Pyropteron triannuliformis ( Freyer, 1843 )

Bartsch, Daniel, Pühringer, Franz, Milla, Liz, Lingenhöle, Arthur & Kallies, Axel, 2021, A molecular phylogeny and revision of the genus Pyropteron Newman, 1832 (Lepidoptera, Sesiidae) reveals unexpected diversity and frequent hostplant switch as a driver of speciation, Zootaxa 4972 (1), pp. 1-75 : 47

Pyropteron triannuliformis ( Freyer, 1843)

( Figs 2a , 22, 25 , 144–149 , 169, 170 , 227, 228 )

Sesia triannuliformis Freyer, 1843 : Neuere Beiträge zur Schmetterlingskunde mit Abbildungen nach der Natur, 5 (68): 35, pl. 404 Fig. 2 . Syntypes: Constantinopel [ Turkey, Istanbul] (lost).

Sesia braconiformis Herrich-Schäffer, 1846 : Systematische Bearbeitung der Schmetterlinge von Europa, zugleich als Text, Revision und Supplement zu Jakop Hübner’s Sammlung europäischer Schmetterlinge. Zweiter Band. Die Schwärmer, Spinner und Eulen (1845), 63, 68, pl. 7 Fig. 35 . Syntypes: Hungary and Croatia (lost).

Sesia armeniaca Bartel, 1906 : Societas entomologica, 20 (22): 170. Holotype: ♂, Armenia, Kulp (lost).

Chamaesphecia balcanica Zukowsky, 1929 : Internationale entomologische Zeitschrift, 23 (2): 21. Lectotype: ♂, near Uesküb [ Macedonia, Skopje] (LNMD).

Bembecia (Synansphecia) ljiljanae Toševski, 1986 : Fragmenta balcanica Musei macedonici scientiarum naturalium, 12 (17): 191, 193 Figs 1–2 . Holotype: ♀, Macedonia, Crnicani , Bogdanci (CIT).

Synansphecia pungeleri Špatenka, 1987 : Zeitschrift der Arbeitsgemeinschaft Österreichischer Entomologen, 39 (1/2): 14. Holotype: ♂, Turkey, Anatolia, Konia (NHMW).

This species shows a substantial geographic and individual variability. Large specimens from the Balkans were described as balcanica Zukowsky, 1929 , small and dark specimens from the same area as ljiljanae Toševski, 1986 , and extensively yellow marked specimens from Armenia as armeniaca Bartel, 1906 . Similar but paler specimens occur in central Anatolia. Laštůvka (1990c) carried out extensive morphological studies of this species, including genitalia dissection of more than 130 specimens from various regions, and found no consistent differences between these forms.

The DNA barcode sequences split P. triannuliformis into at least three distinct clades. Although we did not observe reliable morphological differences between these clades, it is pssible that they constitute distinct species or subspecies. To address this issue comprehensively, it will be required to re-examine the type material of all taxa currently subsurmised under P. triannuliformis . In particular, attention will have to be payed to the morphology and barcodes of specimens from north-western Turkey, the type locality of P. triannuliformis . Given that several distinct barcodes seem to cluster around the Black Sea, it will also be necessary to locate the types or designate neotypes for Sesia triannuliformis , Sesia braconiformis Herrich-Schäffer, 1846 , and Sesia armeniaca Bartel, 1906 to stabilized the nomenclature of this group.

Barcodes. RAxML analysis recovers P. triannuliformis as monophyletic group; however, it shows substantial heterogeneity based on DNA barcode sequences. The average intraspecific differences are very high with 5.5% and reach a maximum of 8.2%, highly indicative of hidden diversity. P. triannuliformis falls into three distinct clusters with high geographic correlation. The first cluster includes specimens from central and south-eastern Europe, the second cluster contains a specimen from Georgia, and the third cluster specimens from Turkey and Armenia.

Biology and habitat. Various species of Rumex constitute the hostplants of this species; the following are listed in the literature: R. conglomeratus , R. crispus , R. palustris Smith , R. thyrsiflorus , R. scutatus Linnaeus , R. acetosa , R. acetosella ( Kallies & Sobczyk 1993, Špatenka et al. 1996, Špatenka et al. 1999, Lepidopterologen-Arbeitsgruppe 2000, Laštůvka & Laštůvka 2001, Laštůvka 2007). Špatenka et al. (1996) also mentioned Geranium sanguineum as occasional hostplant. The larvae, seldom more than one, feed in the main root and develop over one year.

Distribution. This species occurs in a wide, partially scattered range from south-western France, southern and northern Italy, central and eastern Europe, through the Balkans, the Ukraine, the Baltic states, southern Russia and Turkey to the Middle East, the Caucasus region, the Caspian Sea coast and northern Iran ( Laštůvka & Laštůvka 1995, 2001, Špatenka et al. 1999).

Note. Most of the standard literature (e.g. Špatenka et al. 1999) cites 1842 as publication year for this taxon. However, we follow the conclusions of Olivier (2000) and consider 1843 the year of publication.

Specimens examined: 1♀, France, Provence, Manosque, 22.VI.1991, leg. DB; 1♀, France, Provence, Drome, Plaisians , 250m, 26.VII.1992, leg. DB; 3♂, 1♀, Italy, Sicily, prov. Catania, S slope of Mt. Etna , 1800–1900m, e.l. ex Rumex scutatus , larvae 3.VI., adults 14.VI.–1.VII.2013, leg. DB (CDB); 14♂, Italy, Sicily, CT, Etna, Rifugio Sapienza , 1900–2050 m, 12.VI.2001, leg. FP (BOX-2219 F12, DNAtax 02-06, DNAtax 02585, photo 47/5/3-10); 23♂, Italy, Veneto, VR, Monte nr. Sant’ Ambrogio , 480 m, 20.VI .1998 , leg. FP; 1♂, ibid., 10.VII .1999 , leg. FP; 1♂, ibid., 6.VII .2002 , leg. FP; 1♂, ibid., 10.VI .2004 , leg. FP (BOX-2221 E09); 6♂, ibid., 16.VI .2018 , leg. FP (CFP); 2♂, 6♀, Germany, Saxony, Hoyerswerda , e.l. 1.–9.VII .2001 , leg. Sobczyk (CCDB-14564-H02); Poland, Poznan-Nowa Wies , 29.VII .1997 , leg. Bakowski (CCDB-14564-G12) (SMNS); 2♂, Germany, ibid., 3.VII .1993 , leg. Sobczyk; 1♀, ibid., Burg-Burghammer , 28.VI .1992 , leg. Sobczyk; 17♂ ( Figs 144–145 ), 24♀ ( Fig. 146 ), Austria, Styria, Sausalgebirge, Kitzeck , e.l. ex R. acetosa , larvae 3.VI., adults 6.VI.–28.VII .1994 , leg. DB (CDB); 1♀, Austria, Burgenland, Inzenhof S Güssing , [e.l.] VI.1948, leg. R. Pinker; 3♂, 1♀, Austria, Niederösterreich, Gumpoldskirchen N Baden, [e.l.] V .1946, leg. R. Pinker; 1♂, Austria, Niederösterreich, Mannersdorf am Leithageb., e.l. 1928, leg. O. Bubacek; 10♂, Austria, Niederösterreich, Mitterretzbach, Heiliger Stein , 20.VI .1999 , leg. FP; 1♀, Austria, Niederösterreich, Retz , e.l. 14.–19.VI .1928 , leg. Kautz; 1♂, 1♀, ibid., e.l. 4.–8.VII .1929 , leg. H. Reisser; 1♀, ibid., e.l. 4.–11.VII .1932 , leg. H. Reisser; 5♂, Austria, Niederösterreich, Retz–Gollitsch , 300 m, 28.V .2005 , leg. FP (BOX-2221 H06, photo 2005/316-327) (CFP); 1♂, Slovenia, Polovnik , 400–500m, 15.VI .2001 , leg. Predovnik; 1♀, Slovenia, Vrhovo , 200m, e.l. ex Rumex sp. , larva 6.VI., adults 3.VII .2001 , leg. Predovnik; 1♂, Bulgaria, Slivno, E Sliven (Slivno), Str. nach Sinite Kamani , 570 m, 10.VII .2014 , leg. FP; 1♂, 1♀, Bulgaria, Burgas, Cape Akra N Chernomorets , 1 m, 7.VII .2014 , leg. FP (photo IMG_25085-110 (♀ )); 2♂, Bulgaria, Burgas, Chernomorets , 3 m, 7.VII .2014 , leg. FP; 4♂, Bulgaria, Burgas, S Sozopol , 10 m, 2.VII .2014 , leg. FP; 5♂, Bulgaria, Burgas, Nos Agalina NE Dyuni , 20–30 m, 5.VII .2014 , leg. FP; 5♂, ibid., 12.VII .2014 , leg. FP; 17♂, Bulgaria, Burgas, St. Thomas (Sveti Toma) S Djuni , 10 m, 1.VII .2014 , leg. FP; 11♂, ibid., 27.VI .2016 , leg. FP; 1♂, Bulgaria, Burgas, Arkutino , 5–20 m, 6.VII .2014 , leg. FP; 1♂, ibid., 8.VII .2014 , leg. FP; 7♂, ibid., 11.VII .2014 , leg. FP; 6♂, ibid., 23.VI .2016 , leg. FP; 2♂, ibid., 28.VI .2016 , leg. FP; 1♂, ibid., 30.VI .2014 , leg. FP (photo IMG_24617-25); 1♂, ibid., 30.VI .2014 , leg. FP; 3♂, Bulgaria, Burgas, S Kiten , 10 m, 11.VII .2014 , leg. FP; 3♂, Greece, Achaia (Peloponnese), Kalogria W Kato Achaia , 5 m, 5.VI .1997 , leg. FP (photo 28/6/29-29/1/8, gen. prep. FP 98/65); 2♂, ibid., 7.VI .1997 , leg. FP; 7♂, ibid., 14.VI .1997 , leg. FP; 25♂, ibid., 18.VI .1997 , leg. FP; 1♂, Greece, Evritania, 8 km W Karpenisi (= Karpenision, Timfristos Mts ), 17.VI .2002 , leg. FP; 1♂, Greece, Komotini, Skaloti ( Rhodopen ), 1090 m, 5.VII .2003 , leg. FP; 1♂, Greece, Larissa, Ossa Mts, SW Stomio (n), 1000 m, 20.VI .2002 , leg. FP (gen. prep. FP 02/34); 1♂, ibid., 1050 m, 2.VII .2003 , leg. FP (BOX-2221 G01) (CFP); 5♂, Greece, Timfi Oros, Ioannina, vic. Kepesovo , 900m, 22.VI .2005 , leg. DB; 2♂, 2♀, Greece, Gravena, Karpero , e.l. ex Rumex sp. , larva 16.V., adult 20.–25.VI .2012 , leg. DB; 1♂, Greece, Monodentri , 1500m, 20.VI .1997 , leg. DB; 1♂, Greece, Timfi Oros, Ioannina, Vrissohori , Aoosbridge , 800m, 25.VI .2005 , leg. DB; 1♂, Greece, Larissa, Ossa Oros , 1km S Karitza, 450m, 8.VII .2008 , leg. DB (BOLD DB-Lep-00123); 3♂, Greece, Florina , 800–900m, 20.VI.–7.VII .1990 , leg. Arheiliger (CDB); 1♀, Greece, Pindos, Timfristos , 1800m, 14.VI .1999 , leg. AL (Bartsch gen. prep. 2019-42) ( Fig. 228 ) (CAL); 1♂, Jugoslawia, Porec , 26.VI .1988 , leg. H. Popp (Bartsch gen. prep. 2019-54) ( Fig. 169 ); 1♀, same data, 20.VI .1988 , (Bartsch gen. prep. 2019-55) ( Fig. 227 ); 2♂, 1♀, Turkey, Bolu, Mengen , 700m, 2.–6.VII .1995 , leg. Riefenstahl; 1♂ ( Figs 147–148 ), Turkey, Bolu, Yenicaga , 1000m, 30.VI .1995 , leg. Riefenstahl; 5♂, Turkey, Sinop, Bayabat , 300m, 24.VI .1995 , leg. Riefenstahl (Bartsch gen. prep. 2019-56) ( Fig. 170 ); 3♂, Turkey, Ordu, Güncyca , 1500m, 22.VI .1995 , leg. Riefenstahl; 3♂, Turkey, Ordu, Fatsa , 18.VI .1995 , leg. Riefenstahl; (SMNS); 5♂, Turkey, Nevsehir, Mustafapasa valley , 1200–1400m, 24.VI .2002 , leg. DB; 1♂, Turkey, Konya, Fasikan Gecidi , 1700m, 30.VI .2002 , leg. DB; 1♂, 1♀, Turkey, Konya, 45km S Beysehir, Bakaran , 1420m, 14.VI .2002 , leg. DB; 1♂, Turkey, Artvin, Kackar Daglari , 1500m, 16.VII .1995 , leg. DB; 2♀ ( Fig. 149 ), Armenia, Ararat prov., vic. Lanjar, St. Karapet , 1880–2000m, 8.VII .2013 , leg. DB; 1♂, Iran, Teheran prov., Elburz Mts , 3km NNW Shemshak, 2860m, 24.VII.2003, leg. Ebert & Trusch (CDB).

Figures

FIGURE 1a. Phylogenetic tree for Palaearctic Synanthedonini plus Nearctic Carmenta based on RAxML analysis of COI barcodes.

FIGURE 2a. Genitalia structures of Pyropteron, males, above P. chrysidiformis chrysidiformis, below P. triannuliformis.

FIGURES 19–26. Pyropteron Biology. 19. Pyropteron aistleitneri, female, reared, Spain, Castile, Torrestio (image Zdeněk Laštůvka). 20. Pyropteron kautzi, female, reared, Spain, Andalusia, Sierra Nevada (image FP). 21. Pyropteron mannii, female, reared, Bulgaria, Michurin (image Zdeněk Laštůvka). 22. Pyropteron triannuliformis male, reared, Italia, Sicily, Mt. Etna (image DB). 23. Pyropteron jordanicum sp. nov., Jordan, Ajloun (image Thomas Stalling). 24. Pyropteron umbrifera, Greece, Epirus, Salaora. 25. Pyropteron triannuliformis habitat with host plant Rumex scutatus, Italia, Sicily, Mt. Etna. 26. Pyropteron umbrifera, habitat with host plant Limonium brevipetiolatum, Greece, Epirus, Salaora (images DB).

FIGURES 135–149. Pyropteron species. 135–137. Pyropteron manni, (135–136) ♂, (137) ♀, Turkey, Bolu. 138–140. Pyropteron hera, (138–139) ♂, Turkey, Saimbeyli, (140) ♀, Turkey, Adiyaman. 141–143. Pyropteron atypica, (141–142) ♂, Turkey, Hakkari, (143) ♂, Iran, Esfahan. 144–149. Pyropteron triannuliformis, (144–146) ♂, ♀, Austria, Styria, (147–148) ♂, Turkey, Bolu, (149) ♂, Armenia, Ararat prov. Scale 10mm.

FIGURES 165–170. Pyropteron species, male genitalia. 165. Pyropteron pipiziformis, ♂, Syria, Ballozeh, genitalia slide DB 2019-28. 166–168. Pyropteron jordanicum, ♂, Israel, genitalia slides DB 2019-27, AL 101, AL 102, note variation of crista sacculi. 169–170. Pyropteron triannuliformis, ♂, (169) Jugoslawia [Croatia], Porec, genitalia slide DB 2019-54, (170) Turkey, Sinop, genitalia slide DB 2019-56. Scale 1mm.

FIGURES 226–232. Pyropteron species, female genitalia. 226. Pyropteron jordanicum sp. nov., ♀, Israel, slide DB 2020-06. 227–228. Pyropteron triannuliformis, ♀, (227) Jugoslawia [Croatia], Porec, slide DB 2019-55; (228) Greece, Pindos, slide DB 2019-42. 229–230. Pyropteron leucomelaena leucomelaena, ♀, (229) Turkey, Konya, slide DB 2019-51, (230) France, Luberon, slide DB 2019-53, 231. P. leucomelaena blaesii ssp. nov., ♀, Morocco, Middle Atlas, slide DB 2020-12. 232. Pyropteron borreyi, ♀, Morocco, High Atlas, slide DB 2007-51. Scale 1mm.

FIGURES 31–38. Pyropteron Biology. 31–34. Pyropteron erodiiphaga, (31) male, reared, (32) larva in root of Erodium arborescens, (33) habitat with host plant Erodium arborescens, Tunisia, Djerba (images Friedhelm Mai), (34) male, Morocco, Middle Alas, Col du Zad. 35. Pyropteron affinis, male, Austria, Tirol, Zirl (images FP). 36–38. Pyropteron meriaeformis, (36) habitat with host plant Rumex acetosella, France, Corsica, Ghisonaccia (image DB), (37) male, reared, Portugal, Guarda, Pinzio (image Zdeněk Laštůvka), (38) female, old specimen, France, Corsica, Ghisonaccia (image DB).