Hylaea fasciaria (Linnaeus, 1758)

Sihvonen, Pasi, Skou, Peder, Flamigni, Claudio, Fiumi, Gabriele & Hausmann, Axel, 2014, Revision of the Hylaea fasciaria (Linnaeus, 1758) species group in the western Palaearctic (Lepidoptera: Geometridae, Ennominae), Zootaxa 3768 (4), pp. 469-486 : 471-474

Hylaea fasciaria (Linnaeus, 1758)

fasciaria fasciaria Linnaeus, 1758 , Syst. Nat. (Ed. 10) 1: 521, ( Phalaena ( Geometra )). Europe. (Holo) type female (Linnean Society of London, UK) (examined externally).

biliosata Villers, 1789, Linn. ent. 2: 386, pl. 6, fig. 22 (female), (Phal [aena] Geom [etra]). Syntype (s), [ France]: Bressia [=Bresse, near Lyon]. Scoble (1999) gave the type locality as [ Italy]: Brescia , but this is incorrect.

cleui Leraut, 1993, Entomol. Gall. 4 (4): 235, ( Hylaea ). Holotype male (Muséum National d’Histoire Naturelle, Paris, France), France: Hautes-Alpes, L’Argentiére-la-Bessée (examined externally) [originally as subspecies of fasciaria (Linnaeus) , downgraded from subspecies rank ( Scoble 1999, Leraut 2009] (new synonym). Herewith downgraded from subspecies rank quoting the absence of distinct external features. See Remarks.

neustriaria Hufnagel, 1767, Berlin Mag. 4 (5): 520, ( Phalaena ). Syntype (s), [ Germany]: Berlin region.

prasinaria Denis & Schiffermüller, 1775 , Ankündung syst. Werkes Schmett. Wienergegend: 96, ( Geometra ). Syntype (s), [ Austria]: Vienna district. [Junior primary homonym of Phalaena Geometra prasinaria Hufnagel, 1767 .]

prosapiaria Linnaeus, 1758, Syst. Nat. (Ed. 10) 1: 522, ( Phalaena ( Geometra )). Syntypes (Linnean Society of London, UK), Europe [probably near Åbo (=Turku), Finland] (examined externally).

rufofasciosa Esper, 1794, Die Schmett. 3 Suppl. (5–6): 58, pl. 90, pl. 4, 5, (Ph [alaena] Bomb [yx]).

viridifasciosa Esper, 1794, Die Schmett. 3 Suppl. (5–6): 58, pl. 90, figs 6, 7, (Ph [alaena] Geom [etra]). Syntypes male, female, [Europe].

fasciaria cedricola Wehrli, 1929 , Mitt. Münch. Ent. Ges. 19: 319, pl. 24, fig. 3; pl. 25, fig. 9, 10, ( Ellopia ). Syntypes 9 males, 2 females (Zoologisches Forschungsinstitut und Museum Alexander Koenig, Bonn, Germany), [ Turkey (former Syria)]: Achyr Dagh, Bertiz Jaila, 1800 m (examined, including genitalia) [originally as sp., downgraded from species rank ( Scoble 1999)] (revised status). Herewith downgraded from species to subspecies rank quoting the absence of differential features in the genitalia and the exact barcode-sharing with H. fasciaria . See subspecies description below.

fasciaria flavella Wehrli, 1940 (in Wehrli 1939 –1954), in Seitz, Gross-Schmett. Erde 4 (Suppl.): 322, pl. 24: g, ( Ellopia ), Armenia. [Originally as var, but raised to subspecies rank by Wehrli 1954 (in Wehrli 1939 –1954)]. See subspecies description below.

Description. External characters and pregenital abdomen (diagnostic characters underlined) ( Figures 1 , 6 ): Wingspan male 27–35 mm, female 34–39 mm. Ground colour variable (see Variation), dominant colours being different shades of reddish-brown and green. Medial lines often whitish (see Variation). Medial line angled before costa, basal part moves away from costa (not parallel with costa). Postmedial line angled before costa, weakly angled outwards on inner margin. Medial area often slightly darker than rest of wing, narrowest in middle. Hindwings with postmedial line visible only. Terminal line and fringes near forewing apex normally concolorous with wings. Hindwing postmedial line distinct, curved. Discal spots absent. Wings below as above, but paler. Frons pale-brown to brown-red, thorax and abdomen concolorous with wings. Area between antennae (vertex) white. Antennae white dorsally, male antennae bipectinate, female antennae weakly fasciculate. Hindleg tibia of both sexes with 2+2 spurs. Tympanal organs medium-sized. Sternites and tergites 3–8 of both sexes undifferentiated.

Male genitalia (diagnostic characters underlined) ( Figure 10 ): Uncus setose, subapical part rather wide, apex short, roundish. Socii small, setose. Gnathos absent. Valva narrow, apex wider dorsally, sparsely setose. Valva with subapical spine (occasionally with two spines) in ventral margin. Valva base with narrow, symmetric, sclerotised extension. Transtilla wide plate, anterior margin with two concavities. Juxta small, with two setose patches. Saccus very elongated, weakly curved laterally. Aedeagus narrow, caecum long. Aedeagus with straight additional arm, distance between aedeagus and additional arm narrow, apex not expanded, weakly dentate. Vesica opens at approximately 90 degrees angle. Vesica evenly narrowing tube, base with straight row of microcornuti.

Female genitalia (diagnostic characters underlined) ( Figure 14 ): Papillae anales wide, setose. Apophyses posteriores long, straight. Apophyses anteriores about 1/4 length of apophyses posteriores. Lamella postvaginalis large, horizontally striated, partly sclerotised. Lamella antevaginalis often large, margin roundish, weakly sclerotised ridge. Sterigma with membranous, flower-like frill. Ductus bursae short, weakly sclerotised laterally. Posterior part of corpus bursae narrow, rather long, sclerotised, surface granulate. Anterior part of corpus bursae round, membranous. Signum absent or minute, roundish.

Distribution ( Figure 18 ). Eurasian. In Europe from northern Scandinavia to central Iberian peninsula, central Italy and Greece and from British Isles to Ural mountains. In central Italy the species occurs surely in Tuscany and in the northern Marche, while the identity of specimens from Abruzzo must be confirmed by further research. Outside Europe eastwards through southern Siberia to Transbaikal (Dahuria) and Sakha regions East of Lake Baikal, in Caucasus region (subspecies flavella (Wehrli)) and Turkey (nominate subspecies in the northwesternmost part). In the rest of Turkey replaced by subspecies cedricola (Wehrli) .

Phenology. Bivoltine. In southern Europe from April to May, and from August to September ( Robineau 2007; Redondo et al. 2009), in central Europe from May to October, distinction between generations not clear ( Ebert 2003). In northern Europe (data from Finland, Figure 19 ), from mid-May to October, distinction between generations not clear ( Finnish Entomological database 2013). Caterpillar overwinters.

Biology. Caterpillar feeds on the needles of Pinus sylvestris , Picea abies, Abies alba, Larix decidua ( Mikkola et al. 1989; Ebert 2003; Robineau 2007). Subspecies cedricola (Wehrli) has been reared in captivity on Pseudotsuga menziesii (Bernd Müller, pers. comm.). Adults are nocturnal, attracted to light.

Habitat. Coniferous forests, and less frequently in areas with coniferous trees such as Nordic wetlands. Altitude range from sea level to 1300 m in central Europe ( Ebert 2003) and up to 2300 m in the Pyrenees ( Redondo et al. 2009).

Similar species. All four species in Palaearctic Hylaea fasciaria species group are similar. The diagnostic, external characters shown in Figures 6–9 are somewhat tentative and should not be used in isolation, but should be combined with information on biology, collecting locality, male and female genitalia and DNA barcodes. An overview of diagnostic morphological features is given in Table 1 .

Genetic data. Genetically comparatively homogeneous in Europe and Transcaucasia (n=40, from 11 countries), mean intraspecific variation 0.21%, maximum variation 1.71%. The taxon cedricola from Turkey exactly barcode-sharing (n=10), mean intraspecific variation 0.13%, maximum variation 0.46%. Nearest species: Hylaea mediterranea (minimum pairwise distance 3.3%). See Figure 26 .

Variation ( Figure 1 ). Highly variable. Ground colour varies from grey-reddish to dark grey, to yellowish-green and to various shades of green. Various shades of reddish-brown are dominant in northern Europe, and in northern parts of Scandinavia only these colours exist, whereas in southern Europe various shades of green (f. prasinaria ) are dominant. In many areas both colour morphs coexist. Position, width and colour of medial lines variable, those often stand out weakly in reddish-brown specimens, being almost concolorous with wings, grey or blueish-grey. In green specimens the medial lines are usually white, thinner in females. Numerous infrasubspecific forms have been described, those are summarised in Prout (1912 –1916), Wehrli (1939 –1954) and in Leraut (2009). H. fasciaria ssp. cedricola (Wehrli) ( Figures 1h, 1i ) has wings rather dark green, medial lines are whitish and close to each other.

Forewing margin is weakly concave below apex. Only green specimens are known. We retain taxon valid at subspecies level, due to the concave forewing margin, the narrow medial area and the conspicuous transverse lines not reaching the forewing costa. The taxon is, according to current knowledge, allopatric and restricted to Turkey (and Near East?). H. fasciaria ssp. flavella (Wehrli) ( Figure 1e ) has wings grey-yellowish, and forewing medial line is not visible near costa. We have not had access to extensive materials from the Transcaucasus, apart from two specimens from Georgia. Those were DNA barcoded, and they grouped together with other H. fasciaria specimens. The type specimen of flavella has not been located (collection is not mentioned in the original description), thus we have not been able to establish the identity of the Georgian specimens relative to flavella. We follow Wehrli (1939 –1954, p. 507), who cited Heydemann (1942), in his decision to raise the taxon to subspecies rank, and retain taxon valid at subspecies level. The taxon is, according to current knowledge, allopatric and restricted to Armenia (Transcaucasus). Scoble (1999) did not mention the taxon at all, Viidalepp (1996) consided it valid at subspecies level.

Remarks. H. fasciaria ssp. cleui Leraut , illustrated in Leraut (2009) and Hausmann (2001; fig. 66), ( Fig. 1g is also close) is downgraded from subspecies rank to junior synonym to the nominal subspecies of H. fasciaria (Linnaeus) . Wings are purple-pink to crimson-red and medial lines are ash grey. Taxon is known from southern French Alps. In the adjacent Valesia (southern Switzerland,> 200 specimens examined in the ZSM) such forms are dominant but mixed with green and red forms, potentially supporting the existence of a cline. This indicates that cleui, although locally dominant phenotype, does not constitute a subspecies because it lacks disjunct external features, and it is questionable whether the southern French Alps populations are geographically isolated from other populations in the Alps. DNA barcodes are not available, so far, for French Alps populations.

Figures

FIGURE 18. Distribution area of Hylaea fasciaria (Linnaeus) in the West Palaearctic region. The distribution area, where the species occurs frequently, is shown in grey. The examined records, which appear outside that area, are shown in black symbols.

FIGURE 19. Phenology of H. fasciaria (Linnaeus) in Finland. Data from the Finnish Entomological database (2013).

FIGURE 26. Neighbour joining tree (Kimura 2-parameter distance model for COI-5P marker) for European Hylaea specimens. Terminals with specimen ID-number and species name from BOLD.

FIGURES 1–5. Adults of the Palaearctic Hylaea fasciaria species group. 1a. Hylaea fasciaria (L.) male, 13.vi.1987, Volsk, Saratov Region, South East Russia (coll. Skou). 1b. H. fasciaria male, 6.vii.1972, Skanør, Falsterbo, Skåne, Sweden (coll Skou). 1c. H. fasciaria male, 1.viii.1972, Byerum, Öland, Sweden (coll. Skou). 1d. H. fasciaria male, 18.vii.1970, Dueodde, Bornholm, Denmark (coll. Skou). 1e. H. fasciaria flavella (Wehrli), type, original illustration from Wehrli (1939–1954, plate 24: g), Armenia, without date (coll. unknown). 1f. H. fasciaria female, 6.vii.1972, Skanør, Falsterbo, Skåne, Sweden (coll. Skou). 1g. H. fasciaria female, 29.vii.2000, Terre Rouge, Briancon, Dept. Hautes Alpes, France (coll. Skou). 1h. H. fasciaria cedricola (Wehrli) male, ex ovo, Bey Dağlari, Antalya, Turkey (coll. Müller). 1i. H. fasciaria cedricola female, ex ovo, Bey Dağlari, Antalya, Turkey (coll. Müller). 1j. H. fasciaria male, 27–28.vii.1983, Vorarlberg, 1500 m, Austria (coll. Skou). 1k. H. fasciaria male, 10.vi.2010, Mount Parnassos, 1650 m, Greece (coll. Skou). 1l. H. fasciaria female, 27–28.vii.1983, Vorarlberg, 1500 m, Austria (coll. Skou). 1m. H. fasciaria female, 28.vi.1981, Karpenision, Aetolia, Greece (coll. Skou). 2a. Hylaea mediterranea new species male, 9–10.x.2010, Pizzo della Rondine, Sicily, Italy (coll. Skou), holotype. 2b H. mediterranea male, 9–10.x.2010, Pizzo della Rondine, Sicily, Italy (coll. Skou), paratype. 2c. H. mediterranea female, 9–10.x.2010, Pizzo della Rondine, Sicily, Italy (coll. Skou), paratype. 2d. H. mediterranea female, 5.vi.2005, Castelbuono, Sicily, Italy (coll. Skou), paratype. 3a. Hylaea pinicolaria (Bellier) male, 20–21.vii.2004, Camping Monte Cinto, Haut-Asco, Corsica, France (coll. Skou). 3b. H. pinicolaria male, 20–21.vii.2004, Camping Monte Cinto, Haut-Asco, Corsica, France (coll. Skou). 3c. H. pinicolaria male, 6.vii.1951, La Foce, Vizzavona, Corsica, France (coll. Herbulot, coll. ZSM). 3d. H. pinicolaria female, 23.vi.1994, Haut Asco, Corsica, France (coll. Skou). 4a. Hylaea compararia (Staud.) male, 23.vi.1911, Blida, Glacieres, Algeria (coll. BMNH). 4b. Hylaea compararia female, 13.ix.1911, Glacieres, Blida, Algeria (coll. BMNH). 4c. Hylaea compararia female, 15.ix.1911, Glacieres, Blida, Algeria (coll. BMNH). 5a. Hemithea squalidaria O. G. Costa, original illustration from O. G. Costa, [1848]. Illustration is reproduced under a licence from the Picture Library of the Natural History Museum, London, UK.

FIGURES 6–9. An overview of diagnostic characters in the Palaearctic Hylaea fasciaria species group. Figure 6. H. fasciaria (Linnaeus) (Denmark, 27.vi.1977). Figure 7. H. mediterranea, new species (Italy, Sicily, 5.vi.2005, paratype). Figure 8. H. pinicolaria (Bellier) (France, Corsica, 20.-21.vii.2004). Figure 9. H. compararia (Staudinger) (Algeria, 23.vi.1911).

FIGURES 10–13. Male genitalia of the Palaearctic Hylaea fasciaria species group. Hylaea fasciaria (Linnaeus). 10a. genitalia, Finland: Enonkoski, 16–18.vii.1996, slide PS1344 (coll. P. Sihvonen). 10b. aedeagus, Germany: Bad Harzburg, 2.vii.1988, slide PS1741 (coll. P. Sihvonen). 10c. vesica, Finland: Enonkoski, 16–18.vii.1996, slide PS1344 (coll. P. Sihvonen). 10d. base of vesica, Finland: Enonkoski, 16–18.vii.1996, slide PS1344 (coll. P. Sihvonen). Hylaea mediterranea new species 11a. genitalia, Italy: Sicily, 1000 m, 9–10.x.2010, slide PS1738, holotype (coll Skou). 11b. aedeagus, Italy: Sicily, 1000 m, 9– 10.x.2010, slide PS1738, holotype (coll Skou). 11c. vesica, Italy: Sicily, 1000 m, 9–10.x.2010, slide PS1738, holotype (coll Skou). 11d. base of vesica, Italy: Sicily, 1000 m, 9–10.x.2010, slide PS1738, holotype (coll Skou). Hylaea pinicolaria (Bellier) 12a. genitalia, France: Corsica, 1100 m, 20–21.vii.2004, slide PS1542 (coll. Skou). 12b. aedeagus, France: Corsica, 1100 m, 20–21.vii.2004, slide PS1742 (coll. Skou). 12c. vesica, France: Corsica, 1100 m, 20–21.vii.2004, slide PS1542 (coll. Skou). 12d. base of vesica, France: Corsica, 1100 m, 20–21.vii.2004, slide PS1542 (coll. Skou). Hylea compararia (Staudinger) 13a. genitalia, Algeria: Blida, 23.vi.1911, slide PS1866/BMNH GEO 24840 (coll. BMNH). 13b. aedeagus, Algeria: Blida, 23.vi.1911, slide PS1866/BMNH GEO 24840 (coll. BMNH). 13c. vesica, Algeria: Blida, 5.vi.1908, slide PS1870/BMNH GEO 24844 (coll. BMNH). 13d. base of vesica, Algeria: Blida, 5.vi.1908, slide PS1870/BMNH GEO 24844 (coll. BMNH).

FIGURES 14–17. Female genitalia of the Palaearctic Hylaea fasciaria species group. Circle indicates the point of origin of the ductus seminalis. Hylaea fasciaria (Linnaeus) 14a. genitalia, Finland: Föglö, 28.vii.1995, slide PS1448 (coll. P. Sihvonen). 14b. reduced signum, Finland: Föglö, 28.vii.1995, slide PS1448 (coll. P. Sihvonen). 14c. ostium bursae and adjacent structures, Finland: Föglö, 28.vii.1995, slide PS1448 (coll. P. Sihvonen). Hylaea mediterranea new species 15a. genitalia, Italy: Sicily, 1000 m, 9–10.x.2010, slide PS1739, paratype (coll. Skou). 15b. signum, Italy: Sicily, 1000 m, 9–10.x.2010, slide PS1739, paratype (coll. Skou). 15c. ostium bursae and adjacent structures, Italy: Sicily, 1000 m, 9–10.x.2010, slide PS1739, paratype (coll. Skou). Hylaea pinicolaria (Bellier) 16a. genitalia, France: Corsica, 1100 m, 20–21.vii.2004, slide PS1543 (coll. Skou). 16b. reduced signum, France: Corsica, 1100 m, 20–21.vii.2004, slide PS1543 (coll. Skou). 16c. ostium bursae and adjacent structures, France: Corsica, 1100 m, 20–21.vii.2004, slide PS1543 (coll. Skou). Hylaea compararia (Staudinger) 17a. genitalia, Algeria: Blida, 13.ix.1911, slide PS1868/BMNH GEO 24842, margins of the corpus bursae highlighted (coll. BMNH). 17b. posterior part of corpus bursae (signum absent), Algeria: Blida, 13.ix.1911, slide PS1868/BMNH GEO 24842 (coll. BMNH). 17c. ostium bursae and adjacent structures, Algeria: Blida, 15.ix.1911, slide PS1871/BMNH GEO 24845 (coll. BMNH).