Livendula Hall

Hall, Jason P. W., 2007, Two new genera in the Nymphidiina (Lepidoptera: Riodinidae: Nymphidiini), Zootaxa 1415, pp. 35-42 : 35-39

Livendula Hall , n. gen.

( Figs. 1 A,B; 3; 5)

Type species: Lemonias huebneri Butler, 1867 .

Etymology: The name of this genus is based on the medieval Latin word for lavender, and refers to the almost universal occurrence of this color on the dorsal wings of males in this genus.

Description: MALE: Forewing length 11–19 mm. Wing shape: Both wings typically compact; forewing costa approximately straight, distal margin slightly convex, anal margin straight; hindwing rounded. Ve na t io n: As in Adelotypa (see Stichel (1910–11), under Echenais Hübner , [1819]), four forewing radial veins. Dorsal surface: Ground color of both wings brown or rufous brown; three black spots in discal cell of both wings and two at base of cell Cu2, an additional spot at base of cell Rs on hindwing, all of these encircled with lavender scaling in most species; jagged postdiscal band of black spots on both wings usually associated with variably prominent proximal and distal lavender scaling, a broad white postdiscal band present on both wings in one species ( L. amasis (Hewitson, 1870)) , a variably broad distal white patch on hindwing extending to distal margin present in all derived species; dark submarginal spots faint to absent within hindwing white areas, but prominent elsewhere and almost always encircled with lavender scaling; forewing fringe either brown or with variably prominent white elements in cells Cu2, M3, M1 and R4+5, hindwing fringe either brown or checkered brown and white. Ventral surface: Differs from dorsal surface in following ways: Ground color of both wings pale to medium brown; dorsal lavender scaling replaced by whitish scaling of somewhat similar extent; whitish scaling surrounds basal spots in all species, all black spots more prominent.

Head: Ventral surface of labial palpi generally a mixture of pale white or pale yellow-brown and dark scaling, dorsal surface dark brown, second and third segments elongate; eyes brown and bare, with white or pale brown scaling at margins; frons brown with a thin vertical band of white or pale brown scaling laterally and a broad horizontal pale band ventrally; antennal length approximately 60 to 70% of forewing length, segments brown with an elongate band of white lateral scaling, nudum along inner ventral margin of shaft variably narrow and discontinuous; clubs black, tips orange brown.

Body: Dorsal surface of thorax and abdomen dark brown, ventral surface pale brown to whitish; eighth sternite a simple rectangle with a smoothly rounded posterior margin; all legs pale brown or whitish, tarsus of foreleg unimerous, coxa of medium length for family, midleg and hindleg with a tibial spur and a group of spines at inner distal tip of tibia and spines along inner margin of all tarsal segments.

Genitalia ( Fig. 3 ): Uncus in lateral view approximately rectangular and typically more posteriorly elongate ventrally, with a broad and shallow dorsal indentation medially, lateral “window” dorsally separated in all species except L. balista (Hewitson, 1863) ; falces broad; vinculum slightly sinuate and approximately even in width, saccus elongate; valvae in lateral view approximately rectangular, with a rounded lateral bulge basally, an angular anteroventral margin, a dorsal desclerotized region medially, and two inwardly and upwardly curved, rounded processes posteriorly that are most readily visible as separate valve elements in ventral view, upper process broader and slightly more posteriorly elongate, valvae fused dorsally near tips with a broad and slightly medially raised sclerotized band; aedeagus very long and narrow, with pointed posterior tip opening dorsally or to right, and rounded anterior end opening anteroventrally and slightly to right; everted vesica very long and narrow, with a large and narrow “pitch-fork”-shaped cornutus positioned in middle section of vesica consisting of a long and thick dorsal spine and a shorter and narrower ventral spine attached to a more weakly sclerotized, flattened and approximately rectangular pad, and a small membranous tube exiting laterally from main stem of vesica near anterior tip of cornutus; very long and narrow, strap-like pedicel joins aedeagus in its anterior half.

FEMALE: Differs externally from male in following ways: Forewing length 12–20 mm; both wings more rounded; both wing surfaces paler plain brown, dorsal lavender scaling confined to submargin in most species, with derived species only having lavender scaling encircling basal spots; white hindwing patch in female not necessarily correlated with its presence in male, and when present smaller in extent than in male; one species with a white postdiscal band on forewing ( L. violacea (Butler, 1867)) , another with a white postdiscal band on both wings ( L. amasis ).

Head: Second and third palpal segments slightly more elongate.

Body: Foreleg with spines at inner distal tip of tibia and tarsal segments one to four.

Genitalia ( Fig. 5 ): Corpus bursae elongate and generally narrow, with a pair of medially to posteriorly positioned, variably small, spine-like signa in all species except one ( L. epixanthe (Stichel, 1911)) , which typically have an elongate and convex outer margin at wall of corpus; ductus bursae membranous and very elongate (often of equal length to entire abdomen), with an enlarged “half-moon”-shaped section in posterior half that is hardened and slightly darkened by a dense layer of tiny spines on inner surface; posterior portion of ductus bursae with a variably elongate, straight, sclerotized section, which is entirely sclerotized in vicinity of ostium bursae, but only sclerotized ventrally and laterally for most of its length; membranous ductus seminalis exits ductus bursae dorsally, in vicinity of sclerotized section; ostium bursae a large, triangular, sclerotized plate, with posterior apical portion variably projecting away from abdominal surface to form a sclerotized cone, ostium opening round and posteriorly positioned; papillae anales approximately rectangular.

History of classification: The first species to be described of those treated here in the new genus Livendula was Papilio aristus Stoll, 1790 . All of the remaining species were described in Lemonias Hübner, 1807 , Echenais and Adelotypa , almost entirely in that chronological order, predominantly by Hewitson, Butler and Stichel. It is a reflection of how common and widespread most Livendula species are that seven had been described by the time of Bates’ early catalog (1868). All of the remainder, except one, the common yet overlooked species L. jasonhalli (Brévignon & Gallard, 1999) , had been described by the time of Stichel’s (1910– 11) first catalog. Stichel (1910–11) treated all of the then described species except one in a single species group of Echenais , but he additionally included in that group species from several other genera. He erroneously treated L. amasis as a synonym of Lemonias egaensis (Butler, 1867) , presumably because of the superficial similarity of the females. The arrangement in his second catalog (1930–31) was very similar, but L. amasis was transferred to Echenais as a good species, and L. epixanthe was also raised to species status. Stichel recognized seven and nine species, respectively, in these two catalogs. Through the nomenclatural action of Hemming (1943), all the species previously treated in Echenais became part of Adelotypa . Most recently, Callaghan & Lamas (2004) recognized thirteen species in the Adelotypa aristus group, unjustifiably treating asemna Stichel, 1910, and mollis Butler, 1877, as full species (Hall, in prep.). Eleven species are here recognized in Livendula (n. combs.) and transferred from Adelotypa : amasis (Hewitson, 1870) , aminias (Hewitson, 1863), aristus (Stoll, 1790) , balista (Hewitson, 1863) , epixanthe (Stichel, 1911) , huebneri (Butler, 1867) , jasonhalli (Brévignon & Gallard, 1999) , leucocyana (Geyer, 1837), leucophaea (Hübner, [1821]), pauxilla (Stichel, 1911), and violacea (Butler, 1867) . A complete synonymic list of all Livendula names is given in my revision of the subtribe Nymphidiina (Hall, in prep.).

Diagnosis and systematic position: All members of Livendula , as well as Minotauros described below, possess dorsally fused valve tips and a ventrally positioned spiracle on male abdominal segment three, placing them in the nymphidiine subtribe Nymphidiina. Livendula contains many confusingly similar species, and the genital morphology, particularly of males, is very uniform across the group. The genus is characterized by the presence of variably extensive lavender scaling on the dorsal wings of males in all species except the basal L. balista , the possession in the male genitalia of all species of an extraordinarily long aedeagus and associated pedicel, a unique “pitch-fork”-shaped aedeagal cornutus, a very long saccus, and a unique valve shape, and the possession in the female genitalia of all species of a hardened “half-moon”-shaped swelling near the middle of the ductus bursae, presumably the location in which the male aedeagal cornutus fits during copulation.

Livendula species, as well as those of Minotauros , lack the characters possessed by members of the most derived Nymphidiina clades, such as asymmetrical valve tip lengths and an ancillary spine at the base of the aedeagal cornutus, placing the genus in the basal half of the subtribe (Hall, in prep.). There are no obvious synapomorphies uniting Livendula with any other genus, but the fact that the aedeagal vesica of species in both genera has an unusual membranous extension from the main tube in the vicinity of the cornuti, combined with similarities in wing pattern, suggests that Livendula and Minotauros are very close relatives.

Distribution: Livendula is entirely confined to South America east of the Andes, and ranges from Venezuela to Bolivia, Brazil, and the Guianas. It is possible that all eleven species might be found sympatrically in certain regions of western Amazonia.

Figures

FIGURES 1 – 2. Adults (dorsal surface on left, ventral surface on right). 1 A. Livendula huebneri male, nr. Cacaulândia, S. W. Brazil [Ro] (USNM). 1 B. Livendula huebneri female, nr. Cacaulândia, S. W. Brazil [Ro] (USNM). 2 A. Minotauros lampros male, Estación Científica Yasuní, E. Ecuador (JHKW). 2 B. Minotauros lampros female, Estación Científica Yasuní, E. Ecuador (JHKW).

FIGURES 3 – 4. Male genitalia. 3. Livendula huebneri, Matoury, French Guiana (USNM), with genitalia and cornutus (bottom) in lateral view, and valvae in ventral view (top). 4. Minotauros lampros, upper Río Tiputini, E. Ecuador (JHKW), with genitalia in lateral view, valvae in ventral view (left) and uncus / tegumen in dorsal view (right). Scale bar = 1 mm.

FIGURES 5 – 6. Female genitalia in dorsal view. 5. Livendula huebneri, Parque Nacional Manu, Pakitza, Peru (USNM). 6. Minotauros lampros, Mitú, Colombia (AME). Scale bar = 1 mm.