Eupholidoptera smyrnensis ( Brunner von Wattenwyl, 1882 )

Ciplak, Battal, Heller, Klaus-Gerhard & Willemse, Fer, 2009, 2156, Zootaxa 2156, pp. 1-75: 27-35

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Eupholidoptera smyrnensis ( Brunner von Wattenwyl, 1882 )


Eupholidoptera smyrnensis ( Brunner von Wattenwyl, 1882)  

( 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 syntypes M and F, Makedonia, Bosphorus , Smyrna, Beirut ( NMW, ZMB)   .

Material examined: TURKEY: Izmir: Syntypes 1M, 1F, Efes (Ephesus) (Brunner) ( ZMB); 1F, Sevdikoi b. Izmir (Seydikoy-Izmir) (Labaume, S.G.) ( ZMB); 6M, 10F, Bergama-kinik, 12 km W, 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 km wsw 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 km wsw 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, 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, 4–6km ESE of Edirne, 14.7.1979 (L. Willemse & G. Ramaekers) ( CW); 2M, 1F, 3 km westl. 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); Aegean Islands: Kikladhes: 4M, 1F, Tzia I., Korissia-Ioulis, 2 km S of Korissia, 60m, 12.6.2005, N 37º48’01.6’’ E 024º50’54.9’’ (L. Willemse) ( CW); 6M, Tzia I., Stavroudaki-Krthea, 5–250m, along footpath, 12.6.2005, N 37º38’04.6’’ E 024º19’28.2’’(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. chabrieri   festae- 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 km NW 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. smyrnensis   is one of the most widespread species of the genus, its range covers western Anatolia, the southern Balkan (southern Bulgaria and north-western Greece) and the northern and eastern Aegean islands (Thasos, Lemnos, Samos, Nysiros and Rhodes). In the Balkans its range is bordered by the range of E. chabrieri   in the north and northwest and by that of E. megastyla   in the west and south-west ( Ciplak et al. 2007). However, interestingly this species has recently also been recorded from some western Aegean islands such as Evia and Tzia, belonging to the Cyclades archipelago ( Monnerat et al. 1999). Karabag (1958) reported E. smyrnensis   from Maras (east Mediterranean Anatolia) by refering to Bolivar (1899). However, this species has not been recorded from east Mediterranean Anatolia in subsequent studies ( Salman 1983) and this record may refer to E. marashensis   which was later described from this area.

Remarks on the classification of the continental European species of the E. chabrieri   group. When Ramme (1951) established the genus, he listed five Eupholidoptera species   from mainland Europe ( Greece except the Aegean islands, Italy, Bulgaria, former Yugoslavia, Switzerland, France and Italy) or the western Mediterranean islands: Eupholidoptera chabrieri   , E. bimucronata   , E. epirotica   , E. megastyla   and E. smyrnensis   . Later, La Greca (1959) reported four species, E. chabrieri   , E. hesperica   , E. danconai   and E. garganica   , from Italy with 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 beibienkoi   and E. marani   from Bulgaria. In the review of Eupholidoptera   from former Yugoslavia, Adamovic (1972) presented descriptions of three subspecies of E. chabrieri   ; E. chabrieri galvagnii   , E. chabrieri kaltenbachi   and 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 Eupholidoptera   from this area. E. magnifica   ( Costa, 1863) (or Thamnotrizon magnificum   Costa, 1863) was reduced to subspecies level under E. chabrieri   by La Greca (1959) and later it was synonymized with E. chabrieri schmidti   by Willemse (1980). Another subspecies considered to be a synonym of E. chabrieri schmidti   by 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. chabrieri   by Willemse (1980). Moreover, Willemse (1980) reported both E. chabrieri galvagnii   and E. chabrieri usi   to be synonyms of either E. chabrieri schmidti   or E. chabrieri garganica   . However, Massa (1999) considered each of the taxa present in Italy ( E. chabrieri   , E. schmidti   , E. garganica   and 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. megastyla   and E. chabrieri   ). Other forms reported from this area ( E. chabrieri schmidti   , E. chabrieri galvagnii   , E. chabrieri kaltenbachi   , E. chabrieri usi   , E. garganica   or E. chabrieri garganica   , E. marani   and 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. leucasi   and 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 Greece on 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. kinzelbachi   and 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. megastyla   are 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. kinzelbachi   the suggestion by Nadig (1986) that E. kinzelbachi   does not deserve species status is followed in this text. It is worth noting that there are other geographical distinct populations of E. megastyla   in 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. karatolosi   by Mofidi-Neyestanak & Quicke (2007) also shows the typical characteristics of E. megastyla   in 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. megastyla   and this is the agreed classification in this text too.

The case for the remaining forms, which were named E. chabrieri   complex, 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 France up 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, 1959   from C. Italy (Gargano peninsula) (or E. chabrieri garganica   ).

E. marani Peschev, 1960   from SW Bulgaria.

E. beybienkoi Peschev, 1962   from N. Bulgaria.

E. chabrieri usi Adamovic, 1972   from former Yugoslavia (N. Adriatic Isl.).

E. chabrieri galvagnii Adamovic, 1972   from former Yugoslavia (Hercegovina, Montenegro).

E. chabrieri kaltenbachi Adamovic, 1972   from 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. chabrieri   complex 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 Italy and the other at the opposite eastern side of the Adriatic Sea (north-western mainland and Kerkira island of Greece) ( E. garganica La Greca, 1959   or E. chabrieri garganica La 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 type for the population on the eastern side of the Adriatic Sea. There are other forms of the E. chabrieri   complex defined to have non- or slightly curved unfused parts and enlarged apical arms of titillators such as E. chabrieri usi   , E. chabrieri galvagnii   and 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 chabrieri   while 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 type being common (not unique) in the east and the parallel/convergent type being 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 galvagnii   and E. chabrieri usi   are synonyms of E. chabrieri   in agreement with Willemse (1980). Similarly, we agree with Çıplak et al. (2007) that both of the Bulgarian forms, E. marani   and 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. chabrieri   complex (the frontward curvature of the apical arms that is typical for E. chabrieri garganica   and 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, Albania or 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   .

We considered 6 species ( E. chabrieri   , E. megastyla   , E. hesperica   , E. leucasi   , E. epirotica   and E. cephalonica   ) forming a monophyletic clade, the E. megastyla   subgroup, 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.


Naturhistorisches Museum, Wien


Museum für Naturkunde Berlin (Zoological Collections)


University of Nottingham


Departamento de Geologia, Universidad de Chile














Eupholidoptera smyrnensis ( Brunner von Wattenwyl, 1882 )

Ciplak, Battal, Heller, Klaus-Gerhard & Willemse, Fer 2009

Eupholidoptera smyrnensis ( Brunner von Wattenwyl, 1882 )

Ramme, W. 1951: 198

Pholidoptera smyrnensis ( Brunner von Wattenwyl, 1882 )

Ebner, R. 1919: 157

Olynthoscelis smyrnensis ( Brunner von Wattenwyl, 1882 )

Bolivar, I. 1899: 601

Thamnotrizon smyrnensis

Brunner von Wattenwyl, C. 1882: 336