecies(L— lateral view, V— ventral view) (Scale = 2 mm). 129—  E. sevketi,130—  E. excisa,131—  E. demirsoyi,132—  E. krueperi,133—  E. tauricola,134—  E. mersinensis,135—  E. kykladica,136—  E. spinigera,137—  E. prasina,138—  E. tucherti,139—  E. karabagi,140—  E. unimacula,141—  E. femorata,142—  E. icariensis(Of the figures given above:133 from Salman, 1983).   FIGURES 143–148.Titillators of  Eupholidopteraspecies(d— Dorsal view, L— lateral view, V— ventral view) (Scale= 2 mm). 143—  E. peneri,144—  E. pallipes,145—  E. gemellata,146—  E. akdeniz,147—  E. annulipes,148—  E. marashensis(Of the figures given above: 144, 145 from Willemse & Heller, 2001; 146 from Ünal, 2002).   FIGURES 149–153.Titillators of  Eupholidopteraspecies(d— Dorsal view, L— lateral view, V— ventral view) (Scale= 2 mm). 149—  E. uvarovi,150—  E. palaestinensis,151—  E. lyra,152—  E. ledereri,153—  E. werneri(Of the figures given above 152, 153 from Ramme, 1951).   FIGURES 154–156.Titillators of  Eupholidopteraspecies(d— Dorsal view, L— lateral view, V— ventral view) (Scale = 2 mm). 154—  E. cypria,155—  E. helina,156—  E. smyrnensis(Of the figures given above 156C, 156C from Willemse, 1980). We considered 6 species (  E. chabrieri, E. megastyla, E. hesperica, E. leucasi, E. epiroticaand E. cephalonica) forming a monophyletic clade, the  E. megastylasubgroup, which is present in continental Europe and the western Mediterranean islands. Other forms of these species may be geographical forms which are not considered in this text. Further detailed studies using several morphometric characters or DNA data may provide a better insight for subspecies classification of these forms. NMW, ZMB Macedonia Beirut Bosphorus Smyrna 26 27 1 syntype