Neotrop Entomol DOI 10.1007/s13744-015-0320-7
SYSTEMATICS, MORPHOLOGY AND PHYSIOLOGY
Revision of the Genus Pseudaethria Schaus (Lepidoptera, Erebidae) LR PINHEIRO1, W MEY2 1
Museu de Zoologia, Univ de São Paulo, São Paulo, SP, Brasil Museum für Naturkunde an der Humboldt-Univ, Leibniz Institut for Evolution and Biodiversity Research, Berlin, Germany
2
Keywords Lectotype designation, Neotropical, new combination Correspondence LR Pinheiro, Museu de Zoologia da Universidade de São Paulo, Avenida Nazaré 481, Ipiranga, 04263-000 São Paulo, SP, Brasil;
[email protected] Edited by André VL Freitas – Unicamp Received 22 September 2014 and accepted 15 July 2015
Abstract The genus Pseudaethria Schaus is revised and redescribed based on morphological characters of male and female adults. Its type species, Pseudaethria cessogae Schaus, was found out to be a junior subjective synonym of Heliura cosmosomodes Dognin. Therefore, the new combination Pseudaethria cosmosomodes is proposed along with another one: Pseudaethria analis Gaede new combination. A lectotype is designated to P. cessogae, which was described from an undetermined number of specimens. The distribution of the species is discussed as well as its systematic placement.
* Sociedade Entomológica do Brasil 2015
Introduction Ctenuchina is a group predominantly Neotropical that counts with approximately 1000 described species, the exact number being hard to know, given the dated catalogues for the group (Zerny 1912, Draudt 1915, 1919). Due to the large diversity and lack of research on moths from this biogeographic region, many of its species and genera are so far known merely from their original descriptions and brief mentions in catalogues. Some were not even illustrated, which increases the problem of the recognition of these taxa. The monotypic genus Pseudaethria Schaus is an example of such problem. It was described to accommodate Pseudaethria cessogae Schaus, a clear-winged moth thought by its author to be related to Aethria Hübner (Erebidae). However, no comparison was presented to distinguish or to stress the morphological traits that associate the two genera. Pseudaethria was cited by Watson et al (1980) and more recently by Ferro et al (2012), and these mere citations— one catalographic and one note of distribution, respectively—are the only mention in the literature of this taxon. During visits to the Museum für Naturkunde an der Humboldt-Universität, Berlin (ZMHB) and to the National Museum of Natural History, Washington, D.C. (USNM), it was
noticed that Pseudaethria cessogae is a junior synonym of Heliura cosmosomodes Dognin, and that Pseudaethria includes a species misplaced in Pseudosphex Hübner (Erebidae). This study aims to provide an unequivocal recognition of Pseudaethria and its species, and to facilitate future studies on the systematic affinities of the genus by the detailed description of its morphological characters.
Material and Methods The material studied is deposited at the following institutions: Natural History Museum, London, England (BMNH); Museu Nacional do Rio de Janeiro, Rio de Janeiro, Brazil (MNRJ); Museu de Zoologia da Universidade de São Paulo, São Paulo, Brazil (MZSP); National Museum of Natural History, Washington, D.C., USA (USNM); and Museum für Naturkunde an der Humboldt-Universität, Berlin, Germany (ZMHB). The morphological study followed standard procedures (Winter 2000). Wing venation was observed directly under the stereomicroscope with the help of a delicate brush. Male and female abdomens were soaked in a solution of 10% KOH at room temperature for approximately 24 h. The genitalia and the abdominal pelts were kept either in small plastic vials with
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glycerine or slide mounted following the procedure described in Robinson (1976), according to the protocol of the institution of origin of each specimen. Illustrations were made with a camera lucida attached to a stereomicroscope. Subsequently, they were processed in ink, following corrections on Adobe Illustrator CS5. The genitalic terminology follows Klots (1970). The classification used in this work follows Jacobson & Weller (2002) and Zahiri et al (2010).
Results and Discussion The observations made resulted in the following taxonomic changes. The lectotype designation is made to enhance stability of nomenclature. Pseudaethria Schaus Pseudaethria Schaus 1924: 10. Type species: Pseudaethria cessogae Schaus 1924: 11 (by monotypy), new synonym of Heliura cosmosomodes Dognin 1916. Diagnosis. This is a very distinctive genus, readily recognized by the yellow coloration predominant on the body, and the almost completely hyaline wings. Legs predominantly yellow with brown distal segments. Forewings with subcostal and radial veins hidden by dark brown scales, the other veins exposed. M2 and M3 not stalked. Hindwings reduced in size, with M3 and CuA1 with a very long stalk, and both anal veins present. First abdominal segments not constricted. Coremata present. Last abdominal tergite with silver scales. Uncus with rounded apex. Transtila membranous. Valvae densely covered by thick setae at the internal side of the tip. Aedeagus with the posterior half turned dorsally. Remarks. Even though the name of this genus suggests a close relationship with Aethria Hübner, a thorough examination of this genus revealed that this assumption is unlikely to be true. The shape of body and wings, and the hindwing venation are different, as well as the general coloration pattern. Besides, genera thought to be close to Aethria, such as Trichura Hübner, Dinia Walker, Corematura Butler, and Riccia Travassos Filho (Erebidae), with body and wing shape and venation similar to that of Aethria, have very different genitalic characters, including dorsal projections arising between the tegumen and uncus and long aedeagus (Travassos Filho 1938, 1954, 1957), not present in Pseudaethria. An extensive survey was made across potentially closely related genera, both in the literature (Butler 1877, Hampson 1898–1901, 1915, Druce 1881–1900, Schaus 1892, Rothschild 1912, 1913, Draudt 1917) and in collections consulted (the latter limited to external characters and overall resemblance), but no putative closely related genus could be
found. Many ctenuchine genera include species with external resemblance to Pseudaethria, which is not unexpected in a taxon full of wasp mimics, but the phylogenetic affinities of this taxon remain to be discovered. Pseudaethria cosmosomodes (Dognin), new combination (Figs 1, 2, 4, 5, 6, 7, 8, 9, 10, 11, 12, 13) Heliura cosmosomodes Dognin 1916: 4. Holotype ♂, by monotypy. BRAZIL, Saô [São] Paulo (Donckier), with six labels: “Type No 30783 U.S.N.M.”; “Heliura cosmosomodes type ♂ Dogn. Hmpsn 7.16”; “Dognin collection”; “Heliura sp. nat. in B.M.”; “São Paulo Brésil”; and “Kb-Dia-Nr. 1572 B. Kreusel dok.” (USNM) [examined]. Pseudaethria cessogae Schaus 1924: 11. Lectotype hereby designated, male. BRAZIL [Santa Catarina], Joinville (from Julius Arp collection), with six labels: “Type No. 25888 U.S.N.M.”; “Pseudaethria cessogae type Schs”; “312”; “J.ville coll. J. Arp”; “Collection WmSchaus”; and “Kb-Dia-Nr. 1786 B. Kreusel dok.” (USNM) [examined]. New synonym. Diagnosis ♂ and ♀. This species is characterized by the following external traits: subapical antennal segments white, palpi and thoracic segments entirely yellow, including the tegulae and patagia, T1–7 and S2–6 also yellow. Distinctive genitalic traits include the symmetrical valvae, rounded saccus and very short aedeagus. Redescription ♂ and ♀. Head. Proboscis light brown. Labial palpi, frons, vertex, occipital and genal areas of the head yellow. Paraocular area glabrous, tegument light-colored. Scape, pedicel, and the internal surface of the first flagellomere yellow. Second and third flagellomeres brown with a few white scales on their internal surface. Rest of the antennae dark brown, except for the subapical flagellomeres, white dorsally. Thorax. Entirely yellow. Legs predominantly yellow, foretibiae and tarsi of all pairs of legs brown. Forewings. Proximal area yellow, margins and veins dark brown. Narrow line of brown scales forming a post-discal bar. Rest of the surface of forewings covered by tiny setae. Venation: R1 diverging before the discal cell; R2 diverging near the apex, right before R5, which diverges more or less half the way between R2 and R3 +R4. M2 and M3 without a common stalk. Distance between CuA1 and CuA2 approximately the same as the distance between the former and M3 (Fig 4). Hindwings. Proximal area yellow, margins and veins dark brown. Most of the rest of the hindwings’ surface covered by tiny setae, a small region around the stalk of vein Cu completely hyaline. Venation: M3 and CuA1 with a long stalk, M2 diverging separately from it. 1A and 2A present (Fig 4). Abdomen. T1–7 yellow; T8 silver, posterior margin with long brown scales. S2–6 yellow; S7–8 brown, S8 with long brown scales laterally.
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Figs 1–5 Type specimens, wing venation, and androconia of 1 Heliura cosmosomodes n. comb., holotype; 2 Pseudaethria cessogae, lectotype; 3 Chrysostola analis, holotype; 4 wing venation of Pseudaethria cosmosomodes n. comb. (upper part) and Pseudaethria analis n. comb. (lower part); 5 detail of coremata (c) and lateral pouch (p).
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Figs 6–9 Pseudaethria cosmosomodes n. comb., male abdomen and genitalia. 6 Detail of T8 of P. cosmosomodes showing the sacular projections on its anterior margin; 7 genitalia in lateral view, aedeagus removed; 8 same, posterior view; 9 aedeagus, lateral view.
Proximal segments not constricted. Tergo-pleural slits as long as T1. Males: T8 sclerotized, approximately rectangular, with two very thin sacular projections
(SP) with approximately the same length of the tergite. Coremata (C) present in intersegmental membrane 7–8, beside a pouch (P), also probably glandular, with many
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long scales. Females: T8 considerably larger and more heavily sclerotized than the other tergites. S8 also larger and more sclerotized than the remaining sternites, more or less square, in contrast with the rectangular S2–7. Male genitalia. Tegumen considerably wider than the vinculum, with a dorsal triangular indentation on its anterior margin. Limits between tegumen and vinculum membranous. Saccus little developed, rounded. Uncus large, proximal half compressed laterally, distal half bent ventrally. Sparse setae covering the dorsal surface of the uncus. Apex not compressed, rounded. Scaphium heavily sclerotized, forming an H-shaped structure. Juxta sclerotized, shield-like. Valvae long, proximal half less sclerotized and wider than the distal half. Apex densely covered by thick setae, contrasting with the few delicate setae on the ventral surface of the proximal half. Aedeagus very short and pointed. Coecun undeveloped, vesica with minute cornuti. Female genitalia. T8 membranous dorsally and lightly sclerotized laterally. Pheromone glands digitiform, slightly longer than the papilla anales. Anterior apodemes with approximately one third of the length of the posterior apodemes. Antevaginal lamella heavily sclerotized, densely covered by brown scales. Sclerotization of the antevaginal lamella forming lateral flaps fusing with the post-vaginal lamella, ventrally membranous. Ostium slightly turned to the right. Ductus bursae shorter than corpus bursae, sclerotized ventrally and membranous on the remaining surfaces. Corpus bursae more or less rounded, membranous, with a small and rounded patch of minute signa. Distribution. To the present knowledge, this species occurs in the humid Atlantic forest from southeast and south Brazil, and the interior semidecidual forests from Paraguay. Etymology. The specific epithet cosmosomodes seems to come from the genus Cosmosoma Hübner (Erebidae), which comprises many species bearing superficial resemblance to P. cosmosomodes n. comb. Material examined. (10 ♂ and 3 ♀). BRAZIL. Espírito Santo, Santa Tereza, [year of] 1967, Paulo Elias col., 1 male (MNRJ); Minas Gerais, Nova Lima, 850 m, 8.x.1985, V. O. Becker col., col. Becker 63078, 1 female (USNM); Santa Catarina, no precise locality or date, F. Johnson col., 1 male (USNM); Hansa Humboldt, viii.1932, A. Maller, 1 male (BMNH); idem, xi.1936, A. Maller, 1 female (BMNH); Jaraguá do Sul, ix.1934, F. H. Hoffmann, Rothschild Bequest B. M. 1939–1, 1 male (BMNH); idem, viii.1935, A. Maller, 1 male (BMNH); Jville [Joinville], coll. J. Arp, [catalog number] 956, 1 male (MZSP); idem, no collector or date, [catalog number] 957, 1 male (MZSP); idem, no collector or date, [catalog number] 71/735, 1 male (MZSP); São Paulo,
Campos do Jordão (Lefèvre), 1200 m, 13–15.ii.1953, Travassos Filho & Travassos col., 1 male (MZSP); idem, 13–20.ix.1952, d’Almeida, L.T.F. [Lauro Travassos Filho] and Pd. Pereira col., 1 male (MZSP); PARAGUAY, Mbororo, Villarica, 1932, F. Hoffmanns, 1 female (BMNH). Remarks. Dognin (1916) mentioned that Pseudaethria cosmosomodes n. comb. has the aspect of a Cosmosoma. At the same time, he did not explain why he described the species in Heliura Butler 1876, a ctenuchine genus mainly composed of fully wing-scaled moths with no resemblance to his species whatsoever (the type species of Heliura, H. rhodophila, was illustrated in Hampson 1898, p. 483, fig. 267, and Draudt 1917, plate 24 row f). Pseudaethria cessogae was described from an undetermined number of males. Schaus (1924) mentioned “Type Cat. N° 25888”, but because these numbers could also apply to groups of individuals (Robert Poole, personal communication), it is not possible to know whether it really corresponded to the single individual found at the USNM (including the main collection). Schaus (1924) also mentioned that his genus was close to Aethria, but without any comment on which of its species it is close. A thorough search across Aethria showed that there are no species particularly close to P. cosmosomodes n. comb. Pseudaethria analis (Gaede), new combination (Figs 3, 4, 14, 15). Chrysostola analis Gaede 1926: 128. Holotype ♂, by monotypy. BOLIVIA, Rio Songo, 1200 m (Garlepp), with eight labels: “Type”; “Rio Songo (1200 m) Bolivia (Yungas) 1895–6 Garlepp”; “Coll. Staudinger”; “825b 44.”; “near postica neck (Hps.)”; “Chrysostola analis Type Gaede”; “Kb-Dia-Nr. 87 B. Kreusel dok.”, and “genitalia slide Mey 81/14” (ZMHB) [examined]. Diagnosis ♂. This species differs from Pseudaethria cosmosomodes n. comb. mainly in the coloration of the head and patagia, dark brown instead of yellow, as in that species. Pseudaethria analis n. comb. is also distinct in having the distal third of the antennae dark brown, the tips of the palpi brown, and T7, as well as the posterior half of S7, also dark brown scaled. There are also genitalic differences in the asymmetrical valvae, pointy saccus, and the relatively long aedeagus. Redescription ♂. Head. Proboscis light brown. Labial palpi yellow with brown tips. Frons and vertex dark brown. Genal area yellow. Occipital area brown, with some bluish scales. Paraocular area glabrous, tegumen light colored. Scape, pedicel, and flagellomeres dark brown. Rest of the antennae dark brown, except for the subapical flagellomeres, white dorsally. Thorax. Patagia brown, anterior margin with iridescent bluish scales. Tegulae, yellow, with some scattered brown scales. Mesothorax,
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Figs 10–13 Pseudaethria cosmosomodes n. comb., female abdomen and genitalia. 10 Abdomen; 11 female genitalia, lateral view; 12 same, dorsal view; 13 same, ventral view.
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metathorax, episterna, and epimera yellow. Legs predominantly yellow, tarsi of mid and hindlegs brown; hindtibia with a brown streak on dorsal side. Forelegs missing in the holotype. Forewings. Proximal area yellow, margins and veins dark brown. Narrow line of brown scales forming a post-discal bar. Rest of the surface of forewings covered by tiny setae. Venation: R2 branching after R5; M2 and M3 not stalked. Three lines of scales resembling veins occurring in cells R 4-R5 , R5 -M 1, and M1-M2 (Fig 4). Hindwings. Proximal area yellow, margins and veins dark brown. Most of the rest of the hindwings’ surface covered by tiny setae, a small region around the stalk of vein Cu completely hyaline. Venation: M 2 branching at a small distance from the stalk of M3 and CuA1. 1A and 2A running parallel, very close to each other (Fig 4). Abdomen. T1–6 yellow; T7 yellow on the anterior half, and brown on the posterior half, except for the posterior margin, with silver scales. T8 brown with silver scales dorsally. S2–6 yellow; S7 yellow anteriorly and brown posteriorly; S8 entirely brown. Proximal segments not constricted. Tergo-pleural slits as long as T1. Males: T8 sclerotized, sub-rectangular, with two very thin secular projections with approximately the same length of the tergite. Coremata present in intersegmen-
tal membrane 7–8, beside a pouch, also probably glandular, with a bundle of short, club-like androconial scales. Male genitalia. Tegumen considerably wider than vinculum, with a dorsal triangular indentation on its anterior margin. Limits between tegumen and vinculum clear, membranous. Saccus little developed, triangular. Uncus large, proximal half compressed laterally, distal part turned ventrally. Sparse setae covering the dorsal surface of the uncus. Apex not compressed, rounded. Juxta composed of a pair of elongated ribbons. Valvae long, proximal half less sclerotized and wider than the distal half, which ends with an acute tip. Apex densely covered by thick setae, contrasting with the few delicate setae situated on the ventral surface of the proximal half; valvae asymmetric, left one longest with inwardly curved tip, right valve bifid. Aedeagus moderately long and pointed. Coecum undeveloped, vesica with minute cornuti. Etymology. The specific epithet is probably an allusion to the peculiar silver scales on the last abdominal tergite. Remarks. According to Gaede (1926), this species is similar to Pseudosphex discoplaga (Schaus). The latter was
Figs 14–15 Pseudaethria analis n. comb., male genitalia. 14 Genitalia in posterior view, aedeagus removed; 15 aedeagus, lateral view.
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examined, and the resemblance is probably analogous, as both species (as well as many others in Pseudosphex and other genera, such as Sphecosoma Butler and Isanthrene Hübner) (Erebidae) share characters related to wasp resemblance, e.g., hyaline wings, body shape, and coloration. Acknowledgments The first author is grateful to the curators and technicians who granted her access to the collections and facilities under their care: Alessandro Giusti, Geoff Martin, and Martin Honey (BMNH); Miguel Monné and Alexandre Soares (MNRJ); Marcelo Duarte (MZSP); and Donald Harvey (USNM). The manuscript was improved by suggestions and comments of Simeão Moraes and three anonymous reviewers. This research was funded by Fapesp (grants 2009/11159-5 and 2012/02444-0).
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