Lyces, WALKER, 1854

Miller, James S, 2009, Generic Revision Of The Dioptinae (Lepidoptera: Noctuoidea: Notodontidae) Part 2: Josiini, Bulletin of the American Museum of Natural History 2009 (321), pp. 675-1022 : 753-762

publication ID	https://doi.org/ 10.1206/321.1-1
persistent identifier	https://treatment.plazi.org/id/03FF87E0-FFFB-9E16-BF5E-1434FB0549A8
treatment provided by	Felipe
scientific name	Lyces
status

Treatment

LYCES WALKER, 1854 View in CoL

Figures 320–325 View Fig View Fig View Fig View Fig View Fig View Fig ; plates 29–31

Lyces Walker 1854: 366 View in CoL . Type species: Lyces angulosa Walker, 1854 View in CoL (by original designation).

Leptactea Prout, 1918: 397 View in CoL . Type species: Actea minuta Druce, 1885b View in CoL (by original designation). New synonymy.

DIAGNOSIS: Taken as a whole, Lyces shows remarkable variation in wing pattern. This variability includes: Species with a yellow or orange-yellow transverse FW band and a corresponding yellow or orange-yellow central area in the HW (e.g., L. angulosa and L. constricta , pl. 29); taxa with longitudinal yellow or orange FW and HW stripes (e.g., L. striata and L. cruciata , pl. 31); as well as species with a yellow transverse FW band and a completely dark HW (e.g., L. ignorata and L. ena , pl. 29). Anomalous wing patterns include those of L. banana , L. fluonia , and L. vulturata (pls. 29, 30)—all of which exhibit an oblique yellow FW stripe—as well as L. minuta (pl. 31), where the orange markings of the FW include an unusual longitudinal streak behind vein Sc, and a quadrate transverse band. Lyces solaris (pl. 30) possesses wings reminiscent of some Scea species (e.g., S. angustimargo , pl. 33). This spectrum of patterns notwithstanding, comprehensive analysis of adult morphology strongly supports the monophyly of a Lyces clade (fig. 283), separate from Josia .

Lyces species can be recognized by the following suite of characters: labial palpus porrect, barely ascending (figs. 320A, B, E– H); Lp2 almost straight, slightly shorter than Lp1; Lp3 bullet shaped, rarely elongate ( L. attenuata ); front and labial palpus uniformly dark brown to blackish brown, rarely (some Patula Group species) with white or orange markings; eye moderately large (fig. 320A– D), postgena relatively wide; FW vein M 1 arising from anterolateral angle of DC (fig. 320I), either touching or almost touching base of radial sector; male and female abdomen elongate, acute distally (especially in males). The male genitalia are variable, but always have an extremely large, membranous BO (e.g., figs. 322A, 323A), narrow, laterally compressed socii, and a relatively long vesica (e.g., fig. 323B, G). In females, the base of the CB usually forms a sclerotized, vaselike structure near its junction with the DB (figs. 323C, 323H, 324C, 324H, 325B, 325G). Internally, this structure is densely spinose.

In Lyces species with longitudinal FW stripes, such as members of the Patula View in CoL Group (pl. 30), it is important to recognize that these stripes fall along the wing more anteriorly than the ones found in Josia View in CoL ( Forbes, 1931; Miller, 1996). In Lyces View in CoL , the majority of the stripe lies within the DC, anterior to the cubitus. The stripe in Josia View in CoL , on the other hand, invariably straddles the cubitus (fig. 336H), its trailing margin extending posteriorly well beyond the anal fold.

REDESCRIPTION: Male. FW length 5 10.5– 23.0 mm. Head (fig. 320A–H): Labial palpus extending almost horizontally, barely ascending; Lp1 relatively long, curved upward, with a wedge-shaped fringe of long scales below; Lp2 straight and fairly wide, shorter than Lp1, usually tightly scaled, without a fringe; Lp3 bullet-shaped; clypeus scaleless; frontal scales short, pointing ventrally from below antennal bases, then horizontally above clypeus; eye moderately large, postgena wide; scales of vertex relatively short, pointing anteriorly; antennal rami thin, variable in length, ranging from moderately long (e.g., L. striata ) to short ( L. gopala ), approximately 12 terminal segments simple; antennal shaft widened from base, tapering distally.

Thorax: Epiphysis moderately wide, approximately two-thirds length of tibia, not extending beyond its apex; tegula moderately long, slightly less than two-thirds length of mesoscutum, ventral angle acute; metathoracic tympanum kettledrum shaped, drum extremely large, occupying almost entire metepimeron; tympanal membrane large, ovoid, oriented horizontally.

Forewing (fig. 320I; pls. 29–31): Broad to slightly elongate; veins Rs 2 –Rs 4 in the pattern [2+3]+4; M 1 arising from anterolateral angle of DC, either touching (e.g., L. aurimutua , L. flavissima ) or almost touching (e.g., L. angulosa , L. ena ) base of radial sector; DC much longer than one-half FW length; veins M 3 and CuA 1 long stalked; wing pattern variable, ground color dark brown to black; most species either with a yellow or orange-yellow transverse band crossing distal margin of DC ( Angulosa Group), or with a longitudinal stripe ( Patula Group).

Hind wing (fig. 320I; pls. 29–31): Broad and full, rounded at apex; M 3 and CuA 1 long stalked; pattern highly variable; ground color dark brown to black, central area concolorous (e.g., L. fluonia , L. banana ), with a yellow or orange central area of various sizes and shapes (e.g., L. attenuata ), or with a longitudinal orange stripe (e.g., L. aurimutua ).

Abdomen: Elongate, acute distally; often uniformly brownish gray to blackish gray above and below; some species with thin, longitudinal whitish (e.g., L. longistria ) or orange ( L. cruciata ) pleural stripes; venter sometimes either with a faint white stripe along midline (e.g., L. longistria ), or completely white (e.g., L. ena , L. cruciata ).

Terminalia (figs. 321A, B, D–G, I, J; 322A–D; 323A, B, D–G, I, J; 324A, B, D, E, F, G, I, J): Tg8 short, roughly quadrate, occasionally tapered posteriorly (e.g., L. fluonia , L. vulturata ), anterior margin simple, posterior margin either with a shallow, wide excavation ( Angulosa Group), or with a deep, U-shaped mesal excavation ( Patula, Eterusialis groups); St8 longer and wider than Tg8, narrowing posteriorly, sometimes markedly so (e.g., L. flavissima ); anterior margin of St8 with a short, acute mesal apodeme (longer in L. flavissima ), posterior margin either simple ( Angulosa Group), or with a deep, often narrow, U-shaped or V-shaped mesal excavation; socii/uncus complex highly variable in shape, narrowly attached to tegumen; uncus usually long, thin and acute at apex, arching downward from base, sometimes with a large, bladelike dorsal flange (e.g., L. angulosa , L. ariaca , L. ena ); socii variable in shape, usually laterally compressed, sometimes acutely tapered (e.g., L. striata , L. attenuata ), occasionally with bristlelike dorsal setae (e.g., L. ariaca ); tegumen wide ( Angulosa Group), or moderately wide to narrow; tegumen taller than vinculum, or rarely shorter ( Eterusialis Group); vinculum narrow; saccus apparently absent, ventral margin of genitalia broadly U-shaped; valva large, completely membranous except for costa and base, BO extremely large, occupying almost entire valva, expanded along outer margin; costa narrow, variable in length, sometimes extremely short ( Eterusialis Group), frequently with an acute (e.g., L. patula , L. ariaca ) or bladelike (e.g., L. cruciata ) process approximately one-half way out; apex bearing a short, sclerotized process of variable shape, usually somewhat truncate (e.g., L. fluonia , L. solaris ), rarely acute ( L. angulosa ); valva bases frequently joining below ventral margin of genitalia to form a W-shaped structure; arms of transtilla wide, pointing sharply downward, sometimes crenulate along dorsal margin (e.g., L. cruciata ); transtillar arms meeting at midline to form a large, deeply concave medial plate, its lateral margins sometimes curled inward; most species with a triangular patch of spicules in manica above transtillar plate, spicules sometimes long and densely arranged (e.g., L. cruciata ); aedeagus extremely wide and bulbous at base, abruptly tapered distally; apex of aedeagus forming a small point; vesica as long as, or longer than aedeagus, variable in shape, always extending upward; vesica wide and arising at a sharp angle from aedeagus (e.g., Angulosa and Eterusialis groups), or long and narrow, curled anteriorly ( Patula Group); vesica with numerous spinelike cornuti, variable in size; often with a single, isolated cornutus at apex of vesica much larger than others (many Patula Group species).

Female. FW length 5 9.5–24.0 mm. Head: Similar to male, except antenna ciliate.

Thorax: Similar to male.

Forewing: Longer and broader than male (pls. 29–31), yellow/orange-yellow maculations wider.

Hind wing: Longer and broader than male, outer margin more rounded, yellow/orange-yellow pattern wider; frenulum comprising two bristles.

Abdomen: Elongate as in male, but wider.

Terminalia (321C, H; 322E; 323C, H; 324C, H): Tg7 long and wide, slightly narrower posteriorly, anterior margin simple, posterior margin with a deep, wide, often poorly defined, U-shaped mesal excavation; posterior two-thirds of Tg7 more sclerotized than anterior third, a transverse depression dividing the two sections; St7 long and wide, equal in length to Tg7, anterior margin simple, posterior margin with a deep, narrow Ushaped mesal excavation; Tg8 completely membranous, membrane fragile; AA extremely thin, either short and delicate ( Angulosa and Eterusialis groups) or moderately long, straight ( Patula Group); A8 pleuron membranous, with a thin, straplike sclerite along anterior margin; PP moderately long, extremely thin; PA small, almost always with a dorsal lobe, sometimes ( Angulosa Group) with ventrolateral margin heavily sclerotized, forming a bladelike lateral flange; area of PVP either completely membranous, or rarely lightly sclerotized; a short, half-moon–shaped antevaginal plate sometimes present (e.g., L. ena , L. auriaca ); DB short, narrow or moderately wide, membranous; CB large, sometimes (e.g., L. flavissima , L. angulosa ) extremely so, roughly ovoid, midsection broadly sclerotized above; base of CB variable, either with a complex series of folds and a large, laterally compressed dorsal appendix ( Angulosa and Eterusialis groups), or narrowed and smoothly sclerotized to form a vaselike constriction ( Patula Group); CB almost always with a large set of internal spines near base (absent in L. angulosa ); signum located ventrally, bird shaped, comprising a wide, transverse sclerite with long, internal spines at each end, central portion of signum protruding from CB membrane; DS arising dorsally from a large, laterally compressed, often sclerotized, appendix, base of DS often curled, pigtail shaped.

DISTRIBUTION: The Eterusialis Group (two species) is strictly South American, whereas the other two Lyces species groups occur from southern Mexico south to Bolivia and southeastern Brazil. The majority of Patula Group species are endemic to Andean cloud forests at elevations between 1500 and 3000 meters—a few have been recorded from 4000 meters. The remaining taxa, such a L. ena ( Angulosa Group) and L. cruciata ( Patula Group), live in lowland forests.

BIOLOGY: Immature stages have been discovered for 10 Lyces species (table 6)— more than a third of the genus. Their hosts seem to contrast with those of Josia . Almost all Lyces hosts belong in the Passiflora subgenus Granadilla , whereas Josia larvae are associated most commonly with Passiflora species currently assigned to the subgenus Decaloba . A similar dichotomy occurs within Heliconius (Smiley, 1985) ; the erato - charitonia species group specializes on Decaloba , whereas the numata -melpomene group is restricted to Granadilla . Plants in these two subgenera show basic differences. The leaves of most Granadilla species are glabrous and thick, whereas those in Plectostemma are pubescent and delicate (see, e.g., Holm-Nielsen et al., 1988). Whether this difference in host-plant utilization between Lyces and Josia is correlated with divergence in plant secondary chemicals within Passiflora , or whether other biological determinants are at play, remains to be investigated. Plant chemistry is only one of the factors determining Passiflora species usage in Heliconius ( Smiley and Wisdom, 1985) , other variables being plant nutritive value and plant-associated predators. It will be interesting to fill in the missing host-plant data for the remainder of the Josiini in search of general patterns.

In contrast to Josia caterpillars (pl. 39L– O), Lyces larvae generally show simpler abdominal coloration—less reticulate and more geometric (pl. 39J, K). Other features of their morphology are the same ( Miller and Otero, 1994; Miller, 1996).

DISCUSSION: In previous treatments of the Dioptinae , the taxa now residing in Lyces were placed in Josia , which then included many more species than are recognized here. Earlier authors ( Prout, 1918; Hering, 1925; Forbes, 1931) were obviously confused by the diversity of wing patterns occurring in the Josiini , and with few other character systems available to them, were unsure how to proceed regarding the group’s taxonomy. Their solution was to create a large diverse genus called Josia , which we now know, was polyphyletic ( Miller, 1996; Miller, Brower and DeSalle, 1997).

The genus name Lyces , prior to now not widely used, is attributed to Walker (1854), who described two species in the genus, angulosa and flavissima . Kirby (1892) later designated angulosa as the type, listing six species of Lyces , only four of which— angulosa , eterusialis , flavissima , and fornax —currently reside in the Josiini . According to the classification proposed here (appendix 2), all four of these are correctly assigned to Lyces . Schaus (1892) added a fifth, describing maera from Brazil, a species that Hering (1925) subsequently moved to its proper position in Erbessa . Prout (1918) placed Lyces in synonymy with Josia , applying the name to his ‘‘Section III’’. Prout recognized 13 species in ‘‘Section III’’, almost all of which belong in Lyces as conceived here. Both Hering (1925) and Bryk (1930) abandoned the use of Lyces in any capacity. According to my concept, Lyces includes 25 species in three species groups.

Unlike some genera of the Dioptinae , where there are numerous undescribed species and few names, I discovered fewer species in Lyces than names. In this paper, seven new junior synonyms are proposed, and one new species is described. Additional study will, in all likelihood, reduce the number of valid species even further. I know of only a handful of undescribed Lyces in collections, most of these represented by a single specimen collected at an exotic locality.

The three species groups defined here are built largely on characters from genitalia. Wing patterns vary dramatically within Lyces , but other features, such as the labial palpi and male antennae, are relatively uniform throughout (fig. 320). By and large, I believe these species groups are robust. At the very least they can act as hypotheses for future testing.