ZOOTAXAISSN 1175-5326 (print edition)

ISSN 1175-5334 (online edition)Copyright © 2013 Magnolia Press

Zootaxa 3682 (1): 001–044 www.mapress.com/zootaxa/ Article

http://dx.doi.org/10.11646/zootaxa.3682.1.1http://zoobank.org/urn:lsid:zoobank.org:pub:6D48EE4D-05EA-4A2D-88A6-6FBBEA9FD737

A Revision of the New World genus Lepidophora Westwood, 1835 (Diptera, Bombyliidae, Ecliminae) with a key to the species

PAULA FERNANDA MOTTA RODRIGUES1,2,3 & CARLOS JOSÉ EINICKER LAMAS1,4

1Museu de Zoologia da Universidade de São Paulo. Avenida Nazaré, 481 Ipiranga, 04263-000. São Paulo, SP, Brazil. 2Programa de Pós-Graduação em Ciências Biológicas (Zoologia) do Instituto de Biociências, Universidade de São Paulo.3Coordenação de Aperfeiçoamento de Pessoal de Nível Superior (CAPES), fellow.4Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq), fellow.E-mails: pfmotta@usp.br; einicker@usp.br

Table of contents

Abstract . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1Material and Methods . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3Taxonomy . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4Lepidophora Westwood, 1835 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4Key to species of Lepidophora Westwood . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8Lepidophora acroleuca Painter, 1930 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9Lepidophora culiciformis Walker, 1850 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13Lepidophora cuneata Painter, 1939 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18Lepidophora lutea Painter, 1962 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 26Lepidophora secutor Walker, 1857 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 30Lepidophora trypoxylona Hall, 1981 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 34Lepidophora vetusta Walker, 1857 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 37Acknowledgments . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 41References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 44

Abstract

The bombyliid genus Lepidophora has a distribution restricted to the New World. The genus is composed by eight species, which are revised and redescribed herein: five restricted to the Neotropical region (L. acroleuca Painter, 1930, L. culici-formis Walker, 1850, L. cuneata Painter, 1939, L. secutor Walker, 1857 and L. trypoxylona Hall, 1981), two restricted to the Nearctic region (L. lepidocera (Wiedemann, 1828) and L. lutea Painter, 1962) and one species occurring in Nearctic and Neotropical regions (L. vetusta Walker, 1857). The main characters of the external morphology of adults and male and female terminalia are illustrated. An key to species is presented.

Key words: bee flies; Lepidophora; New World; taxonomy.

Introduction

Westwood (1835) erected the genus Lepidophora with Ploas aegeriiformis Gray, 1832 the type species by monotypy. The genus was initially placed in subfamily Toxophorinae. In that paper, Westwood suggested the similarity between Ploas aegeriiformis and Toxophora lepidocera Wiedemann, 1828. Subsequent authors adopted that synonymy, although it was not formalized in Westwood’s (1835) manuscript.

Becker (1913) erected the subfamily Cylleniinae to include Amictus Wiedemann, Cyllenia Latreille, CyrtomyiaBigot, Eclimus Loew, Epibates Osten Sacken, Henica Macquart, Toxophora Meigen, Sphenoidoptera Williston and Lepidophora, based on similar habitus and wing venation. These genera were previously placed in Toxophorinae.

Accepted by Neal Evenhuis: 26 Apr. 2013; published: 26 Jun. 2013 1

Bezzi (1924) did not agree with the placement of Lepidophora and Toxophora in Cylleniinae, and placed them back in Toxophorinae.

The first revision of Lepidophora was presented by Painter (1925) and included a new record of the species Lepidophora appendiculata Macquart, 1846 at the type locality (Texas, United States). Painter redescribed, included notes of occurrence and presented an identification key to four of the five species so far known: L. appendiculata, L. lepidocera (Wiedemann, 1828), L. vetusta Walker, 1857 and L. secutor Walker, 1857. In that paper, the antennae of L. secutor and L. lepidocera were illustrated to show the difference between the lengths of the scales on the postpedicel of these species. L. culiciformis Walker, 1850 was not revised because there were no further records of it since the original description. However, the author reproduced the original description to make it more accessible.

Painter (1930) added a new species to the genus, Lepidophora acroleuca, collected during his studies in the Republic of Honduras.

Painter (1939) described another species for the genus, Lepidophora cuneata, studying specimens from the collection of “Deutsches Entomologisches Institut”, Berlin-Dahlem. The author presented illustrations of the wing and the pupal case, but did not describe the latter.

Paramonov (1949) revised the species of Lepidophora and presented a catalog and an identification key to adults. The species revised in that study were: L. acroleuca, L. appendiculata, L. culiciformis, L. cuneata, L. lepidocera, L. secutor and L. vetusta. The species L. lepidocera and L. vetusta were redescribed. With regard to the others, the author only reproduced the original descriptions or Painter’s (1925) redescriptions. Paramonov also described a new species, Lepidophora austeni, from Uganda, the only Afrotropical species of the genus and the first with record outside the New World. L. austeni differs from other Lepidophora species in having three submarginal cells (instead of two) and the wing completely dark, but Paramonov did not consider these characters relevant to justify the establishment of a new genus. Paramonov also observed that the genus Lepidophora was more closely related to the genus Toxophora, both of which he placed in Toxophorinae.

Painter (in Painter & Painter 1962) added one more new species to the genus, Lepidophora lutea, during his examination of types of Bombyliidae housed in American and European museums. Painter and Painter (1962) observed three distinct species with records from the United States. L. vetusta, the most different of these, was recorded from Texas; L. lutea occurred in the northern United States; while L. lepidocera occurs in the south. The authors also stated that the two latter species may, with future study, be shown to be subspecies. Also as a result of that study, the species L. appendiculata was considered a new synonym of L. lepidocera. Painter noted two significant errors in Painter´s (1925) revision of Lepidophora where he redescribed L. lepidocera as L. appendiculata and L. lutea as L. lepidocera.

François (1964) observed that despite the common features of L. austeni in relation to other species of Lepidophora (scales on the wing, palpus formed by two palpomeres, diminutive style and four posterior cells), the differences between them were much more significant. Based on that observation, François erected the genus Palintonus to include the single Afrotropical species, P. austeni (Paramonov, 1949). A complete redescription of the species, with illustrations of the wing, abdomen and male terminalia was presented. Following that study, the genus Lepidophora was once again restricted to the New World.

Hull (1973) redescribed the genus Lepidophora, indicated that their immatures live in the nests of wasps, and presented a list of known hosts of L. lepidocera. The author initially erected the tribe Lepidophorini within Toxophorinae and included Lepidophora along with the genera Marmasoma White, Cyrtomyia Bigot and Palintonus François. These genera are distinguished from Toxophora Meigen (placed within tribe Toxophorini) by the subglobular head, very swollen occiput and the spikelike occipital bristles. After preparation of Hull´s (1973) work was completed, he revised the subfamilies Toxophorinae and Bombyliinae on the basis of pupal characters and placed the tribe Lepidophorini within Bombyliinae.

Painter and Painter (1974) examined the type-series of Neotropical species of Lepidophora housed in American and European museums. The authors presented redescriptions of the external morphology of L. culiciformis, L. cuneata and L. secutor and an identification key to the Lepidophora species of South America. Painter and Painter observed that the genera Lepidophora and Cyrtomyia were morphologically very similar in having the pronotum armed with abundant thorns.

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Hall (1976), in his study on the Bombyliidae of Chile, proposed that the genus Cyrtomyia was the Chilean counterpart of Lepidophora, since the former is located just to the east of the Andes while the latter is found just west.

Hall (1981) described the new species Lepidophora trypoxylona from an observation of specimens previously identified as L. vetusta reared from nests of the wasp Trypoxylon (Trypargilum) tenoctitlan Richards. Descriptions of adults and pupal case, as well as illustrations of the latter, were presented. Hall considered eight species in the genus Lepidophora, all restricted to the New World: five restricted to the Neotropical region (L. acroleuca, L. culiciformis, L. cuneata, L. secutor and L. trypoxylona), two restricted to the Nearctic region (L. lepidocera and L. lutea) and one species occurring in both the Nearctic and Neotropical regions (L. vetusta).

Theodor (1983) described and illustrated the male and female terminalia of Bombyliidae and positioned the genera of Lepidophorini sensu Hull (1973) in the subfamily Toxophorinae, but showed reluctance with this placement, as these genera differed from Toxophora in the morphology of the sand chamber, the spermathecae and male terminalia.

Greathead (1988) redescribed and compared the genus Tillyardomyia Tonnoir with other more closely related genera of Bombyliidae. As a result, he concluded that Tillyardomyia belongs to subfamily Ecliminae and redefined this subfamily to also include the genera Cyrtomyia, Eclimus, Lepidophora, Marmasoma, Palintonus and Thevenetimyia. In addition, Greathead conducted a phenetic analysis based on 22 morphological characters. That analysis confirmed the intuitive division of the subfamily into three groups of genera and their geographical distribution. The phenogram of affinities proposed by Greathead showed a close relationship between Lepidophora and Cyrtomyia and the genus Palintonus appeared related to these genera.

Yeates (1994) presented a phylogenetic hypothesis for Bombyliidae based on morphological characters and proposed a classification for the family based on cladistic analysis. The results showed that the scales on the wing and the broad pronotum evolved independently in the genera Lepidophora and Toxophora confirming the relationship suggested by Hull (1973) and Greathead (1988), but unlike the latter, Yeates reduced the Ecliminae to a tribe within in Bombyliinae (and included Lepidophora in that tribe), since the Eclimini appeared as a “more distal” member the Bombyliinae. The synapomorphies that support Lepidophora are: 1) pedicel extremely elongate (often more than five times longer than wide); 2) eyes of the male relatively wide apart; 3) tentorial arm very narrow; and 4) enlarged prothorax. Yeates obtained two different resolutions for the placement of the Eclimini genera and both resolutions revealed a sister-group relationship between the genera Lepidophora and Cyrtomyia. In Yeates’s (1994) study, he included in the Eclimini the genera Lepidophora, Cyrtomyia, Paratoxophora Engel (which was for the first time considered part of this taxon), Thevenetimyia, Eclimus and Marmasoma. However, Yeates pointed out that the inclusion in a future analysis of representatives of the genera Palintonus, Tillyardomyia and a large taxonomic and geographical sample of Thevenetimyia may resolve the relationships between the genera and species of the tribe.

Yeates and Greathead (1997) described two kinds of cleptoparasitism in Bombyliidae, one of them present only in Lepidophora where the bee fly larva feeds only on the host larva’s provisions, causing the host larva to starve.

Evenhuis and Greathead (1999) returned the subfamily status to Ecliminae, based on morphological differences and followed this placement in their keys in Greathead and Evenhuis (2001), where they provided a history of the classification of the subfamily. The Ecliminae are elongated, usually without bristles and have a modified sand chamber in the female terminalia. The subfamily includes nine genera: Alepidophora Cockerell (only known from fossils), Cyrtomyia, Eclimus, Lepidophora, Marmasoma¸ Palintonus, Paratoxophora, Thevenetimyia and Tillyardomyia, with three of them (Cyrtomyia, Lepidophora and Thevenetimyia) occurring in the Neotropical region. Evenhuis and Greathead followed the Hall’s (1981) concept, retaining the New World genus Lepidophora with eight species.

Lamas and Lopes (2004) described and illustrated the pupal case of L. culiciformis, expanded the known geographic record for the species and presented a list of hosts.

Material and Methods

The studied material belongs to the following institutions: American Museum of Natural History (AMNH), New York, United States; The Natural History Museum (BMNH), London, England; California Academy of Sciences

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(CAS), California, United States; Coleção Entomológica Padre Jesus Santiago Moure (DZUP), Departamento de Zoologia, Universidade Federal do Paraná, Paraná, Brazil; Instituto Nacional de Pesquisas da Amazônia (INPA), Manaus, Brazil; Museu Nacional, Universidade Federal do Rio de Janeiro (MNRJ), Rio de Janeiro, Brazil; Museu Paraense Emílio Goeldi (MPEG), Pará, Brazil; Museu de Zoologia da Universidade de São Paulo (MZUSP), São Paulo, Brazil; Forschungsinstitut und Museum Senckenberg (SMFD), Frankfurt, Germany; University Museum, University of Oxford (UMO), Oxford, England; Smithsonian National Museum of Natural History (USNM), Washington, United States; Museum für Naturkunde (ZMHB), Humboldt-Universität, Berlin, Germany.

Genitalia were macerated in 10% KOH at room temperature for one day to remove soft tissue, then rinsed in distilled water, dehydrated in an increasing ethanol series (30%, 50%, 70% and 95%) and dissected in glycerin. Genitalia preparations were placed in glycerin in a genitalia vial mounted on the pin beneath the specimen.

Terminology follows Cumming and Wood (2009), except for the pleural and genitalic morphology where we follow Yeates (1994), for wings we follow Greathead and Evenhuis (2001), and for antennae we follow Stuckenberg (1999).

Species distribution data were compiled in a Microsoft Office Excel spreadsheet using specimen labels and literature data. Records without geographic coordinates were georeferenced using online gazetteers. The geographical coordinates, used in the distribution map, are presented in brackets, after the locality of each specimen, in the list of other material examined. The distribution map was produced using ARCVIEW 3.2 (ESRI, Redlands, CA).

Taxonomy

Lepidophora Westwood, 1835

Lepidophora Westwood, 1835: 447. Type species: Ploas aegeriiformis Gray, 1832 [= Toxophora lepidocera Wiedemann, 1828], designated by monotypy; Macquart, 1840: 119; Guérin-Ménéville, 1844: 538; Macquart, 1846: 350; Bigot, 1857: 292; Walker, 1857: 145; Bigot, 1858: 585; Osten Sacken, 1858: 25, 44; Loew, 1860: 247, 270; Osten Sacken, 1877: 229, 265; Osten Sacken, 1878: 94; Osten Sacken, 1886: 76; Osten Sacken, 1887: 160; Bigot, 1892: 347; Coquillett, 1894: 92; Johnson, 1895: 325; Williston, 1901: 294; Aldrich, 1905: 240; Bezzi, 1908: 30; Kertész, 1909: 90; Verrall, 1909: 509–511; Coquillett, 1910: 559; Graenicher, 1910: 33, 35; Johnson, 1910: 747; Becker, 1913: 437; Cockerell, 1913: 231; Graenicher, 1913: 179; Johnson, 1913: 58; Cockerell, 1914: 234; White, 1916: 189, 190; Bezzi, 1924: 350; Cole et al., 1924: 184, 190; Painter, 1925: 119–127; Hallock and Parker, 1926: 6; Edwards, 1930: 164; Painter, 1930: 804; Curran, 1931: 43; Paramonov, 1931: 119; Curran, 1934: 196, 197; Brimley, 1938: 341; Painter, 1939: 43–45; Paramonov, 1939: 65; Paramonov, 1949: 631–643; D’Andretta and Carrera, 1950: 351–353, 373; D’Andretta and Carrera, 1952: 296; Gibson and Carrillo, 1959: 169; Painter and Painter, 1962: 50–53; François, 1964: 323–326; Painter and Painter, 1965: 425–426; Krombein, 1967: 5, 69, 89, 150, 202, 255, 395, 396, 408–410; Hall, 1969: 2, 3; Liepa, 1969: 8, 18; Hennig, 1973: 42; Hull, 1973: 236–240; Painter and Painter, 1974: 89–94; Hall, 1976: 3, 119–120; Merle, 1975: 9, 59, 151; Lachaise and Bowden, 1976: 331; Painter et al.,1978: 38.21; Rohlfein and Ewald, 1979: 222; Hall, 1981a: 591, 592, 597; Hall, 1981b: 161–163; Theodor, 1983: 83; Greathead, 1988: 14, 16–20; Yeates, 1994: 21, 28, 48, 64, 72–74, 82, 113, 125, 144, 148, 150, 151, 163, 183, 189, 190; Yeates and Greathead, 1997: 166, 170, 179; Evenhuis and Greathead, 1999: 208, 209; Sabrosky, 1999: 173; Coville et al., 2000: 28, 39–41, 47; Lamas and Lopes, 2004: 1–4; Hernández, 2007: 145, 156, 158; Kits et al., 2008: 2–4, 6, 27.

Diagnosis. Male dichoptic, eyes separated by distance equivalent to width of the ocellar tubercle (Fig. 1A); ocellar tubercle of the female with short and thin proclinate bristles; postpedicel with scales (Fig. 2); proboscis shorter than antenna; occiput with long and thin brown bristles, two kinds of scales, thin and spatulate; pronotum well developed; anepisternum with bristles on upper and posterior thirds; meropleurite with long hairs; scutellum with two kinds of scales, thin and spatulate; femur I with long scales on posterior surface, longitudinal row of bristles on posterodorsal surface, apical bristles on anteroventral and anterodorsal surfaces; femur III with dark scales on anterior and dorsal surfaces, apical crown of bristles; tibia I with longitudinal row of bristles on posterior surface; tibia II with dark scales on anterior surface, longitudinal row of bristles on posterior surface; abdominal segment I with long light hairs on anterior margin.

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Redescription of genus

MaleLengths. Body 10.71–21.73 mm; wing 7.86–15.14 mm.

Head. Antenna long, scales on all segments (Fig. 2); postpedicel conical, longitudinal groove on inner surface; clypeus margin with long and spatulate scales; proboscis long, but shorter than antenna (Fig. 1C); palpus formed by two palpomeres, longitudinal groove on ventral surface, short dark brown bristles; occiput with long strong bristles around the occipital foramen; lower surface of the head with long hairs (Fig. 1C).

Thorax. Pronotum arched in lateral view with spatulate scales, anterior margin with row of long and strong proclinate dark brown bristles, laterally with thin scales on all surface and row of long and strong dark brown bristles placed on medial 1/3; scutum arched with short scales, hairs, sparse bristles; posterior margin of the scutellum dark brown with medium bristles and long and strong bristles.

FIGURE 1A–D. Lepidophora culiciformis Walker, 1850: (A) Male head, dorsal view; (B) Female head, dorsal view; (C) Female head, lateral view; (D) Wing.

Wings. Variable pigmentation patterns (Fig.3); short and spatulate scales on both surfaces, costal vein with two rows of modified bristles (toothed) of variable extent; stem vein with long spatulate scales on base; basicosta with tuft of long spatulate scales (Fig. 1D); R2+3 and R4 veins recurved apically (Fig. 1D).

Legs. Tarsi with short dark brown bristles, more abundant ventrally; pulvilli light brown, 2/3 length of the claws; claws light brown, dark brown apex. Leg I. Tarsomere I with strong apical dark brown bristles on dorsal and ventral surfaces, other tarsomeres with these bristles on ventral surface. Leg III. Tarsi with strong apical dark brown bristles on ventral surface.

Abdomen. Long with short and spatulate dark brown scales; scales of different colors, kinds and length

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laterally. Male terminalia. Epandrium short, sub-rectangular, posterior margin rounded with long dark brown bristles and dark brown scales, short and spatulate on posterior ½; gonocoxite elongated, apex rounded in lateral view; hypandrium not fused to gonocoxite; basiphallus wide; epiphallus long, apex rounded in lateral view; distiphallus short and thin; gonostylus robust, apex dilated with pointed apical process.

FIGURE 2A–I. Antenna of Lepidophora Westwood species: (A) L. acroleuca Painter, 1930; (B) L. culiciformis Walker, 1850; (C) L. cuneata Painter, 1939; (D) postpedicel of L. lepidocera (Wiedemann, 1828); (E) L. lepidocera; (F) L. lutea Painter, 1962; (G) L. secutor Walker, 1857; (H) L. trypoxylona Hall, 1981; (I) L. vetusta Walker, 1857. (Bars = 1mm)

FemaleLengths. Body 9.71–9.57 mm; wing 7.0–14.00 mm.

Similar to male except: eyes separated by distance equivalent to 3.0 times width of the ocellar tubercle (Fig. 1B); frons with short and thin dark brown bristles on upper ½, some long and strong laterally, proclinate; costal vein with only short bristles. Female genitalia. Spermathecae very short, spermathecal bulbs fungiform with numerous tubules; spermathecal ducts in two sections (basal and apical), divided by sperm pump; apical

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spermathecal duct as long as sperm pump; sperm pump short, surrounded by microtubules; sclerotized collars well developed with ornamentation, delimiting the sperm pump; basal spermathecal ducts connected directly to the furca, without forming a common spermathecal duct; furca “U” shaped formed by two bars.

FIGURE 3A–H. Wings of Lepidophora Westwood species: (A) L. acroleuca Painter, 1930; (B) L. culiciformis Walker, 1850; (C) L. cuneata Painter, 1939; (D) L. lepidocera (Wiedemann, 1828); (E) L. lutea Painter, 1962; (F) L. secutor Walker, 1857; (G) L. trypoxylona Hall, 1981; (H) L. vetusta Walker, 1857.

Geographical Records (Fig. 4)

Nearctic. Canada (Ontario), United States (Alabama, South Carolina, Colorado, Florida, Georgia, Illinois, Indiana, Iowa, Kansas, Louisiana, Maine, Maryland, Michigan, Minnesota, Missouri, New Jersey, New York, Ohio, Oklahoma, Pennsylvania, Tennessee, Texas, Virginia), Mexico (Colima, Guerrero, Nayarit, Sinaloa, Sonora,

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Tamaulipas, Veracruz-Llave). Neotropical. Bolivia, Brazil (Acre, Amazonas, Bahia, Ceará, Espírito Santo, Goiás, Mato Grosso, Mato Grosso do Sul, Minas Gerais, Pará, Rondônia, Roraima, São Paulo), Colombia (Amazonas), Costa Rica (Alajuela, Guanacaste, Heredia), Guatemala (Chiquimula, Escuintla, Guatemala, Retalhuleu), Guiana, Honduras (Colon), Mexico (Chiapas, Jalisco, Morelos, Oaxaca, San Luis Potosi), Nicaragua (Esteli, Managua),Paraguay (Central), Peru (Amazonas, Loreto), Trinidad and Tobago, Venezuela (Amazonas).

FIGURE 4. Distributional pattern of Lepidophora Westwood species through the New World.

Key to species of Lepidophora Westwood

1. Wings dark brown with lighter areas on center of the cells and along the posterior margin, without hyaline areas (Figs. 3D–E); body predominantly with yellow and/or yellowish white and dark brown scales; postpedicel with sparse short dark brown scales and bristles on proximal ½ of outer surface (Figs. 2D–F); basicosta with tuft of long yellow or yellowish white scales; abdominal segment V with tuft of long and spatulate yellow, yellowish white or white scales laterally. . . . . . . . . . . . . . . . . . . 2

- Wings dark brown with hyaline or milky white areas (Figs. 3A–C and F–I); body predominantly with white and dark brown or golden and dark brown scales; postpedicel with long dark brown scales on outer surface (Figs. 2A–C and F–H); basicosta with tuft of long white or dark brown scales; abdominal segment V with only dark brown scales laterally . . . . . . . . . . . . . . . . . . . 3

2. Abdominal segment IV with short and spatulate yellow, yellowish white or white scales laterally... . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Lepidophora lutea Painter, 1962

- Abdominal segment IV with short and spatulate dark brown scales laterally ........ Lepidophora lepidocera (Wiedemann, 1828)3. Abdomen clavate or slightly clavate; abdominal segment V with dark brown scales of different kinds and lengths laterally. . . 4- Abdomen cylindrical, segments with approximately the same width; abdominal segment V with long and spatulate dark brown

scales or short and spatulate white scales laterally . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 54. Wings with hyaline areas (Fig. 3F) or milky white on distal 1/3 (Fig. 3A); abdominal segment I with long white and brown

hairs; postpedicel with long and numerous dark brown scales on proximal 1/3 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6

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- Wings with brown areas around the cells and on veins on distal 1/3 (Figs. 3C and H); abdominal segment I with long white or brown hairs; postpedicel with long and numerous dark brown scales on proximal 2/3 or on proximal ½ . . . . . . . . . . . . . . . . . 7

5. Basicosta with tuft of long brown scales; abdominal segment V with long and spatulate dark brown scales laterally; postpedi-cel with dark brown scales on all outer surface, long and numerous on proximal 2/3, whose length exceeds the length of post-pedicel (Fig. 2B); tergite VI with short dark brown covering scales . . . . . . . . . . . . . . . Lepidophora culiciformis Walker, 1850

- Basicosta with tuft of long brown and white scales; abdominal segment V with short white covering scales laterally; postpedi-cel with dark brown scales on proximal 2/3 of outer surface, long and numerous on proximal ½, whose length does not exceed the length of postpedicel (Fig. 2H); tergite VI with central tuft of short and spatulate white scales . . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Lepidophora trypoxylona Hall, 1981

6. Wings milky white and yellow veins on distal 1/3 (Fig. 3A); abdomen slightly clavate; abdominal segment V with short dark brown covering scales laterally; postpedicel with long dark brown scales that exceed its length (Fig. 2A) . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Lepidophora acroleuca Painter, 1930

- Wings totally hyaline on distal 1/3, dark veins (Fig. 3F); abdomen clavate; abdominal segment V with tuft of long and spatu-late dark brown scales on distal 1/3 laterally; postpedicel with dark brown scales that do not exceed its length . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Lepidophora secutor Walker, 1857

7. Abdominal segment I with long brown hairs; abdominal segment V with short dark brown covering scales; postpedicel with dark brown scales on proximal 2/3 of outer surface, long and numerous on proximal ½, whose length does not exceed the length of postpedicel (Fig. 2I); abdominal segment VIII with tuft of long and pedunculate dark brown scales laterally . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Lepidophora vetusta Walker, 1857

- Abdominal segment I with long white hairs; abdominal segment V with long and spatulate dark brown scales; postpedicel with dark brown scales on all outer surface, long and numerous on proximal 2/3, whose length exceeds the length of postpedicel (Fig. 2C); abdominal segment VIII with tuft of long and spatulate dark brown scales laterally . . . . . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Lepidophora cuneata Painter, 1939

Lepidophora acroleuca Painter, 1930 (Figs. 2A, 3A, 5A–B, 7)

Lepidophora acroleuca Painter, 1930: 804. Holotype male, HONDURAS, Puerto Castilla (USNM). Paramonov, 1949: 633; Hull, 1973: 238; Evenhuis and Greathead, 1999: 208.

Type material examined. Holotype: HONDURAS: Colon: Puerto Castilla [16.02, -85.97], 20.iv.26 (R.H. Painter col.), 1 male (USNM).

Other material examined. BRAZIL: Amazonas: Tefé [-3.37, -64.70], xii.24, 1 male (AMNH) [F6160–H. Bassler Collection Acc. 33591].

Diagnosis. Lepidophora acroleuca is similar to L. cuneata and L. vetusta in having a tuft composed of two colors of hairs near the posterior spiracle; slightly clavate abdomen; and abdominal segment I with long white and brown hairs. Lepidophora acroleuca differs from L. cuneata and L. vetusta in having the wing apex milky white with yellow veins (Fig. 3A); coxa II with long and strong bristles on anterodorsal and posterodorsal surfaces; femur II with short and spatulate brown and white scales on ventral surface; and abdominal segment V with short dark brown covering scales laterally.

Redescription of maleLengths. Body 15.85–17.14 mm; wing 13.14–13.35 mm.

Head. As wide as thorax; ocellar tubercle dark brown with gray pruinescence, short and thin proclinate dark brown bristles; frons polished dark brown on upper ½, gray pruinescence on lower ½, long and thin dark brown bristles and short and spatulate dark brown scales around antennal base; antenna (Fig. 2A) polished dark brown with gray pruinescence; scape 2.8 times longer than pedicel; pedicel slightly shorter than postpedicel; scape and pedicel with short and spatulate dark brown scales, long on ventral surface, medium dark brown bristles; scape with long and thin dark brown bristles on proximal 1/3; pedicel with short and strong dark brown bristles on apex and base; postpedicel with sparse short and strong dark brown bristles on base, row of long and thin dark brown scales on proximal 2/3 of outer surface, numerous and long on proximal 1/3, whose length does not exceed the length of postpedicel; face polished light brown, gray pruinescence on lower 1/3, short, medium and long spatulate dark brown scales more concentrated on clypeus margin, tuft of short and spatulate white scales along the margin of eyes at height of antennal base, long dark brown bristles bordering the eyes on upper 2/3; proboscis polished brown about ¾ length of antenna, short dark brown bristles, sparse medium bristles, except on apex of labellum,

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FIGURE 5A–F. Lepidophora Westwood species: A–B: L. acroleuca Painter, 1930, holotype male: (A) dorsal habitus; (B) lateral habitus; (C) L. culiciformis Walker, 1850, holotype female, dorsal habitus; D–E: L. cuneata Painter, 1939, lateral habitus; (D) holotype male; (E) paratype female; (F) L. lepidocera (Wiedemann, 1828), syntype female, dorsal habitus.

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FIGURE 6A–F. Lepidophora Westwood species: A–B: L. lutea Painter, 1962, holotype male; (A) dorsal habitus; (B) lateral habitus; (C) L. secutor Walker, 1857, holotype male, dorsal habitus; D–E: L. trypoxylona Hall, 1981, holotype male; (D) dorsal habitus; (E) lateral habitus; (F) L. vetusta Walker, 1857, lectotype male, dorsal habitus.

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where they are short and white, labellum 1/3 length of proboscis; palpus light brown, little more than ½ length of proboscis, palpomere I 2.3 times longer than palpomere II; occiput dark brown with gray pruinescence, short and spatulate dark brown scales on upper and lower thirds along the margin of eyes, long and thin posteriorly on lower 1/3, short and spatulate white scales on medial 1/3 along the margin of eyes, long posteriorly around the occipital foramen; lower surface of head with long brown hairs.

Thorax. Pronotum polished brown with gray pruinescence, long and thin brown bristles, anterior margin with long and thin light brown scales followed by short and spatulate dark brown scales on dorsal surface, laterally with long and thin white scales and sparse brown scales; scutum polished dark brown, margins light brown, short brown hairs, white and dark brown covering scales; postfrontal lobe with long and thin brown and white scales, long and strong dark brown bristles on anterior 1/3, medium on medial 1/3, short and thin on posterior 1/3, lower margin with long brown bristles; supra alar callus with long and thin brown and white scales, long and strong dark brown bristles, medium dark brown bristles around; postalar callus with long and strong and medium dark brown bristles, long and thin light brown and white scales, anteroinferior corner with long white and brown hairs; pleurae polished light brown with gray pruinescence; notopleuron with sparse short and spatulate white scales, long and thin white and brown scales, long brown hairs; anepisternum with sparse short and spatulate white scales, long and thin white and brown scales, long brown hairs; proepimeron with long and thin brown and white scales on the posterosuperior corner; proepisternum with sparse medium dark brown bristles, long and thin white and brown scales; katepisternum with long and thin white scales and long brown hairs on medial portion; meropleurite with long white and brown hairs on posterior 1/3; metepisternum with short white hairs; anepimeron, metepimeron, laterotergite and metanotum bare; tuft of short white and brown hairs near the posterior spiracle; halter with stem yellowish brown and short and thin white scales, knob yellowish white with sparse short and thin dark brown scales; scutellum polished dark brown, short and spatulate dark brown scales, posterior margin with short and spatulate and long and thin white scales.

Wings (Fig. 3A). Dark brown until the distal 1/3 of r1 cell, lighter areas along the posterior margin; r4, distal ½ of r2+3 and distal 1/3 of r1 milky white, R1 vein and distal 1/3 of R2+3 yellow; alula hyaline; short and spatulate dark brown scales on both surfaces of wing, concentrated on proximal ½ of r1, proximal 1/3 of r5 and r-m crossvein;

costal vein with two rows of toothed bristles that extend from distal ½ until the R4; costal vein and R1 with short dark brown bristles, short and spatulate dark brown scales, long and spatulate dark brown scales on base; stem vein with long and spatulate dark brown scales on base; basicosta with tuft of long and spatulate dark brown scales; anal cell closed, CuA2 and A1 meeting before reaching the margin of wing; calypter whitish, margin with long white and brown hairs on proximal 2/3, short brown and white scales on distal 1/3; r-m crossvein positioned on ½ of discal cell.

Legs. Polished light brown with gray pruinescence. Leg I. Coxa with long and thin white scales and long brown hairs, long and strong brown bristles on distal 1/3 of anterodorsal surface; trochanter with short and thin brown and white scales; femur with short and spatulate scales, dark brown on anterior and dorsal surfaces, white on posterior surface, sparse dark brown and white on ventral surface, long light brown hairs on posterior and ventral surfaces, longitudinal row of short and strong dark brown bristles on posterodorsal surface, strong apical dark brown bristles on anteroventral and anterodorsal surfaces; tibia with short dark brown bristles more concentrated ventrally, short and spatulate scales, dark brown on anterior, dorsal and ventral surfaces, white on posterior surface, longitudinal row of short and strong dark brown bristles on anterodorsal, posterodorsal and posterior surfaces, strong apical dark brown bristles on dorsal (3), ventral (2), posteroventral (1) and posterior (1) surfaces; tarsi with short and spatulate dark brown scales. Leg II. Coxa with long and thin white scales and long brown hairs, long and strong dark brown bristles on distal 1/3 of anterodorsal and posterodorsal surfaces; trochanter with short and thin brown and white scales; femur with short and spatulate scales, dark brown on anterior and dorsal surfaces, white on posterior surface, sparse dark brown and white on ventral surface, long light brown hairs on posterior and ventral surfaces, longitudinal row of short and strong dark brown bristles on distal ½ of anteroventral surface, strong apical dark brown bristle on posteroventral surface; tibia with short dark brown bristles, short and spatulate scales, dark brown on anterior, dorsal and ventral surfaces, white on posterior surface, longitudinal row of short and strong dark brown bristles on anterodorsal, posterodorsal, posterior, posteroventral and anteroventral surfaces, strong apical dark brown bristles on dorsal (3), ventral (2), posteroventral (1), posterior (1), anteroventral (1) and anterior (1) surfaces; tarsi with short and spatulate dark brown scales. Leg III. Coxa with long and thin white scales and long brown hairs, long and strong dark brown bristles on distal 1/3 of anterodorsal and posterodorsal surfaces;

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trochanter with short and thin brown and white scales; femur with short and spatulate dark brown scales, longitudinal row of short and strong dark brown bristles on anteroventral and ventral surfaces, apical crown of strong dark brown bristles; tibia with short dark brown bristles, short and spatulate dark brown scales, longitudinal row of short and strong dark brown bristles on anterodorsal, posteroventral, dorsal and anteroventral surfaces, strong apical dark brown bristles on ventral (3), posterior (1), posterodorsal (1), anterodorsal (1), anteroventral (1) and anterior (1) surfaces; tarsi with short and spatulate dark brown scales.

Abdomen. Polished dark brown, slightly clavate, segments II–IV narrower than others; segment I with long white hairs on anterior margin followed by long brown hairs until the anterior ½; segments II and III with sparse long brown hairs on tergites and laterally; sternites I–IV with long brown hairs, the other sternites with long dark brown hairs; segments I–IV with short and spatulate white scales laterally; segment V with short dark brown covering scales; segments VI–VIII with tuft of long and spatulate dark brown scales laterally. Male terminalia (Fig. 7). Gonocoxites 2.5 times longer than wide; gonocoxal apodeme long, apex rounded in dorsal view and dilated in lateral view; hypandrium subtriangular, anterior margin convex; basiphallus rounded in lateral view; epiphallus exceeding the apex of distiphallus in lateral view; epiphallus and distiphallus not exceeding the posterior margin of gonocoxite in lateral view; lateral aedeagal apodeme not exceeding the lateral margin of gonocoxite in dorsal view; ejaculatory apodeme long, rounded in lateral view, exceeding the anterior margin of gonocoxite in dorsal view.

FemaleUnknown.

Geographical Records (Fig. 4)

Neotropical. Honduras (Colon) and Brazil (Amazonas). Remarks. The redescription was based on only two male specimens, the holotype from Puerto Castilla

(Honduras), and a specimen from the Amazonas (Brazil), which creates a large sampling problem, and prevented the observation of possible intraspecific polymorphisms and the establishment of the geographical distribution of the species, and it does not resolve the unknown morphology of female.

Lepidophora culiciformis Walker, 1850(Figs. 1, 2B, 3B, 5C, 8)

Lepidophora culiciformis Walker, 1850: xcvii. Holotype female, BRAZIL, Pará (BMNH). Osten Sacken, 1887: 160; Painter, 1925: 119, 126; Paramonov, 1949: 631–632, 637, 643; D’Andretta and Carrera, 1950: 352–353; Hull, 1973: 238; Painter and Painter, 1974: 89–90; Painter et al.,1978: 38.21; Evenhuis and Greathead, 1999: 208; Lamas and Lopes, 2004: 1–4.

Type material examined. Holotype: BRAZIL: Pará: 1 female (BMNH) [50–2].Other material examined. BRAZIL: Pará: São João de Pirabas [-0.77, -47.17], Boa Esperança, 18–

24.v.1990 (A.L. Henriques coll.), 1 female (MPEG) [Trap Malaise]; Benevides [-1.37, -48.25], 408 km 06, 25.vi.1981 (E.L. Oliveira coll.), 1 female (MPEG); Amazonas: Manaus, vii.1935 (G.V. Vredenburg coll.), 1 female (BMNH) [B.M. 1935–615]; idem, BR 174, ZF 3, Km 23 [-2.4, -59.87], 59º52'W 2º23'S, 10.ix.1987 (M.V.B. Garcia coll.), 1 male (USNM) [Proj. DBFF. WWF Res: 3402-388-15]; idem, idem, 28.ii.1987 (M.V.B. Garcia coll.), 1 male (USNM) [Proj. DBFF. WWF Res: 1207–nº 13014]; idem, idem, 13.x.1987 (M.V.B. Garcia coll.), 1 male (USNM) [Proj. DBFF. WWF Res: 3402-390-1]; idem, 59º52'W 2º24'S, 02.xii.1987 (M.V.B. Garcia coll.), 2 females (USNM) [Conv. INPA - WWF Res: 3402-496-13 and 14]; idem, ZF 3 km, 20.ii.1987 (M.V. Garcia), 1 male (USNM) [2]; Rio Solimões [-3.25, -58.97], 3º15'S 58º58'W, 16.x.1991 (M.V.B. Garcia coll.), 4 females and 3 males (USNM) [731-2, 769-2, 734-2, 736-1, 732-1, 732-1, 732-2]; Iranduba, Rio Solimões, Ilha da Manchataria [-3.25, -60.10], 16.x.1991 (M.V. Garcia coll.), 1 female (INPA) [774-2]; Pará: Belém, Mocambo [-6.72, -69.97], 6.vii.1981 (Edimar Lima coll.), 1 male (MPEG); Primavera [-7.27, -62.33], 19.xi.1987 (P. Tadeu coll.), 1 female (MPEG); Minas Gerais: Marlieria [-19.72, -42.75], 17.viii.2003 (Y. Antonini coll.), 1 male (MZUSP) [182]; idem, 23.ix.2005, 1 male (MZUSP) [collected in the nest–emerged 26.x.2005]; idem, 9.i.2003 (Y. Antonini coll.), 1 male (MZUSP) [Lepidophora-81]; idem, 23.xii.1997 (C.D.P. Coelho coll.), 1 female (MNRJ) [MNRJ 7759]; idem,

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FIGURE 7A–C. L. acroleuca Painter, 1930, male terminalia: (A) lateral view; (B) dorsal view; (C) ventral view.

25.xii.1997 (C.D.P. Coelho coll.), 1 male (MNRJ) [MNRJ 7758]; São Paulo: São José do Rio Preto, Est. Ecol. Paulo de Faria [-20.03, -49.38], 21.xi.1998 (Garofalo, Gazola Serrano leg.), 1 female (MZUSP); idem, 14.iv.1998 (Garofalo, Gazola Serrano leg.), 1 male (MZUSP); idem, 30.ix.1998 (Garofalo, Gazola Serrano leg.), 1 male (MZUSP); idem, 14.v.1999 (Garofalo, Gazola Serrano leg.), 1 female (MZUSP); idem, 21.xi.1998 (Garofalo, Gazola Serrano leg.), 1 female (MZUSP); Matão [-21.63, 48.54], 21º37'40"S 48º32'15"W, Mata Semidecídua,

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02.xi.2007 (V.C. Silva & D.S. Amorim colls.), 1 female (MZUSP) [Biota Fapesp, Varredura]; idem, Faz. Cambuhy, 1997 (Serrano, Jesus leg.), 1 female (MZUSP).

Diagnosis. Lepidophora culiciformis is similar to L. trypoxylona in having femur I with two colors of long scales on posterior surface, long scales on ventral surface and long hairs on anterior and dorsal surfaces. Lepidophora culiciformis differs from L. trypoxylona in having scutum with short golden and dark brown scales; coxa III with long strong bristles on anterodorsal surface; tibia III with white scales on posterior surface and apical bristles on ventral, posterior, posteroventral and anteroventral surfaces; and abdominal segment I with long yellow hairs on anterior margin.

Redescription of femaleLengths. Body 9.71–19.00 mm; wing 7.00–13.57 mm.

Head (Figs. 1B and C). Wider than thorax; ocellar tubercle dark brown with gray pruinescence; frons brown with gray pruinescence, lower ½ with medium dark brown bristles, short and spatulate dark brown scales around the antennal base, slightly longer laterally; antenna (Fig. 2B) dark brown with gray pruinescence; scape 2.5 times longer than pedicel; pedicel slightly longer than postpedicel; scape and pedicel with short and spatulate dark brown scales, long on ventral surface, medium dark brown bristles more concentrated ventrally; pedicel with short and strong dark brown bristles on apex and base; postpedicel with sparse short and strong dark brown bristles on base, row of long and thin dark brown scales on all outer surface, numerous and long on proximal 2/3, whose length exceeds the length of postpedicel; face light brown with gray pruinescence, upper 1/3 with short, medium and long spatulate dark brown scales more concentrated on clypeus margin, medium white scales above the uppermost portion of clypeus margin, tuft of short and spatulate white scales along the margin of eyes at height of antennal base and long dark brown bristles bordering the eyes on upper 2/3; proboscis polished light brown, about ½ length of antenna, short and medium dark brown bristles except on apex of labellum, where they are short and white, labellum about ½ length of proboscis; palpus light brown with gray pruinescence, little more than ½ length of proboscis, palpomere I as long as palpomere II; occiput dark brown with gray pruinescence, short and spatulate dark brown scales on upper and lower thirds along the margin of eyes, long and thin posteriorly on lower 1/3, short and spatulate white scales on medial 1/3 along the margin of eyes, long posteriorly, around the occipital foramen; lower surface of head with long white hairs.

Thorax. Pronotum polished dark brown with gray pruinescence, long and thin brown bristles, anterior margin with long and thin white scales followed by short and spatulate white scales on all dorsal surface, long and thin white scales laterally; scutum dark brown with gray pruinescence, short dark brown hairs, short golden and dark brown covering scales, margins slightly lighter; postfrontal lobe with long and thin yellow scales on posterior ½, long and strong dark brown bristles on anterior 1/3, medium on medial 1/3, short and thin on posterior 1/3; supra alar callus with long and thin yellow scales on upper margin, long and strong dark brown bristles, medium around; postalar callus with long and strong and medium dark brown bristles, anterior 1/3 with long and thin yellow scales, posterior 1/3 with long and thin white scales, upper ½ with long yellow hairs; pleurae light brown with gray pruinescence; anepisternum with short and spatulate white scales, long yellow hairs, posterosuperior corner with tuft of long and thin yellow scales; proepimeron with long and thin brown and white scales on posterosuperior corner; proepisternum with sparse medium dark brown bristles, long and thin white scales; katepisternum with long and thin white and brown scales, long dark brown hairs on anterior and upper halves; meropleurite with long brown and light brown hairs; metepisternum with short white hairs; metepimeron with short white hairs on medial portion; notopleuron, anepimeron, laterotergite and metanotum bare; tuft of short white and brown hairs near the posterior spiracle; halter with stem light brown and short and thin dark brown scales, knob yellowish white with short and thin dark brown scales; scutellum polished brown with gray pruinescence, short and spatulate dark brown scales, posterior margin with short and spatulate and long and thin white scales.

Wings (Figs. 1D and 3B). Brown, hyaline areas on apex of r1 cell, distal 1/3 of r2+3, distal ½ of r5, along the posterior margin and alula; short and spatulate dark brown scales on both surfaces of wing concentrated on proximal 2/3 of r1, proximal ½ of br and upper ½ of m2; costal vein and R1 vein with short dark brown bristles, spatulate dark brown scales, long dark brown scales on base; stem vein with long and spatulate dark brown scales on base; basicosta with tuft of long and spatulate dark brown scales; anal cell open on margin of wing by distance equivalent to 1/3 length of r-m crossvein; calypter whitish, dark brown margin, small dark brown spot on proximal 1/3, margin with white and dark brown long hairs on proximal 2/3, short white and brown scales on distal 1/3; r-m crossvein positioned on ½ of discal cell; small projection toward the apex of wing on curve of R2+3.

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Legs. Polished light brown. Leg I. Coxa with long brown hairs; trochanter with short brown hairs; femur with long brown hairs, short and spatulate scales, light brown and white on posterior and anterior surfaces, dark brown on anterior and dorsal surfaces, sparse long and thin light brown and white scales on posterior and ventral surfaces, longitudinal row of short and strong dark brown bristles on posterior surface, strong apical dark brown bristles on anteroventral and anterodorsal surfaces; tibia with short dark brown bristles more concentrated ventrally, longitudinal row of short and strong dark brown bristles on posterodorsal, posterior and ventral surfaces, strong apical dark brown bristles on dorsal (3), ventral (4), posteroventral (1) and posterodorsal (1) surfaces; tarsi with short and spatulate dark brown scales, tarsomere I with longitudinal row of dark brown bristles on ventral surface. Leg II. Coxa with long white hairs, long and strong dark brown bristles on distal 1/3 of anterodorsal surface; trochanter with short brown hairs; femur with short and spatulate scales, brown on ventral surface, white on posterior surface, dark brown on anterior and dorsal surfaces, long brown hairs on ventral, anterior and posterior surfaces, longitudinal row of short and strong dark brown bristles on distal ½ of anteroventral surface, strong apical dark brown bristles on posteroventral surface; tibia with short dark brown bristles, short and spatulate scales, white on posterior surface, dark brown on anterior, dorsal and ventral surfaces, longitudinal row of short and strong dark brown bristles on anterodorsal, posterodorsal and anteroventral surfaces, strong apical dark brown bristles on dorsal (3), ventral (2), posteroventral (1), posterior (1) and anteroventral (1) surfaces. Leg III. Coxa with long white hairs, long and strong dark brown bristles on distal 1/3 of anterodorsal surface; trochanter with short brown hairs; femur with short and spatulate dark brown scales, longitudinal row of short and strong dark brown bristles on anteroventral surface and medial 1/3 of ventral surface, crown of strong apical dark brown bristles; tibia with short dark brown bristles and spatulate scales, longitudinal row of short and strong dark brown bristles on anterodorsal, posterodorsal, posterior, posteroventral and anteroventral surfaces, strong apical dark brown bristles on ventral (4), posteroventral (1), posterior (1), anteroventral (1), anterior (1), anterodorsal (1) and posterodorsal (1) surfaces.

Abdomen. Light brown, cylindrical, segments about the same width, except VI, which is somewhat broader; segment I with long brown hairs on anterior margin followed by long dark brown hairs; segments II and III with long brown hairs on sternites and laterally; segments I–IV with short and spatulate white scales laterally; segments V–VIII with tuft of long and spatulate dark brown scales laterally. Female genitalia (Figs. 8D–E). Sperm pump less than ¼ length of spermathecal duct; tubules on proximal 1/3 of spermathecal bulb; tubules on apex and until to just above the proximal ½ of apical spermathecal duct; furca with bars dorsally and laterally extended.

MaleLengths. Body 10.71–21.73 mm; wing 10.57–14.71 mm.

Similar to female except: head narrower than thorax; ocellar tubercle with long and thin proclinate dark brown bristles; frons polished dark brown on upper ½, lower ½ with long and thin dark brown bristles; clypeus only with dark brown scales; pronotum with long and thin brown scales on anterior margin, long and thin white and brown scales laterally; scutum polished brown with gray pruinescence; postfrontal lobe with long and thin dark brown scales on posterior ½; supra alar callus with long and thin dark brown scales on anterior margin; postalar callus with long and thin dark brown scales on anterior 1/3, long dark brown hairs on lower ½; pleurae polished light brown with gray pruinescence; anepisternum with long dark brown hairs, tuft of long and thin brown scales on posterosuperior corner; meropleurite with short and thin white scales on posterior 1/3; halter with stem brown and short and thin white scales, knob orange brown with short and thin white scales on base; costal vein with two rows of toothed bristles that extend from distal ½ until the R2+3 vein; anal cell closed, CuA2 and A1 are just before reaching the margin of wing; legs with gray pruinescence; coxa I with long and thin brown and white scales, long and thin dark brown bristles, long brown hairs; trochanter I with short and thin brown scales; tibia I with short and spatulate scales, white on posterior surface, dark brown on anterior, dorsal and ventral surfaces; coxa II with long and thin light brown and white scales, long dark brown and white hairs; trochanter II with short and thin brown scales. Male terminalia (Figs. 8A–C). Gonocoxite 4.3 times longer than wide; gonocoxal apodeme long, apex rounded in dorsal view; hypandrium semicircular, anterior margin slightly convex; basiphallus rounded in lateral view; epiphallus reach over the apex of distiphallus in lateral view; epiphallus and distiphallus not exceeding the posterior margin of gonocoxite in lateral view; lateral aedeagal apodeme not exceeding the lateral margin of gonocoxite in dorsal view; ejaculatory apodeme long, rounded, exceeding the anterior margin of gonocoxite in dorsal view.

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FIGURE 8A–E. L. culiciformis Walker, 1850: A–C male terminalia: (A) lateral view; (B) dorsal view; (C) ventral view; D–Efemale genitalia: (D) spermathecae; (E) furca.

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Geographical Records (Fig. 4)

Neotropical. Brazil (Amazonas, Minas Gerais, Pará, São Paulo).Remarks. The following variations were observed in the sample series (males and females): face polished

dark brown; proboscis polished dark brown; palpus dark brown; occiput with short and spatulate light brown scales on medial 1/3 along the margin of the eyes, long posteriorly around the occipital foramen; the lower region of the head with long dark brown hairs; anterior margin of the pronotum with long and thin light brown, or white and yellowish brown, or dark brown scales; dark brown scales on dorsal surface of the pronotum; lower margin of the supra alar callus with long and thin dark brown scales.

The former curator of the Natural History Museum´s Diptera collection in London, J. E. Chainey, in 1986 added a holotype label to the type of L. culiciformis examined in this study based on information affixed to the type specimen and a label handwritten by Harold Oldroyd (“though unlabeled, it is almost certainly type of L. culiciformis Wlk. 1/38. H.O.”).

Lepidophora cuneata Painter, 1939 (Figs. 2C, 3C, 5D–E, 9)

Lepidophora cuneata Painter, 1939: 43. Holotype male, PARAGUAY: Barcequillo (USNM). Paramonov, 1949: 633, 637; D’Andretta and Carrera, 1950: 353; Hull, 1973: 238; Painter and Painter, 1974: 89, 92; Merle, 1975: 151; Painter et al.,1978: 38.21; Rohlfein and Ewald, 1979: 222; Evenhuis and Greathead, 1999: 208.

Type material examined. Holotype and allotype: PARAGUAY: Central: San Lorenzo, Barcequillo [-25.33, -57.53], 2.ii.1937, 1 male and 1 female, respectively (USNM) [Type n° 101454 – n° 84].

Other material examined. BRAZIL: Ceará: Limoeiro [-7.05, -38.82], vi.1940 (Shannon and Alves colls.), 1 female (MZUSP) [20.089]; Bahia: Ibotirama, Povoado Cipó [12°08´08”S/43°19´06”W], 26–27.i.2009 (Nihei, Figueiredo, Almeida & Cezar colls.), 1 male (MZUSP); Brumado [-14.22, 41.67], 8.v.1975 (C. & P. Elias leg.), 1 male (DZUP); Mato Grosso: 1 female (ZMHB) [Mattogrosso Rohde–Lepidophora cuneata Paint. det. J.C. Hall–Zool. Mus. Berlin]; Poconé, Faz. Rio Clarinho, Trilha da Torre, Varredura [S16°36´24” W56°43´25.1”], 15.vii.2012 (Lamas, Nihei & eq. colls.), 1 male (MZUSP) [SISBIOTA CNPq/FAPESP]; Goiás: Vianópolis [-16.67, -48.58], iii.1930 (R. Spitz coll.), 1 male (MNRJ) [I - EMBRAPA - MNRJ 725]; Minas Gerais: Pirapora [-17.35, -77,93], 11–13.xi.1919 (Cornell U. Exp.), 1 female (AMNH); Goiás: Mineiros [-17.57, -52.55], xi.1994 (Paulo R. Magno coll.), 1 male (MNRJ) [MNRJ 7250]; Minas Gerais: Serra do Cipó [-19.08, -43.08], 12.ix.1998 (D. Yanega coll.), 1 female (MNRJ); Mato Grosso do Sul: Corumbá, BEP – Transição Ciliar/Paratudal, Malaise 02 [S19°34´20.09” W57°00´57.09”], 29.ix–14.x.2012 (Lamas, Nihei & eq. colls.), 1 female (MZUSP) [SISBIOTA CNPq/FAPESP]; Aquidauana, Reserva Ecológica UEMS, Vegetação aberta/Flor. Est. Decidual, Malaise 08 [S20°25´59” W55°39´20.8”], 26.x–11.xi.2011 (Lamas, Nihei & eq. colls.), 1 female (MZUSP) [SISBIOTA CNPq/FAPESP]; LOCALITY UNKNOWN: 1 male (MZUSP) [18569-62684]; Chapada?, 1 female (AMNH) [S.W. Williston Collection].

Diagnosis. Lepidophora cuneata is similar to L. vetusta in having the same color pattern of the wings in females, epiphallus exceeding the posterior margin of the gonocoxites and distiphallus reaching the posterior margin of the gonocoxites in lateral view. Lepidophora cuneata differs from L. vetusta in having pronotum brown, margins light brown; coxa III with long strong bristles on dorsal surface; femur II with white scales on dorsal surface; tibia III with apical bristles on ventral, posteroventral, posterior, anteroventral, anterodorsal, posterodorsal and dorsal surfaces; abdominal segment I with long white hairs; and tergites V and VII with short white and dark brown covering scales.

Redescription of maleLengths. Body 15.92–18.14 mm; wing 11.64–12.86 mm.

Head. Wider than thorax; ocellar tubercle brown with gray pruinescence, medium and thin proclinate dark brown bristles; frons polished dark brown on upper ½, light brown with gray pruinescence on lower ½, short, medium and long thin dark brown bristles and short and spatulate dark brown scales around the antennal base; antenna (Fig. 2C) brown with gray pruinescence, base of scape orange; scape 2.4 times longer than pedicel; pedicel

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slightly longer than postpedicel; scape and pedicel with short and spatulate dark brown scales, long on ventral surface, medium dark brown bristles; scape with long and thin dark brown bristles on proximal 1/3; pedicel with short and strong dark brown bristles on base and apex, short and spatulate white scales, sparse light brown scales on dorsal surface; postpedicel dark brown with gray pruinescence, sparse short and strong dark brown bristles on base, row of long and thin dark brown scales on all outer surface, long and numerous on proximal 2/3, whose length exceeds the length of postpedicel; face polished light brown, gray pruinescence on lower 1/3, upper 1/3 with short, medium and long dark brown scales more concentrated on clypeus margin, medium white and light brown scales above the uppermost portion of clypeus margin, long dark brown bristles bordering the eyes on upper 2/3; proboscis polished brown, 3/5 length of antenna, short dark brown bristles except on apex of labellum, where they are light brown, labellum about ¼ length of proboscis; palpus polished brown with gray pruinescence, 2/5 length of proboscis, palpomere I little longer than palpomere II; occiput dark brown with gray pruinescence, short and spatulate dark brown scales on upper and lower thirds along the margin of eyes, long and thin posteriorly on lower 1/3, short and spatulate white and light brown scales on medial 1/3 along the margin of eyes, long and spatulate and long and thin posteriorly around the occipital foramen; lower surface of head with long brown hairs.

Thorax. Pronotum polished dark brown with gray pruinescence, long and thin brown bristles, posterior margin and lateral light brown, anterior margin with long and thin light brown and white scales followed by short and spatulate dark brown scales on all dorsal surface, laterally with white scales and sparse long and thin dark brown scales; scutum dark brown with gray pruinescence, margins orange brown, short brown hairs, short white covering scales; postfrontal lobe with sparse long and thin white scales on lower 1/3, short and spatulate dark brown scales on upper and posterior thirds, long and strong dark brown bristles on anterior 1/3, medium on medial 1/3, short and thin on posterior 1/3; supra alar callus with sparse long and thin white and brown scales, long and strong dark brown bristles, sparse medium dark brown bristles around; postalar callus with long and strong and medium dark brown bristles, long and thin white scales, long light brown and white hairs; pleurae polished light brown with gray pruinescence; notopleuron with short and spatulate and long and thin white scales; anepisternum orange on upper and posterior thirds, central dark brown spot, greater on anterior ½, short and spatulate white scales, posterior margin with tuft of long and thin brown and white scales, anteroinferior corner with long light brown hairs; proepimeron with long and thin white scales on posterosuperior corner; proepisternum with sparse medium light brown bristles, long and thin white scales; katepisternum with sparse long and thin white and brown scales, long light brown hairs more concentrated on lower 1/3; meropleurite with long light brown hairs on posterior 1/3; anepimeron, metepisternum, metepimeron, laterotergite and metanotum bare; tuft of short white and light brown hairs near the posterior spiracle; halter with yellowish brown stem and short and thin white scales, knob yellowish white with sparse short and thin dark brown scales; scutellum polished dark brown, margins orange, short and spatulate dark brown scales, posterior margin with long and spatulate and long and thin white scales.

Wings (Fig. 3C). Brown until the distal 1/3 of R1 vein, height of discal cell and surrounding the R2+3; r4, m1 and m2 cells, center of distal ½ of r1 and center of discal cell lighter, other areas hyaline; short and spatulate brown scales on both surfaces of wing concentrated on apex of r1 and until the height of lower 1/3 of wing; costal vein with two

rows of toothed bristles that extend of distal ½ of wing until the R2+3; costal vein and R1 with short dark brown bristles and short and spatulate dark brown scales, long dark brown scales on base; stem vein with long and spatulate dark brown scales on base; basicosta with tuft of long and spatulate dark brown scales; anal cell closed, CuA2 and A1 closed on margin of wing; calypter whitish, small dark brown spot on proximal 1/3, margin with long white and dark brown hairs on proximal 2/3, short white scales on distal 1/3; r-m crossvein positioned on ½ of discal cell; small projections directed to the apex of wing on curve of R2+3 and R4 shortly after the bifurcation with R5.

Legs. Polished orange brown with gray pruinescence. Leg I. Coxa with sparse long and thin brown and white scales and brown and dark brown hairs, long and strong apical dark brown bristles on anterodorsal surface; trochanter with short and thin dark brown scales, sparse short brown hairs; femur with short and spatulate scales, white and light brown on posterior surface, dark brown on anterior, dorsal and ventral surfaces, longitudinal row of short and strong dark brown bristles on posterodorsal surface, strong apical dark brown bristles on anteroventral surface; tibia with short dark brown bristles more concentrated on anterior and ventral surfaces, short and spatulate scales, white on posterior and ventral surfaces, dark brown on anterior and dorsal surfaces, longitudinal row of short and strong dark brown bristles on posterodorsal and posterior surfaces, strong apical dark brown bristles on dorsal (3), ventral (2), posteroventral (1) and posterior (1) surfaces; tarsi with sparse short and spatulate dark brown

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scales. Leg II. Coxa with long and thin light brown and white scales and long dark brown and light brown hairs, long and strong dark brown bristles on distal 1/3 of anteroventral and anterodorsal surfaces; trochanter with short and thin dark brown scales, sparse short brown hairs; femur with short and spatulate scales, brown on ventral and anterior surfaces, white on posterior and dorsal surfaces, long brown hairs on ventral and posterior surfaces, longitudinal row of short and strong dark brown bristles on distal ½ of anteroventral surface, strong apical dark brown bristle on posteroventral surface; tibia with short dark brown bristles, short and spatulate scales, white on dorsal and posterior surfaces, dark brown on anterior and ventral surfaces, longitudinal row of short and strong dark brown bristles on anterodorsal, posterodorsal, posterior, posteroventral and ventral surfaces, strong apical dark brown bristles on dorsal (3), ventral (2), posterior (1), posteroventral (1), anteroventral (1) and anterior (1) surfaces; tarsi with sparse short and spatulate dark brown scales. Leg III. Coxa with long and thin light brown scales and long brown and dark brown hairs, long and strong dark brown bristles on distal 1/3 of dorsal surface; trochanter with short and thin dark brown scales, sparse short brown hairs; femur with short and spatulate scales, brown on ventral and posterior surfaces, dark brown on dorsal and anterior surfaces, longitudinal row of short and strong dark brown bristles on distal ½ of anteroventral and ventral surfaces, crown of strong apical dark brown bristles; tibia with short dark brown bristles and spatulate scales, longitudinal row of short and strong dark brown bristles on anterodorsal, posterodorsal, posterior, posteroventral and anteroventral surfaces, strong apical dark brown bristles on ventral (2), posteroventral (1), posterior (1), anteroventral (2), anterodorsal (1), posterodorsal (1) and dorsal (1) surfaces; tarsi with short and spatulate dark brown scales.

Abdomen. Polished brown, lateral and margins of segment IV light orange brown, slightly clavate, segments III and IV narrower than others; segment I with long white hairs on anterior margin; sternites I–IV with long white hairs, other sternites with long dark brown hairs; segments I–IV with short and spatulate white scales laterally; tergites V and VII with short and spatulate white scales; segments V–VIII with tuft of long and spatulate dark brown scales laterally. Male terminalia (Figs. 9A–C). Gonocoxite 2.2 times longer than wide; gonocoxal apodeme long, apex rounded in dorsal view and dilated in lateral view; hypandrium semicircular, borders slightly rounded; basiphallus rounded with rounded medial projection in lateral view; distiphallus not exceeding the height of epiphallus and reaching the posterior margin of gonocoxite in lateral view; epiphallus dilated dorsally, apex surpassing the posterior margin of gonocoxite in lateral view; lateral aedeagal apodeme not exceeding the lateral margin of gonocoxite in dorsal view; ejaculatory apodeme long, rounded in lateral view, exceeding the anterior margin of gonocoxite in dorsal view.

FemaleLengths. Body 12.28–18.28 mm; wing 8.71–11.86 mm.

Similar to male, except: frons with medium dark brown bristles on lower ½; pedicel about the same length of postpedicel; scape without long and thin dark brown bristles on proximal 1/3; pedicel without white scales on dorsal surface; postfrontal lobe without dark brown scales on upper and lower thirds; scutellum polished orange brown; wings dark brown, center of distal ½ de R1 and center of discal cell lighter; costal vein with short and spatulate white scales on base; basicosta with tuft of long and spatulate white, light brown and dark brown scales; calypter whitish light brown; femur I with short and spatulate light brown and white scales on ventral surface, long and spatulate light brown scales on dorsal surface; tarsomere I without row of dark brown bristles on ventral surface; coxa II with long dark brown, light brown and white hairs; femur II with short and spatulate scales, dark brown on anterior and dorsal surfaces, light brown and white on posterior and ventral surfaces; coxa III with long and thin light brown and white scales and long brown, dark brown and white hairs. Female genitalia (Figs. 9D–E). Sperm pump just under 1/5 length of spermathecal duct; tubules on proximal ½ of spermathecal bulb; tubules on apex and proximal ½ of apical spermathecal duct; furca with bars dilated anteriorly, posteriorly and laterally.

Geographical Records (Fig. 4)

Neotropical. Bolivia, Paraguay (Central), Trinidad and Tobago, Brazil (Amazonas, Bahia, Ceará, Goiás, Mato Grosso, Mato Grosso do Sul, Minas Gerais).

Remarks. The following variations were observed in the sample series (males and females): antenna dark brown, base of scape lighter; clypeus only with dark brown scales.

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FIGURE 9A–E. L. cuneata Painter, 1939: A–C male terminalia: (A) lateral view; (B) dorsal view; (C) ventral view; D–Efemale genitalia: (D) spermathecae; (E) furca.

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Lepidophora lepidocera (Wiedemann, 1828) (Figs. 2D–E, 3D, 5F, 10)

Toxophora lepidocera Wiedemann, 1828: 360. Syntype female, LOCALITY UNKNOWN (SMFD). Westwood, 1835: 447; Osten Sacken, 1887: 160; Johnson, 1895: 321; Painter, 1925: 119–122; Curran, 1931: 43; Paramonov, 1949: 631–634, 636–639; Painter and Painter, 1962: 50–51; Painter and Painter, 1965: 426; Krombein, 1967: 5, 69, 150, 202, 395, 408; Hull, 1973: 16, 238; Merle, 1975: 59, 151; Painter et al.,1978: 38.21; Hall, 1981: 161; Theodor, 1983: 83; Yeates and Greathead, 1997: 166, 170, 179; Evenhuis and Greathead, 1999: 208; Lamas and Lopes, 2004: 1.

Ploas aegeriiformis Gray in Griffith and Pidgeon, 1832: pl. 128, Fig. 6. Holotype male, LOCALITY UNKNOWN (UMO). Macquart, 1840: 119; Macquart, 1846: 350; Osten Sacken, 1858: 44; Osten Sacken, 1887: 160; Aldrich, 1905: 240; Coquillett, 1910: 599; Graenicher, 1910: 33, 35; Johnson, 1910: 747; Johnson, 1913: 58; Cole et al., 1924: 190; Curran, 1931: 43; Paramonov, 1949: 631, 639; Painter and Painter, 1962: 50. Correct original spelling determined by First Reviser in Sabrosky (1999: 173).

Toxophora appendiculata Macquart, 1846: 246(118); Osten Sacken, 1887: 160; Aldrich, 1905: 240; Painter, 1925: 119–122; Paramonov, 1949: 633, 634, 636; Painter and Painter, 1962: 51; Krombein, 1967: 255; Hull, 1973: 16, 238.

Ploas aegeriformis, incorrect original spelling of aegeriiformis (Gray in Griffith and Pidgeon, 1832: 792).Ploas oegeriformis, incorrect original spelling of aegeriiformis (Gray in Griffith and Pidgeon, 1832: 779).

Type material examined. Holotype: LOCALITY UNKNOWN: 1 male (UMO) [Ploas aegeriiformis]; Syntype: LOCALITY UNKNOWN: [Brazilia Freireiss?], 1 female (SMFD) [Toxophora lepidocera].

Other material examined. UNITED STATES: Minnesota: Itasca State Park [47.20, -95.20], 24.vii.1914, 1 male (USNM); New York: Texas Hollow, Schuyler County, Bennettsburg [42.42, -76.81], 26.vii.1983, 1 male (AMNH); Illinois: Palos Park [41.67, -87.83], 22.vii.1923 (Owen Bryant coll.), 1 male (CAS); Ohio: Akron [41.08, -81.52], 1949 (W.D. Stockton coll.), 1 male (CAS) [California State Univ., Long Beach 1996 donation to Calif. Acad. Sci.]; New Jersey: Lakehurst [40.01, -74.31], 26.viii.1934, 1 male (USNM) [A.L Melander Collection 1961]; Colorado: Lafayette [40.00, -105.09], 15.i.1960 (F.M. Hull coll.), 1 female (BMNH) [J. Bowden collection - BMNH(E) 2003–159]; New Jersey: Burlington County, Oswego Lake [39.72, -74.49], 30.viii.1974 (A.S. Menke coll.), 1 female (USNM); Missouri: Higginsville [36.07, -93.72], 19.vii.2003 (E.A. Dodge coll.), 1 female (CAS); Maryland: Prince George County, Patuxent Refuge [39.10, -77.30], 19.vii.1965 (P.H. Thompson coll.), 1 male (USNM); Beltsville [39.03, -76.91], 31.vii.1966 (D.R. Smith coll.), 1 male (USNM); Kansas: Douglas County, Natural History Reserve [38.97, -95.23], 5 mi. Lawrence, 28.vii.1979 (N.E. Woodley coll.), 1 male (USNM); Virginia: near Vienna [38.90, -77,26], 12.ix.1937 (ABGurney coll.), 1 female (USNM); Nelson County [38.32, -79.47],30.vii.1928 (W. Robinson coll.), 1 male (USNM); Princess Anne County, 1.3 km W Pungo [36.72, -76.02], 4.viii.1984 (D.S. Bogar & W.E. Steiner colls.), 1 male (USNM); Oklahoma: Quinton [35.12, -95.37], 10.vi.1934 (A.E. Pritchard coll.), 1 male (USNM) [R.H. Painter Collection 1975]; idem (J. Stankavich coll.), 1 male (CAS); Alabama: Sheffield [34.76, -87.70], 4.viii.1944 (G.E. Bohart coll.), 1 male (CAS); Texas: 1 female (ZMHB) [Texas Boll. – 7375 – Zool. Mus. Berlin]; Dallas County, Irving [32.81, -96.95], 21.ix.1982 (Russell A. Rahn leg.), 1 female (USNM); Georgia: 1 female (BMNH) [107]; 1 female (ZMHB) [Esclier - 1314 - Zool. Mus. Berlin]; Tifton [31.45, -85.51], 16.x.1996, 1 female (USNM) [ALMelander collection 1961]; Texas: Jasper County [30.92, -94.00], 20.x.1965 (L.D. McWilliams III leg.), 1 male (USNM) [on flowers]; College Station [30.63, -69.33], 11.xi.1919 (H.J. Reinhard coll.), 1 female (USNM); idem, 11.viii.1919 (idem), 1 male (USNM); idem, 11.vi.19 (H.J. Reinhard Collector), 1 female (ZMHB) [Lepidophora appendiculata Mqt. det. H.J. Reinhard – Lepidophora/L. appendiculata Bgt. – Zool. Mus. Berlin]; Brownwood [30.11, -93.73], 14.viii.1921 (R.H. Painter coll.), 1 female (USNM); Florida: Lacoochee [28.46, -82.17], 9.viii.1939 (E.G. Wegenek coll.), 1 female (USNM) [Lepidophora lepidocera Wied Painter 1960 Homotype]; 1 male (USNM) [Collection C.V. Riley]; Highlands County, Lake Placid [27.29, -81.36], 13.iii.1958 (K.V. Krombein coll.), 1 male (USNM) [Frank M. Hull Collection C.N.C. 1981]; idem, 14.ix.1960 (K.V. Krombein coll.), 1 female (USNM) [Reared Podium carolina nest (Roflower) - Sphecidae- Frank M. Hull Collection C.N.C. 1981]; idem, 13.vii.1948 (E.L. Todd coll.), 1 male and 1 female (USNM) [R.H. Painter Collection 1975]; 10–40 miles of Everglades City [25.86, -81.38], 30.viii.1931 (Bradley & Knorr coll.), 1 female (USNM) [R.H. Painter Collection 1975]; Highlands County, Lake Placid, Archbold Biological Station [27.18, -81.35], 13.v.1961 (H.E. & M.A. Evans colls.), 1 male (USNM) [R.H. Painter Collection 1975]; idem, 11.v.1961 (H.E. & M.A. Evans colls.), 1 male (USNM) [H.E. & M.A. Evans–Biol. Note nº 1696 H.E. Evans–R.H. Painter Collection 1975]; idem, 2.viii.1985 (M. Deyrup coll.), 1 female (USNM) [Malaise Trap Trail 2 SS0]; LOCALITY UNKNOWN: “America Boreal”, 1 male (BMNH) [Bigot Collection–B.M. 1960–539].

Diagnosis. Lepidophora lepidocera is similar to L. lutea in having postpedicel with sparse short and thin scales

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on proximal ½ of outer surface; body usually covered with yellow, or yellowish white and dark brown scales; scutellum with two color of short scales; dark brown wings with lighter areas on center of the cells and along the posterior margin, without hyaline areas; basicosta with tuft of long yellow or yellowish white scales; coxa I with light hairs; coxa III with light bristles; abdominal segment V with long light scales forming central tuft laterally; and short gonocoxal apodeme. Lepidophora lepidocera differs from L. lutea in having pronotum with short and spatulate dark brown and yellow scales; tibia I with longitudinal row of bristles on dorsal and posterodorsal surfaces; tibia III with longitudinal row of bristles on ventral surface; and abdominal segment IV with short dark brown scales laterally.

Redescription of maleLengths. Body 11.57–21.43 mm; wing 7.86–14.28 mm.

Head. Slightly narrower than thorax; ocellar tubercle dark brown with gray pruinescence, long and thin proclinate dark brown bristles; frons dark brown, gray pruinescence on lower ½, long and thin dark brown bristles and short and spatulate dark brown scales around the antennal base; antenna (Fig. 2E) dark brown with gray pruinescence; scape 2.0 times longer than pedicel; pedicel slightly longer than postpedicel; scape with short and spatulate dark brown scales, long on ventral surface, medium dark brown bristles; pedicel with short and thin dark brown scales, long on ventral surface, medium dark brown bristles, short and strong dark brown bristles on apex; postpedicel (Fig. 2D) with sparse dark brown bristles, short and strong on base, outer surface with medium bristles on proximal ½, sparse short and thin dark brown scales on proximal 1/3; face polished dark brown, gray pruinescence on lower and upper thirds, short and spatulate dark brown scales, medium and long on upper 1/3, more concentrated on clypeus margin, tuft of short and spatulate white scales along the margin of eyes at height of antennal base, long dark brown bristles bordering the eyes on upper 2/3; proboscis polished dark brown, about ¾ length of antenna and short dark brown bristles, labellum ½ length of proboscis; palpus polished dark brown, a little less than ½ length of proboscis, palpomere I as long as palpomere II; occiput dark brown with gray pruinescence, short and spatulate dark brown scales on upper and lower thirds along of the margin of eyes, long and thin posteriorly on lower 1/3, short and spatulate white scales on medial 1/3 along of the margin of eyes, long and spatulate posteriorly around the occipital foramen; lower surface of head with long white hairs.

Thorax. Pronotum polished dark brown with gray pruinescence, long and thin dark brown bristles, sparse long and thin white scales, anterior margin with long and thin white scales followed by dark brown scales, long and thin white scales laterally; scutum polished dark brown, margins slightly lighter, sparse short brown hairs and white covering scales, dark brown covering scales on posterior ½, short and long yellowish white scales on posterior 1/3; postfrontal lobe with long and thin white scales and sparse long dark brown bristles, long and strong dark brown bristles on anterior 1/3, medium on medial 1/3, short and thin on posterior 1/3; supra alar callus with long and thin white scales, long and strong dark brown bristles, medium dark brown bristles around; postalar callus with long and strong and medium dark brown bristles, lower ½ with long and thin white scales and hairs; pleurae polished light brown with gray pruinescence; anepisternum grayish brown on upper 1/3, short and spatulate white scales, long brown hairs, margins with long and thin white scales, posterosuperior corner with tuft of long and thin white scales and hairs; proepimeron with long and thin white scales on posterosuperior corner; proepisternum with sparse medium dark brown bristles, long and thin white scales; katepisternum grayish brown on lower ½, long and thin white scales, upper and lower margins with long white hairs; anepimeron with sparse short white hairs on lower 1/3; meropleurite with long white hairs on posterior 1/3; metepisternum with short white hairs on lower 1/3, sparse on medial portion; metepimeron with sparse short white hairs on lower 1/3; notopleuron, laterotergite and metanotum bare; tuft of short white hairs near the posterior spiracle; halter with stem yellowish brown and short and thin white scales, knob yellowish white with short and thin dark brown scales; scutellum polished dark brown, short and spatulate dark brown and yellow scales, posterior margin with long and spatulate and long and thin yellow scales.

Wings (Fig. 3D). Brown, lighter areas on center of discal cell, center of ½ of r1 and r2+3 and along the posterior margin; alula hyaline; short and spatulate dark brown scales on both surfaces of wing concentrated until the distal 1/3 of r1 and r2+3 and even at height of proximal ½ of discal cell and upper margin of anal cell; costal vein with two rows of toothed bristles that extend from proximal 1/3, at height of beginning of discal cell, until the apex of R2+3

vein; costal vein and R1 with short dark brown bristles, small tuft of short and spatulate white and dark brown scales on base; costal vein and stem vein with long and spatulate white and dark brown scales on base; basicosta with tuft

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of long and spatulate yellowish white scales; anal cell open on wing margin by distance equivalent to 1/3 length of r-m crossvein; calypter whitish, dark brown spot on proximal 1/3, margin with long white hairs on proximal 2/3, short white scales on distal 1/3; r-m crossvein positioned just beyond the ½ of discal cell; small projections on base of R4 forming a “V”.

Legs. Polished brown. Leg I. Coxa with gray pruinescence, long and thin white scales and long and short white hairs, long and strong yellow bristles on distal 1/3 of anterodorsal surface; trochanter with short and thin white scales; femur with short and spatulate scales, dark brown on anterior and dorsal surfaces, white on ventral and posterior surfaces, long and thin white scales on posterior surface, longitudinal row of short and strong dark brown bristles on posterodorsal surface, strong apical dark brown bristles on anteroventral and anterodorsal surfaces; tibia with short dark brown bristles more concentrated on anterior and ventral surfaces, short and spatulate scales, dark brown on anterior, dorsal and ventral surfaces, white on posterior surface, longitudinal row of short and strong dark brown bristles on posterodorsal, posteroventral and dorsal surfaces, strong apical dark brown bristles on dorsal (2), posterior (1), anteroventral (1), ventral (1) and posteroventral (1) surfaces; tarsi with short and spatulate dark brown scales. Leg II. Coxa with gray pruinescence, long and thin white scales and long and short white hairs, long and strong dark brown bristles on distal 1/3 of anteroventral surface; trochanter with short and thin white scales, short white hairs; femur with short and spatulate scales, dark brown on anterior and dorsal surfaces, white on posterior and ventral surfaces, long white and light brown hairs on posterior and ventral surfaces, longitudinal row of short and strong dark brown bristles on anteroventral surface, strong dark brown bristle on distal 1/3 of posteroventral surface; tibia with short dark brown bristles, short and spatulate scales, dark brown on anterior, ventral and dorsal surfaces, white on posterior surface, longitudinal row of short and strong dark brown bristles on anterodorsal, posterodorsal, posteroventral, posterior and anteroventral surfaces, strong apical dark brown bristles on dorsal (3), posterior (2), ventral (1), posteroventral (1) and anterior (2) surfaces; tarsi with short and spatulate dark brown scales. Leg III. Coxa with gray pruinescence, long and thin white scales and long and short white hairs, long and strong yellow bristles on distal 1/3 of posterodorsal surface; trochanter with short and thin white scales, short white hairs; femur with short and spatulate scales, dark brown on anterior and dorsal surfaces, white on posterior and ventral surfaces, longitudinal row of short and strong dark brown bristles on anteroventral and ventral surfaces, apical crown of strong dark brown bristles; tibia with short dark brown bristles, short and spatulate dark brown scales, longitudinal row of short and strong dark brown bristles on anteroventral, anterodorsal, posteroventral, posterodorsal, ventral and posterior surfaces, strong apical dark brown bristles on posterior (1), posteroventral (1), ventral (3), anterior (1), dorsal (2) and anteroventral (1) surfaces; tarsi with short and spatulate dark brown scales.

Abdomen. Polished brown with gray pruinescence, cylindrical, segments about the same width; segment I with long white hairs on anterior margin followed by long brown and dark brown hairs; tergites II and III with long white and light brown hairs; sternites I and II with long white hairs, other sternites with long dark brown hairs; segments I–III and V with short and spatulate white scales, short dark brown covering scales and long white and brown hairs laterally; segment IV with dark brown covering scales laterally; segment V with tuft of long and spatulate yellow scales laterally; segments VI–VIII with tuft of long and spatulate dark brown scales laterally. Male terminalia (Figs. 10A–C). Gonocoxites 2.3 times longer than wide; gonocoxal apodeme short, apex pointed in dorsal view, enlarged in lateral view; hypandrium semicircular, borders slightly rounded, anterior margin slightly concave; basiphallus with slightly straight margin and rounded borders in lateral view; distiphallus extends to just before the apex of epiphallus in lateral view; epiphallus and distiphallus not exceeding the posterior margin of gonocoxite in lateral view; lateral aedeagal apodeme not exceeding the lateral margin of gonocoxite in dorsal view; ejaculatory apodeme long spatulate, exceeding the anterior margin of gonocoxite in dorsal view.

FemaleLengths. Body 13.71–18.71 mm; wing 9.71–12.57 mm.

Similar to male, except: ocellar tubercle light brown; frons with medium dark brown bristles on lower ½; base of scape lighter; postpedicel with short and strong dark brown bristles on proximal 2/3 of outer surface; labellum 1/5 length of proboscis; ventral surface of wing with short and spatulate dark brown and white scales; costal vein and stem vein with only short and spatulate dark brown scales on base; femur I with short and spatulate white scales on dorsal surface; coxa II with long and strong yellow bristles on distal 1/3 of anterodorsal surface; coxa III

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FIGURE 10A–E. L. lepidocera (Wiedemann, 1828): A–C male terminalia: (A) lateral view; (B) dorsal view; (C) ventral view; D–E female genitalia: (D) spermathecae; (E) furca.

Zootaxa 3682 (1) © 2013 Magnolia Press · 25REVISION OF LEPIDOPHORA WESTWOOD

with long and strong yellow bristles on distal 1/3 of anterodosal surface; abdominal segments I–III with short and spatulate yellowish white scales laterally. Female genitalia (Figs. 10D–E). Sperm pump less than 1/5 length of spermathecal duct; tubules on proximal ½ of spermathecal bulb; tubules on apical spermathecal duct, except on a small area just above the proximal ½; furca with dilated bars dorsally and laterally, slightly dilated posteriorly.

Geographical Records (Fig. 4)

Nearctic. United States (Alabama, South Carolina, Colorado, Florida, Georgia, Illinois, Iowa, Kansas, Louisiana, Maryland, Minnesota, Missouri, New York, New Jersey, Ohio, Oklahoma, Texas, Virginia).

Remarks. The following variations were observed in the sample series (males and females): scales covering the body usually yellow; occiput with short and spatulate white and yellowish white scales on medial 1/3 along the margin of eyes, long and spatulate yellow scales posteriorly around of the occipital foramen; anterior margin of the pronotum with long and thin yellowish white or yellow, or dark brown scales; dorsal surface of pronotum with long and thin yellowish white scales, laterally with long and thin yellow scales; yellow scales on surface of wing.

Lepidophora lutea Painter, 1962 (Figs. 2F, 3E, 6A–B,11)

Lepidophora lutea Painter, 1962: 51. Holotype male, UNITED STATES: New Jersey: Whiting (USNM). Painter and Painter, 1965: 426; Hull, 1973: 238; Hall, 1981a: 592, 597; Yeates, 1994: 21, 72, 125; Evenhuis and Greathead, 1999: 208; Kits et al., 2008: 2–4, 6, 27.

Type material examined. Holotype: UNITED STATES: New Jersey: Whiting [39.95, -74.38], 15.viii.20/6?, 1 male (USNM); paratypes: UNITED STATES: Maine: Rangeley [44.97, -70.64], 9.viii.1936 (C.F. dos Passos coll.), 1 male (AMNH) [Lepidophora lutea R.H. Painter Paratype 1961]; New York: Tuxedo, Sta. Study Insects [41.22, -74.19], 29.vii.1928, 1 male (CAS); idem, 28.viii.1928 (C.H. Curran coll.), 1 male (USNM) [R.H. Painter Collection 1975 - Lepidophora lutea R.H. Painter Paratype 1961]; idem, 25.vii.1928 (C.H. Curran coll.), 1 female (USNM) [Lepidophora lutea R.H. Painter Paratype 1961]; idem, 18.viii.1928 (C.H. Curran coll.), 1 male (USNM) [R.H. Painter Collection 1975 - Lepidophora lutea R.H. Painter Paratype 1961]; idem, 30.vii.1928, 1 female (AMNH) [Lepidophora lutea R.H. Painter Paratype 1961]; New Jersey: "Greenwood" Lake [41.18, -74.33], (A. Nicolay coll.), 1 female (CAS); Pennsylvania: Monroe County, Delaware Water Gap [40,97, -75,13], 1 female (USNM) [R.H. Painter Collection 1975–Lepidophora lutea R.H. Painter Paratype 1961–Collection of Mrs. A.T. Slosson Ac. 26226].

Other material examined. UNITED STATES: Ohio: Holmes County, Richland [39.19, -82.60], 15.vii.1934 (C.F. Walker & E.S. Thomas colls.), 1 female (USNM) [R.H. Painter Collection 1975]. 1 male (USNM) [Insect Book Pl. 18 fig.21 Tex.?]; CANADA: Ontario: 15 mi. SE Kenora [49.77, -94.47], 7.viii.1979 (J.R. Powers. coll.), 1 male (CAS); idem, 4.viii.1979 (J.R. Powers. coll.), 2 males (CAS); UNITED STATES: Minnesota: Eagles Nest [47.84, -92.10], 31.vii.1959 (W.V. Balduf coll.), 1 male (USNM); idem, 5.viii.1957 (W.V. Balduf coll.), 1 female (USNM) [Fly in flower honey lop]; Michigan: Grand Rapids [46.76, -89.27], 16.ix.1946, 1 male (USNM); idem, 11.ix.1946, 1 male (USNM); Presque Isle County, Ocqueoc Lake [45.47, -84.11], 18–23.vii.1987 (R.W. & E.R. Hodges colls.), 1 male and 1 female (USNM); New York: Monroe County, Penfield, Ellison Park [43.15, -77.52], 31.vii.1981 (F.G. Howarth coll.), 1 female (USNM) [Bishop Museum]; Illinois: Palos Park [41.67, -87.83], 22.vii.1923 (Owen Bryant coll.), 4 males (2–USNM and 2–CAS); Pennsylvania: Lake Winola [41.51, -75.85], 25.viii.1916 (E.G. Anderson coll.), 1 male (USNM) [on Solidago]; New York: Orange County, Cornwall on Hudson, Black Rock Forest [41.41, -74.02], UTM 18-4585527N 582809E, 26.vii.2003 (V. Giles & J.G. Rozen colls.), 1 male (AMNH); Tuxedo, Sta. Study Insects [41.22, -74.19], 29.vii.1928, 1 female (USNM) [S.W. Bromley Collection 1955]; Pennsylvania: State College [40.79, -77.86], 8.iv.1912, 1 female (USNM); Lehigh Gap [40.78, -75.61], 20.vii.2002, 1 male (USNM); Indiana: Lafayette [40.42, -86.87], 10.vii.1914 (J.J. Davis coll.), 1 male (USNM) [Collection J.M. Aldrich]; Pennsylvania: Huntingdon County, Corn Propsts Mills [40.41, -77.98], 26.ix.1973 (D. Dee Wilder coll.), 1 male (CAS) [D. Dee Wilder Collection]; 5 mi. NW Davidsburg [39.98, -76.89], 3.ix.1962 (Paul J. Spangler coll.), 1 female (USNM); Ohio: Muskineum County, Norwich [39.98, -81.79],

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1.ix.1971 (A.S. Menke coll.), 1 male (USNM); Hocking County [39.83, -82.80], 28.viii.1943, 1 male (USNM) [O. SW. Bromley Collection 1955]; Pennsylvania: Blue Ridge Summit [39.72, -77.47], 8.viii.1920 (J.M. Aldrich coll.), 1 female (USNM); Maryland: Morgan County, 7.2 mi W. Berkeley Springs [39.36, -77.05], 17.vii.1981 (G.F. & J.F. Hevel colls.), 1 male and 1 female (USNM); Bowie [39.01, -76.78], 1–30.vi.1945 (DD Expt.?), 1 female (USNM); Virginia: Nelson County [38.32, -79.47], 30.vii.1928 (W. Robinson coll.), 1 male (USNM); Buckeye [37.69, -77.32], 31.vii.1939, 2 females (USNM); Wingina [37.64, -78.72], 14.viii.1928 (W. Robinson coll.), 1 female (USNM); Speedwell [36,82, -81,18], 11.viii.1938, 1 female (USNM); Lee County, Cumberland Mountains [36.75, -83.20], (H.G. Hubard coll.), 2 females (USNM); Sulphur Springs [36.72, -83.08], 17.vii.1911 (W. Robinson coll.), 1 female (USNM); Tennessee: Miller Johnson City [36.31, -82.35], 8.x.1965, 1 female (USNM) [Student Collection - WT 383]; Georgia: Perry [32.46, -83.73], 9.vii.1945 (P.W. Fattig coll.), 1 female (USNM).

Diagnosis. Lepidophora lutea is similar to L. lepidocera (see diagnosis of L. lepidocera) but differs in having notopleuron with long white hairs; femur I with short and spatulate white scales on dorsal surface; tibia II with longitudinal row of bristles on anterodorsal, posterodorsal, posteroventral and anteroventral surfaces; abdominal tergite III with long brown hairs; abdominal segment IV with short yellow or yellowish white scales laterally; and abdominal segment V with tuft of long and spatulate yellow, yellowish white, or white scales laterally.

Redescription of maleLengths. Body 11.86–19.86 mm; wing 9.14–14.86 mm.

Head. As wide as thorax; ocellar tubercle polished dark brown, long and thin proclinate dark brown bristles; frons polished dark brown on upper ½, gray pruinescence on lower ½, long and thin dark brown bristles and short and spatulate dark brown scales around the antennal base; antenna (Fig. 2F) dark brown with gray pruinescence, base of scape light brown; scape 2.5 times longer than pedicel; pedicel longer than postpedicel; scape with short and spatulate dark brown scales, long on ventral surface, medium dark brown bristles; pedicel with short and thin dark brown scales, long on ventral surface, medium dark brown bristles and short and strong dark brown bristles on apex; postpedicel with sparse short and strong dark brown bristles on base, medium bristles on proximal ½ of outer surface, sparse short and thin dark brown scales on proximal 1/3 of outer surface; face light brown, gray pruinescence on lower and upper thirds, short, medium and long spatulate dark brown scales more concentrated on clypeus margin, sparse medium white scales above the uppermost portion of clypeus margin, tuft of short and spatulate white scales along the margin of eyes at height of antennal base, long dark brown bristles bordering the eyes on upper 2/3; proboscis polished dark brown, about 5/7 length of antenna with short dark brown bristles, labellum about 1/5 length of proboscis; palpus polished dark brown, about 2/5 length of proboscis, palpomere I longer than palpomere II; occiput dark brown with gray pruinescence, short and spatulate dark brown scales on upper and lower thirds along the margin of eyes, lighter long and thin posteriorly on lower 1/3, short and spatulate white scales on medial 1/3 along the margin of eyes, long posteriorly, long and spatulate yellowish white scales around the occipital foramen; lower surface of head with long white hairs.

Thorax. Pronotum polished dark brown with gray pruinescence, sparse long and thin dark brown bristles and long and spatulate dark brown scales, long and thin yellow scales laterally; scutum polished dark brown, margin slightly lighter, short brown hairs more concentrated on anterior ½, yellowish white covering scales; postfrontal lobe with long and thin yellowish white scales, long and strong dark brown bristles on anterior 1/3, medium on medial 1/3, short and thin on posterior 1/3; supra alar callus with long and thin yellowish white scales, long and strong dark brown bristles, sparse dark brown bristles around; postalar callus with long and strong and medium dark brown bristles, upper ½ with long and thin white scales, anterosuperior corner with tuft of long white hairs, anteroinferior corner with tuft of long and thin white scales; pleurae polished light brown with gray pruinescence; anepisternum and notopleuron with short and spatulate white scales, long and thin white scales, long white hairs; anepisternum grayish brown on upper 1/3 with long brown hairs; katepisternum grayish brown on lower ½ with long and thin white scales, long white hairs; anepimeron with short white hairs on lower 1/3; meropleurite with long white hairs on posterior 1/3; metepisternum with short white hairs on medial portion; metepimeron with sparse white hairs on lower 1/3; proepisternum, proepimeron, laterotergite and metanotum bare; tuft of short white hairs near the posterior spiracle; halter with stem brown and with short and thin yellowish white scales, knob yellowish white with sparse short and thin dark brown scales; scutellum polished dark brown, margin slightly reddish brown with short and spatulate dark brown and yellow scales, posterior margin with long and spatulate and long and thin yellow scales.

Zootaxa 3682 (1) © 2013 Magnolia Press · 27REVISION OF LEPIDOPHORA WESTWOOD

Wings (Fig. 3E). Brown, lighter areas on center of discal cell, center of distal ½ of r1 and r2+3 cells and along

the posterior margin; alula hyaline; short and spatulate dark brown scales on both surfaces of wing concentrated until the distal 1/3 of r1 and r2+3 and until the height of proximal ½ of discal cell and upper margin of anal cell; costal vein with two rows of toothed bristles that extend of proximal 1/3 at height of beginning of discal cell until the apex of R2+3 vein; costal vein and R1 with short dark brown bristles, small tuft of short and spatulate white and dark brown scales on base; costal vein and stem vein with long and spatulate white and dark brown scales on base; basicosta with tuft of long and spatulate yellow scales; anal cell closed, CuA2 and A1 meet on margin of wing; calypter whitish, dark brown spot on proximal 1/3, margin with long white hairs on proximal 2/3, short white scales on distal 1/3; r-m crossvein positioned on ½ of discal cell; small projection on curved apex of R2+3, R4, and just after

the bifurcation with R5 and before the bifurcation on apex of the r2+3. Legs. Polished light brown. Leg I. Coxa with gray pruinescence, long and thin white scales and long and short

white hairs, long and strong yellow bristles on distal 1/3 of anterodorsal surface; trochanter with sparse short and thin white scales; femur with short and spatulate scales, dark brown on anterior and ventral surfaces, white on dorsal and posterior surfaces, longitudinal row of short and strong dark brown bristles on posterodorsal surface, strong apical dark brown bristles on anteroventral and anterodorsal surfaces; tibia with short dark brown bristles more concentrated on anterior and ventral surfaces, short and spatulate scales, dark brown on anterior and dorsal surfaces, white on posterior and ventral surfaces, longitudinal row of short and strong dark brown bristles on posterodorsal, posterior and anterodorsal surfaces, strong apical dark brown bristles on dorsal (2), posterior (1), posterodorsal (1), anteroventral (1), ventral (1) and posteroventral (1) surfaces; tarsi with short and spatulate dark brown scales. Leg II. Coxa with gray pruinescence, long and thin white scales and long and short white hairs, long and strong dark brown bristles on distal 1/3 of anteroventral surface; trochanter with short and thin white scales, short white hairs; femur with short and spatulate scales, dark brown on anterior and dorsal surfaces, white on posterior and ventral surfaces, longitudinal row of short and strong dark brown bristles on anteroventral surface, strong brown bristle on distal 1/3 of posteroventral surface; tibia with short dark brown bristles, short and spatulate scales, dark brown on anterior and ventral surfaces, white on posterior and dorsal surfaces, longitudinal row of short and strong dark brown bristles on anterodorsal, posterodorsal, posteroventral and anteroventral surfaces, strong apical dark brown bristles on dorsal (3), posterior (1), posteroventral (1), ventral (1) and anteroventral (1) surfaces; tarsi with short and spatulate dark brown scales. Leg III. Coxa with gray pruinescence, long and thin white scales and long and short white hairs, long and strong yellow scales on distal 1/3 of posterodorsal surface; trochanter with short and thin white scales, short white hairs; femur with short and spatulate scales, dark brown on anterior and dorsal surfaces, white on posterior and ventral surfaces, longitudinal row of short and strong dark brown bristles on anteroventral and ventral surfaces, crown of strong apical dark brown bristles; tibia with short dark brown bristles and spatulate dark brown scales, longitudinal row of short and strong dark brown bristles on anteroventral, anterodorsal, posteroventral and posterodorsal surfaces, strong apical dark brown bristles on posterodorsal (1), posterior (1), posteroventral (1), ventral (2) and anterior (2) surfaces; tarsi with short and spatulate dark brown scales.

Abdomen. Polished brown, cylindrical, segments about the same width; segment I with long white hairs on anterior margin followed by long brown hairs; tergites II and III with long light brown hairs; sternites I and II with long yellowish white hairs, others sternites with long dark brown hairs; segment IV with short and spatulate white and yellowish white scales laterally; segment V with central tuft of long and spatulate yellowish white scales laterally; segments VI–VIII with tuft of long and spatulate dark brown scales laterally. Male terminalia (Figs. 11A–C). Gonocoxite 1.8 times longer than wide; gonocoxal apodeme short, apex slightly rounded in dorsal view and enlarged in lateral view; hypandrium subtriangular, anterior margin slightly concave; basiphallus with margin slightly straight and borders rounded in lateral view; distiphallus not exceeding the apex of epiphallus in lateral view; epiphallus with apex dilated dorsally in lateral view; epiphallus and distiphallus not exceeding the posterior margin of gonocoxites in lateral view; lateral aedeagal apodeme not exceeding the lateral margin of gonocoxites in dorsal view; ejaculatory apodeme long, rounded, anterior margin exceeding the anterior margin of gonocoxites in dorsal view.

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FIGURE 11A–E. L. lutea Painter, 1962: A–C male terminalia: (A) lateral view; (B) dorsal view; (C) ventral view; D–E female genitalia: (D) spermathecae; (E) furca.

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FemaleLengths. Body 11.86–19.14 mm; wing 7.86–13.71 mm.

Similar to male, except: color of body usually darker; head with the same width of thorax; frons with medium dark brown bristles on lower ½; base of scape with short and spatulate white scales; color of scales of occiput usually white. Female genitalia (Figs. 11D–E). Sperm pump less than 1/5 length of spermathecal duct; tubules on proximal ½ of spermathecal bulb; tubules on proximal ½ of apical spermathecal duct; furca with bars extended anteriorly and posteriorly, slightly extended laterally.

Geographical Records (Fig. 4)

Nearctic. Canada (Ontario), United States (Florida, Georgia, Illinois, Indiana, Louisiana, Maine, Maryland, Michigan, Minnesota, Missouri, New Jersey, New York, Ohio, Pennsylvania, Tennessee, Virginia).

Remarks. The following variations were observed in the sample series (males and females): clypeus margin without white scales; occiput with short and spatulate yellow scales on medial 1/3 along the margin of eyes, long posteriorly, long and spatulate yellow scales around the occipital foramen; yellow scales on the wing.

Lepidophora secutor Walker, 1857(Figs. 2G, 3F, 6C, 12)

Lepidophora secutor Walker, 1857: 145. Holotype male, BRAZIL: Amazonas or Pará (BMNH). Osten Sacken, 1887: 160; Painter, 1925: 119–121, 125; Paramonov, 1949: 631, 633, 638, 640; D’Andretta and Carrera, 1952: 296; Hull, 1973: 238; Painter and Painter, 1974: 89, 92; Painter et al.,1978: 38.21; Evenhuis and Greathead, 1999: 209.

Type material examined. Holotype: BRAZIL: Amaz [Amazonas or Pará], 1 male (BMNH). Other material examined. VENEZUELA: Amazonas: Rio Mavaca Camp [2.03, -65.10], 65º06' W 2º2' N,

150 m., 16–27.iii.1989 (D.A. Grimaldi coll.), 1 male (AMNH) [Phipps FUDECI Exped. By American Museum of Natural History]; COLOMBIA: Amazonas: Rio Caquetá [1.92, -76.67], 9.x.1956 (D.J. Taylor coll.), 1 male (BMNH) [B.M. 1954–110]; BRAZIL: Amazonas: São Gabriel da Cachoeira [-0.13, -67.09], Rio Negro, 10.ix.1927 (J.F. Zikán coll.), 1 female (MZUSP); Pará: 1954 (W.M. Mann coll.), 1 female (USNM) [W.M. Mann, Stanford Expedition, USNM Bombyliidae Collection]; Bragança-Ajuruteua [-1.05, -46.77], 1–4.ix.1988 (F.F. Ramos coll.), 1 female (MPEG) [Armadilha 1,6m suspensa]; Ponta de Pedras [-1.39, -48.87], 06.iii.1979 (M.F. Torres coll.), 1 male (MPEG); Belém [-1.46, -48.50], v.1924 (F.X. Williams coll.), 1 female (USNM); São Domingos do Capim, SESP, Rio Capim [-1.67, -47.78], 2 females (MPEG); Amazonas: Manaus, ZF 3, km 23 [-2.40,-59.87], 01.ix.1987 (M.V. Garcia coll.), 1 female (USNM) [3]; idem, BR 174, km 45 [-2.57, -60.03], 9.iv.1982 (E.L. Oliveira coll.), 1 male (INPA); idem, Est. Aleixo km 4 [-3.11, -60.02], 21.i.1976 (W.E. Kar coll.), 1 female (INPA) [0235]; idem, BR [-2.57, -60.03], x.1994 (Henriques coll.), 1 female (INPA); idem, Campus Univers. [-3.11, -60.02], 9.xii.1979 (J.A. Rafael coll.), 1 female and 1 male (INPA); idem, idem, 31.iii.1979 (J.A. Rafael coll.), 1 female (INPA); idem, idem, 11.vii.1979 (J.A. Rafael coll.), 1 female (INPA) [AR: Malaise]; idem, idem, 9.xii.1978 (J.A. Rafael coll.), 1 female (INPA); idem, idem, 9.ix.1978 (J.A. Rafael coll.), 1 female (INPA) [AR: Malaise]; idem, idem, 2.xii.1978 (J.A. Rafael coll.), 2 males (INPA); idem, idem, 25.xi.1978 (J.A. Rafael coll.), 1 male (INPA); idem, idem, 4.xi.1978 (J.A. Rafael coll.), 1 male (INPA) [AR: Malaise]; capoeira ton stubbst. 101?, 28.x.1924 (Melim S.V. coll.?), 1 male (USNM) [Amaz. Exp. 1923-25]; Iranduba, Rio Solimões, Ilha da Manchataria [-3.25, -60.10], 16.x.1991 (M.V. Garcia coll.), 1 female (INPA) [735–2]; idem, 29.xi.1991 (M.V. Garcia coll.), 1 female (INPA) [911–1]; Rio Solimões [-3.25, -58.97], 3º15' S 58º58' W, 16.x.1991 (M.V. Garcia coll.), 1 male (USNM) [736–2]; Pará: Tucuruí [-3.70, -49.70], Rio Tocantins, Canoal, 28–31.iii.1984, 1 female (MPEG) [Armadilha 1,6m suspensa]; PERU: LOCALITY UNKNOWN: 8.viii.22, 1 female (ZMHB) [Rictiriani G. Tessmann S.G.–Lepidophora secutor Wlk. det. J.C. Hall–Zool. Mus. Berlin]; Loreto: Iquitos [-3.75, -73.25], 28.i.1924, 1 female (AMNH) [F. locu–H. Bassler Collection Acc. 33591]; BRAZIL: Pará: Serra Norte, Rio Salobo [-5.90, -49.88], 24.ii.1984 (M.F. Torres coll.), 1 male (MPEG); PERU: Amazonas: Pucallpa [-6.14, -78.38], 10.i.1964 (J. Schunke coll.), 1 male (BMNH) [B.M. 1964–686]; BRAZIL: Acre: Senador Giomard, Rio Iquiri [-10.17, -67.83], 1 male (MZUSP) [63222–Acre Iquiri Exp. Dep. Zool.]; Rondônia: Ji-Paraná [-10.83, -61.97], 15.viii.1984

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(R.B. Neto coll.), 1 female (MPEG); Mato Grosso: Ribeirão Cascalheira [-12.83, -51.78], 12º50’S 51º47’W, 23.iii.1968 (O.W. Richards coll.), 1 female (BMNH) [B.M. 1968–260]; Goiás: Campinas [-14.33, -49.13], 1936 (R. Spitz coll.), 1 female (MZUSP) [161]; Mato Grosso: Cáceres [-16.06, -57.68], 11.xii.1984 (C. Elias leg), 1 female (DZUP) [POLONOROESTE–DPTº ZOOL. UF–PARANÁ]; Espírito Santo: Guarapari [-20.67, -40.50], ix.1960 (M. Alvarenga leg), 1 male (DZUP) [DZUP 206490]; LOCALITY UNKNOWN: 21.xi.1982 (Ulysses Barbosa coll.), 1 male (INPA); 18.x.1982 (Ulysses Barbosa coll.), 1 male (INPA); 1 male (ZMHB) [Lepidophora secutor Wlk. Tevas – Zool. Mus. Berlin].

Diagnosis. Anterior margin of pronotum with long and thin dark scales; scutum with short golden, dark brown and white scales; posterior margin of scutellum with short golden scales; wings with distal 1/3 completely hyaline; femur I with short and spatulate brown scales on posterior surface; abdomen clavate; abdominal segment V with lateral long dark brown scales only on distal 1/3.

Redescription of maleLengths. Body 12.43–21.14 mm; wing 9.14–15.14 mm.

Head. As wide as thorax; ocellar tubercle dark brown with gray pruinescence, short and thin proclinate dark brown bristles; frons polished dark brown on upper ½, gray pruinescence on lower ½, long and thin dark brown bristles and short and spatulate dark brown scales around the antennal base; antenna (Fig. 2G) dark brown with gray pruinescence; scape 2.5 times longer than pedicel; pedicel slightly longer than postpedicel; scape and pedicel with short and spatulate dark brown scales, long on ventral surface, medium dark brown bristles; scape with long and thin dark brown bristles more concentrated on proximal 1/3; pedicel with short and strong dark brown bristles on apex; postpedicel with sparse short and strong dark brown bristles on base, row of long and thin dark brown scales on all outer surface, numerous and long on proximal 1/3, whose length exceeds the length of postpedicel; face light brown, gray pruinescence on lower 1/3, upper 1/3 with short, medium and long spatulate dark brown scales more concentrated on clypeus margin, dark brown bristles bordering the eyes; proboscis polished dark brown, slightly shorter than antenna, short and medium dark brown bristles except on apex of labellum, where they are short and white, labellum about ¼ length of proboscis; palpus polished brown with gray pruinescence, about ½ length of proboscis, palpomere I longer than palpomere II; occiput dark brown with gray pruinescence, short and spatulate dark brown scales on upper and lower thirds along the margin of eyes, long and thin posteriorly on lower 1/3, short and spatulate light brown and white scales on medial 1/3 along the margin of eyes, long and thin white scales around the occipital foramen; lower surface of head with long light brown hairs.

Thorax. Pronotum dark brown with long and thin dark brown bristles, anterior margin with long and thin dark brown scales, long and thin white and brown scales laterally; scutum dark brown with gray pruinescence, margins slightly lighter, short brown hairs, dark brown covering scales, anterior 1/3 with short white covering scales, medial and posterior thirds with short golden covering scales; postfrontal lobe with long and thin white scales on lower ½, long and strong dark brown bristles on anterior 1/3, medium on medial 1/3, short and thin on posterior 1/3; supra alar callus with long and thin brown scales, long and strong dark brown bristles, sparse dark brown bristles around; postalar callus with long and strong and medium dark brown bristles, long and thin white scales, lower ½ with long light brown hairs; pleurae brown with gray pruinescence; anepisternum with short and spatulate white scales, upper 1/3 with long and thin white scales, anterior and posterior thirds with long and thin brown scales, lower ½ with long brown hairs; katepisternum with long and thin brown scales, long brown hairs on posterior 2/3; meropleurite with long brown hairs; metepisternum with short white and light brown hairs; metepimeron with short white hairs on medial portion of lower ½; notopleuron, proepisternum, proepimeron, anepimeron, laterotergite and metanotum bare; tuft of short light brown hairs near the posterior spiracle; halter with short and thin brown and white scales on stem, knob yellowish white with sparse short and thin dark brown scales; scutellum polished dark brown, short and spatulate dark brown scales, posterior margin with short and spatulate golden scales and long and thin white scales.

Wings (Fig. 3F). Dark brown until the 5/6 of r1 cell and until at height of ½ of r5, apex and posterior margin lighter; alula hyaline; short and spatulate dark brown scales on both surfaces of wing concentrated to proximal ½ of r1, r5 and discal cells, distal 1/3 of cell anal and alula; costal vein with two row of toothed bristles that extend from ½ of wing until just before the apex of r1; costal vein and R1 vein with short dark brown bristles, short and spatulate dark brown scales, long on base; stem vein with long and spatulate dark brown scales on base; basicosta with tuft of long and spatulate dark brown scales; anal cell closed, CuA2 and A1 meet on margin of wing; calypter whitish brown, small dark brown spot on center of distal 1/3, margin with long white and brown hairs on proximal 2/3, short brown scales on distal 1/3; r-m crossvein positioned on ½ of discal cell.

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Legs. Polished light brown. Leg I. Coxa with long and thin dark brown and white scales, long brown hairs; trochanter with sparse short and thin brown scales, short brown hairs; femur with short and spatulate scales, dark brown on anterior, dorsal and sparse on ventral surfaces, white on posterior surface, long and thin brown scales on posterior surface, long brown hairs on posterior and ventral surfaces, longitudinal row of short and strong dark brown bristles on posterodorsal surface, strong apical dark brown bristles on anteroventral and anterodorsal surfaces; tibia with short dark brown bristles more concentrated ventrally, short and spatulate scales, dark brown on anterior, dorsal and ventral surfaces, white on posterior surface, longitudinal row of short and strong dark brown bristles on posterodorsal and posterior surfaces, strong apical dark brown bristles on dorsal (2), ventral (2), posteroventral (1) and posterior (1) surfaces; tarsi with short and spatulate dark brown scales. Leg II. Coxa with long and thin brown and white scales and long brown hairs, long and strong dark brown bristles on distal 1/3 of anterodorsal surface; trochanter with sparse short and thin brown scales, short brown hairs; femur with short and spatulate scales, dark brown on anterior, dorsal and sparse on ventral surfaces, white on posterior surface, long brown hairs on posterior and ventral surfaces, longitudinal row of short and strong dark brown bristles on distal ½ of anteroventral surface, strong apical dark brown bristle on posteroventral surface; tibia with short dark brown bristles, short and spatulate scales, dark brown on anterior, dorsal and ventral surfaces, white on posterior surface, longitudinal row of short and strong dark brown bristles on anterodorsal, posterodorsal, posterior and anteroventral surfaces, strong apical dark brown bristles on dorsal (2), ventral (2), posteroventral (1), posterior (1), anteroventral (1) and anterior (1) surfaces; tarsi with short and spatulate dark brown scales. Leg III. Coxa with long and thin white and light brown scales and long brown hairs, long and strong dark brown bristles on distal 1/3 of posterodorsal and anterodorsal surfaces; trochanter with sparse short and thin brown scales, short brown hairs; femur with short and spatulate dark brown scales, longitudinal row of short and strong dark brown bristles on anterodorsal and ventral surfaces, crown of strong apical dark brown bristles; tibia with short dark brown bristles, short and spatulate dark brown scales, longitudinal row of short and strong dark brown bristles on anterodorsal, posteroventral, posterior, posterodorsal and anteroventral surfaces, strong apical dark brown bristles on ventral (2), anteroventral (2), dorsal (1) and posteroventral (1) surfaces; tarsi with short and spatulate dark brown scales.

Abdomen. Polished dark brown, clavate, segments II–IV much narrower than others; segment I with long white hairs on anterior margin followed by dark brown and light brown hairs until the anterior ½; segment II with long brown hairs laterally and on sternite; segments I–IV with short and spatulate white scales laterally; segments VI–VIII and distal 1/3 of segment V with tuft of long and spatulate dark brown scales laterally. Male terminalia(Figs. 12A–C). Gonocoxite 2.0 times longer than wide; gonocoxal apodeme long, apex rounded in dorsal view; hypandrium subtriangular, anterior margin concave; basiphallus rounded in lateral view; distiphallus not exceeding the apex of epiphallus nor posterior margin of gonocoxites in lateral view; epiphallus reaching the posterior margin of gonocoxites in lateral view; lateral aedeagal apodeme exceeding the lateral margin of gonocoxites in dorsal view; ejaculatory apodeme long, rounded, exceeding the anterior margin of gonocoxites in dorsal view.

FemaleLengths. Body 11.28–18.28 mm; wing 7.14–13.28 mm.

Similar to male, except: frons with medium dark brown bristles on lower ½; face with tuft of short and spatulate white scales along the margin of eyes at height of antennal base; occiput with short and spatulate white scales on medial 1/3 along the margin of eyes; lower surface of head with long white hairs. Female genitalia (Figs. 12D–E). Sperm pump 1/5 length of spermathecal duct; tubules on proximal ½ of spermathecal bulb; tubules on proximal 2/3 of apical spermathecal duct; furca with enlarged bars anteriorly and laterally.

Geographical Records (Fig. 4)

Neotropical. Brazil (Acre, Amazonas, Espírito Santo, Goiás, Mato Grosso, Pará, Rondônia, Roraima), Colombia (Amazonas), Guyana, Peru (Amazonas, Loreto), Venezuela (Amazonas).

Remarks. The following variations were observed in the sample series (males and females): face with gray pruinescence on upper and lower thirds, tuft of short and spatulate white scales along the margin of eyes at height of antennal base; proboscis with gray pruinescence; lower side of head with dark brown, or white and brown hairs; anal cell open.

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FIGURE 12A–E. L. secutor Walker, 1857: A–C male terminalia: (A) lateral view; (B) dorsal view; (C) ventral view; D–Efemale genitalia: (D) spermathecae; (E) furca.

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Lepidophora trypoxylona Hall, 1981 (Figs. 2H, 3G, 6D–E, 13)

Lepidophora trypoxylona Hall, 1981: 161. Holotype male, COSTA RICA: Guanacaste (CAS). Coville et al., 2000: 28, 39–41, 47; Evenhuis and Greathead, 1999: 209.

Type material examined. Holotype: COSTA RICA: Guanacaste: 15 km SW Bagaces [10.53, -85.25], Comelco, 25.ii.1975 (R. Coville coll.), 1 male (CAS) [California Academy of Sciences type n° 13728, trap n° M402C29C3, emerged 19.iii.1975, Host: Trypoxylon (Trypargilum) tenoctitlan]; paratypes: COSTA RICA: Heredia: La Selva, 4 km SE Puerto Viejo [10.47, -84.02], 26.vii.1980 (R.E. Coville coll.), 1 female (CAS) [ex Trap-nest coll., emerged 21.viii.1980, Trap nº C201Fcl]; idem, 16.vii.1980 (R.E. Coville coll.), 1 female (USNM) [ex Trap-nest coll., emerged 17.viii.1980, Trap nº CIFC3]; Guanacaste: 4 km NW Cañas [10.43, -85.10], La Pacifica, 14.ii.1975, (R. Coville coll.), 1 male (CAS) [emerged 10.iii.1975–trap nº M507cl].

Diagnosis. Lepidophora trypoxylona is similar to L. culiciformis (see diagnosis of L. culiciformis) but differs in having frons with spatulate dark brown, translucent and white scales; scape and pedicel with short and thin dark brown bristles; basicosta with tuft of long brown and white scales; coxa II with long light and dark hairs; coxa III with bristles on anterior and posterodorsal surfaces; femur I with apical bristles on dorsal and anterodorsal surfaces; femur III with brown and white scales on posterior surface; anterior margin of abdominal segment I with long dark brown hairs; and tergite VI with central tuft of short and spatulate white scales.

Redescription of maleLengths. Body 16.43–16.57 mm; wing 13.00–13.86 mm.

Head. Wider than thorax; ocellar tubercle dark brown with gray pruinescence, medium and thin proclinate dark brown bristles; frons polished dark brown on upper ½, brown with gray pruinescence on lower ½, long and thin dark brown bristles and short and spatulate dark brown and white scales around the antennal base, some translucent like white with reflected light; antenna (Fig. 2H) polished dark brown with gray pruinescence; scape 2.7 times longer than pedicel; pedicel slightly shorter than postpedicel; scape and pedicel with short and spatulate dark brown scales, long on ventral surface, short and thin dark brown bristles; pedicel with short and strong dark brown bristles on apex; postpedicel with sparse short and strong dark brown bristles on base, row of long and thin dark brown scales on proximal 2/3 of outer surface, numerous and long on proximal ½, whose length does not exceed the length of postpedicel; face polished light brown, gray pruinescence on lower 1/3, upper 1/3 with short, medium and long spatulate dark brown scales more concentrated on clypeus margin, medium spatulate light brown scales above the uppermost portion of clypeus margin, long dark brown bristles bordering the eyes; proboscis polished dark brown, about ½ length of antenna, short and sparse medium dark brown bristles except on apex of labellum, where they are short and light brown, labellum 2/5 length of proboscis; palpus brown, about 2/5 length of proboscis, palpomere I longer than palpomere II; occiput dark brown with gray pruinescence, short and spatulate dark brown scales on upper and lower thirds along the margin of eyes, long and thin posteriorly on lower 1/3, short and spatulate white and light brown scales on medial 1/3 along the margin of eyes, long and spatulate white scales around the occipital foramen; lower surface of head with long brown hairs.

Thorax. Pronotum polished brown with gray pruinescence, long and thin dark brown bristles, anterior margin with long and thin white scales followed by dark brown scales on all dorsal surface, long and thin dark brown and white scales laterally; scutum polished brown with gray pruinescence, margins light brown, short brown hairs, short white and brown covering scales; postfrontal lobe with sparse long dark brown bristles, long and thin white scales on lower 1/3, long and strong dark brown bristles on anterior 1/3, medium on medial 1/3, short and thin on posterior 1/3; supra alar callus with sparse long and thin white scales, long and strong dark brown bristles, medium dark brown bristles around; postalar callus with long and strong and medium dark brown bristles, lower 1/3 with long white hairs; pleurae polished light brown with gray pruinescence; anepisternum and notopleuron with short and spatulate white scales, long and thin white and light brown scales, long brown hairs; anepisternum with dark brown spot on anterior 1/3, posteroinferior corner with tuft of long and thin brown and white scales; proepimeron with long and thin light brown and white scales on posterosuperior corner; proepisternum with sparse medium dark brown bristles, long and thin white scales; katepisternum with long and thin white and brown scales, long brown hairs on anterior and upper halves; meropleurite with long brown and light brown hairs; metepisternum with short white and brown hairs on medial portion; anepimeron, metepimeron, laterotergite and metanotum bare; tuft of

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short brown hairs near the posterior spiracle; halter with stem yellowish brown and short and thin white scales, knob yellowish white with short and thin dark brown scales; scutellum polished brown with short and spatulate brown scales, posterior margin with central tuft of short and spatulate white scales, long and thin white scales.

Wings (Fig. 3G). Brown, hyaline areas on apex of r1 cell, distal 1/3 of r2+3, r4, distal 1/3 of r5 and along the posterior margin; alula hyaline; short and spatulate dark brown scales on both surfaces of wing concentrated on upper 2/3 until before the apex of r1, r2+3 and br, proximal 1/3 of discal cell, proximal ½ of anal cell and alula; costal vein and R1 vein with short dark brown bristles, short and spatulate dark brown scales; costal vein with short and spatulate white scales on base, two rows of toothed bristles that extend from the distal ½ to R2+3; stem vein with long and spatulate dark brown and white scales on base; basicosta with tuft of long and spatulate dark brown and white scales; anal cell closed, CuA2 and A1 meeting on margin of wing; calypter whitish, margin dark brown with long white and dark brown hairs on proximal 2/3, short white scales on distal 1/3, small dark brown spot on proximal 1/3; r-m crossvein positioned just beyond the ½ of discal cell; small projection on apex of R2+3 and base of R4.

Legs. Polished yellowish brown with gray pruinescence. Leg I. Coxa with sparse long and thin brown and white scales, long dark brown hairs; trochanter with short and thin light brown and white scales, sparse short brown hairs; femur with short and spatulate scales, light brown and white on posterior and ventral surfaces, dark brown on anterior and dorsal surfaces, sparse long and thin light brown and white scales on posterior and ventral surfaces, long brown hairs, longitudinal row of short and strong dark brown bristles on posterodorsal surface, strong apical dark brown bristles on anterodorsal surface; tibia with short dark brown bristles more concentrated ventrally, short and spatulate scales, white on posterior surface, dark brown on anterior, dorsal and ventral surfaces, longitudinal row of short and strong dark brown bristles on posterodorsal and posterior surfaces, strong apical dark brown bristles on dorsal (2), ventral (1), posteroventral (1), posterior (1), posterodorsal (1) and anteroventral (2) surfaces; tarsi with short and spatulate dark brown scales, tarsomere I with longitudinal row of short and strong dark brown bristles on ventral surface. Leg II. Coxa with long and thin light brown and white scales and long dark brown and white hairs, long and strong dark brown bristles on distal 1/3 of anterodorsal surface; trochanter with short and thin light brown scales, sparse short brown hairs; femur with short and spatulate scales, brown on ventral surface, white on posterior surface, dark brown on anterior and dorsal surfaces, long brown hairs on ventral surface, longitudinal row of short and strong dark brown bristles on distal ½ of anteroventral surface, strong apical dark brown bristles on posteroventral surface; tibia with short dark brown bristles, short and spatulate scales, white on dorsal and posterior surfaces, brown on anterior and ventral surfaces, longitudinal row of short and strong dark brown bristles on anterodorsal, posterodorsal, posterior, posteroventral and ventral surfaces, strong apical dark brown bristles on dorsal (3), ventral (2), posterior (1), posteroventral (1), anteroventral (1) and anterior (1) surfaces. Leg III. Coxa with long and thin light brown and white scales and long brown hairs, long and strong dark brown bristles on distal 1/3 of posterodorsal and anterodorsal surfaces; trochanter with short and thin light brown scales, short brown and dark brown hairs; femur with short and spatulate scales, brown on anterior, dorsal and ventral surfaces, brown and white on posterior surface, longitudinal row of short and strong dark brown bristles on distal ½ of anteroventral and ventral surfaces, crown of apical dark brown bristles; tibia with short dark brown bristles, short and spatulate dark brown scales, longitudinal row of short and strong dark brown bristles on anterodorsal, posterodorsal, posterior, posteroventral and anteroventral surfaces, strong apical dark brown bristles on ventral (4), posteroventral (1), posterior (1), anteroventral (1), anterior (1), anterodorsal (1) and posterodorsal (1) surfaces; tarsi with short and spatulate dark brown scales.

Abdomen. Polished light brown, cylindrical, segments about the same width; segment I with long dark brown hairs on anterior margin, short white covering scales on posterior margin of tergite; sternites I and II with long brown hairs; segments I–V with short and spatulate white scales and long brown hairs laterally; tergite VI with central tuft of short and spatulate white scales; segments VI–VIII with tuft of long and spatulate dark brown scales laterally. Male terminalia (Figs. 13A–C). Gonocoxites 2.5 times longer than wide; gonocoxal apodeme long, apex rounded in dorsal view and directed ventrally in lateral view; hypandrium subtriangular, anterior margin straight; basiphallus rounded in lateral view; distiphallus extending until just before the apex of epiphallus in lateral view; epiphallus with apex dilated dorsally; epiphallus and distiphallus not exceeding the posterior margin of gonocoxites in lateral view; lateral aedeagal apodeme not exceeding the lateral margin of gonocoxite in dorsal view; ejaculatory apodeme long, rounded, exceeding the anterior margin of gonocoxites in dorsal view.

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FIGURE 13A–E. L. trypoxylona Hall, 1981: A–C male terminalia: (A) lateral view; (B) dorsal view; (C) ventral view; D–Efemale genitalia: (D) spermathecae; (E) furca.

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FemaleLengths. Body 16.43–17.43 mm; wing 11.43–12.86 mm.

Similar to male, except: frons only with dark brown and translucent scales, lower ½ with medium dark brown bristles; lower surface of head with long white and brown hairs; wings dark brown. Female genitalia (Figs. 13D–E). Sperm pump 1/3 length of spermathecal duct; tubules on proximal ½ of spermathecal bulb; tubules on apical spermathecal duct, except for a small area above proximal ½; furca with bars extended dorsally and laterally, slightly extended posteriorly.

Geographical Records (Fig. 4)

Neotropical. Costa Rica (Guanacaste, Heredia).

Lepidophora vetusta Walker, 1857(Figs. 2I, 3H, 6F, 14)

Lepidophora vetusta Walker, 1857: 145. Lectotype male designed by Painter and Painter (1962:52), MEXICO: Veracruz-Llave (BMNH). Williston, 1901: 294; Aldrich, 1905: 240; Painter, 1925: 119–120, 123, 125; Painter, 1939: 44–45; Paramonov, 1949: 631, 633, 638, 641; Painter and Painter, 1962: 50, 52; Painter and Painter, 1965: 426; Hull, 1973: 238; Painter et al.,1978: 38.21; Hall, 1981: 161; Evenhuis and Greathead, 1999: 209.

Type material examined. Lectotype: MEXICO: Veracruz-Llave: 1 male (BMNH) [684]. Other material examined. UNITED STATES: Texas: Harlingen [26.19, -97.70], 8.4?, 1 male (USNM)

[Painter, 1960, Walk BM (Homotype)]. MEXICO: LOCALITY UNKNOWN: (Bigot coll.?), 2 males (BMNH) [B.M. 1960-539]; Sinaloa: San Ignácio, Sierra de las Ventanas [25.92, -108.60], 2.000 ft, (Forrer. coll.), 4 females (BMNH) [B.M. 1904–85, Cent. America: F.D. Godman & O. Salvin]; San Luís Potosi: 5m S Valles [21.98, -99.02],(Hobart Smith coll.?), 1 male (USNM) [6.1332]; 12 mi E Xilitla [21.33, -98.97], elevation 800 ft., 31.vii.1962 (R.H. & E.M. Painter colls.), 1 female (USNM); Tamazunchale [21.27, -98.78], 19.vii.1946 (J. & D. Pallister colls.), 2 males (AMNH); idem, 20.viii.1967 (Gary F. Hevel coll.), 1 male (USNM); Jalisco: Chamela [19.53, -105.08], 30.ix.1985 (R.J. McGinley coll.), 1 male (USNM); Colima: 6 mi. S Colima [19.23, -103.72], elevation 1400 ft., 25.viii.1962 (R.H. & E.M. Painter colls.), 1 female and 1 male (USNM) [R.H. Painter Collection 1975]; Veracruz-Llave: Veracruz-Llave (=Vera Cruz) [19.20, -69.13], 1 male (BMNH) [5466]; idem, 7-28-8-11-56 [7–28.viii–xi.1956?] (R. & K. Dreisbach colls.), 1 male (USNM); Morelos: 62 mi. S Cuernavaca [18.92, -99.25], altitude 4400 ft., 13.v.1966 (R.H. & E.M. Painter colls.), 1 female (USNM) [R.H. Painter Collection 1975]; Veracruz-Llave: Atoyac [18.90, -96.77], (April, H.H.H. coll.), 1 male (BMNH); Guerrero: Huitzuco [18.30, -99.35], iv.1951 (W.G. Owens coll.), 1 male (USNM) [A.L. Melander Collection 1961, R.H. Painter Collection 1975]; 23km NNE Iguala [17.85, -100.37] Jet., 18.x.1986 (E. Fisher coll.), 2 males and 1 female (USNM) [Hwy. 95D]; Oaxaca: Valerio Trujano [17.77, -96.98], 28.vii1937 (M. Emberry coll.), 1 male (CAS); Guerrero: 32 mi. North Chilpancingo [17.55, -99.50] Hy 90 km. 225, 19.ix.1960 (Chas. H. Martin coll.), 1 male (CAS); Carrizal [17.27, -99.73], (W.G. Downs. coll.), 2 females (USNM) [ALMelander Collection 1961, R.H. Painter Collection 1975]; Chiapas: Município Tuxtla Guitierrez [16.75, -93.12], Mirador for Chicoasen Dam at N, end of Cañon El Sumidero, alt. 518 m, 9.ix.1976 (D.E. & J.A. Breedlove cols. - Cal. Acad. Sci.), 1 male (CAS); 5.6 mi SE Chiapa de Corzo [16.70, -93.00] 2500 ft., 16.viii.1966 (D.E. Breedlove & J. Emmel colls.), 1 female (CAS); Municipio de Acala [16.57, -92.80], 20km N Acala on rd. Along Rio Grijalva, 548 m, 30.vii.1981 (D.E. & P.M. Breedlove colls.); GUATEMALA: LOCALITY UNKNOWN: 1997?, 1 female (BMNH) [BMNH(E) 1997–240]; Chiquimula: Guatalon [14.67, -89.40], Santa Adelaida, 1000m, Mar Apr. 31 (iii–iv.1931?) (J. Bequaert coll.), 1 female (AMNH); Guatemala: 33 mi. NE Guatemala City [14.62, -90.53], Altitude 3000 ft, 9.ix.1967 (R.H. & E.M. Painter colls.), 1 female (CAS); Retalhuneu: 51–59 mi. W San Sebastian [14.57, -91.65], altitude 1200 ft., 14.ix.1967 (R.H. & E.M. Painter colls.), 2 males (USNM) [R.H. Painter Collection 1975]; 29 mi. W San Sebastian [14.57, -91.65], altitude 1400 ft.. 14.ix.1967 (R.H. & E.M. Painter colls.) 1 male (USNM) [R.H. Painter Collection 1975]; Escuintla: El Salto [14.30, -90.78], 1934 (F.X. Williams coll.), 1 male and 1 female (USNM); Escuintla/Obispo: La Providencia [14.20, -90.93], 1 female (USNM) [emRoiullard All?, R.H. Painter Collection 1975];

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Esteli: La Trinidad [12.97, -86.24], 609 m, 12.ix.1976 (Edward S. Ross coll.–Cal. Acad. Sci.), 1 female (CAS); Manágua: Managua [12.15, -86.27], X–XII.1950 (Swain coll.?), 1 female (AMNH); COSTA RICA: Guanacaste: Santa Rosa National Park [10.88, -85.77], 300m, x.1982 (D.H. Janzen & W. Hallwachs colls.), 1 female (AMNH); idem, 6–20.vii.1978 (D.H. Janzen coll.), 1 female (AMNH); idem, 300m, iii.1983 (D.H. Janzen & W. Hallwachs colls.), 1 female (AMNH); idem, 12.xii.1978–10.i.1979 (D.H. Janzen coll.), 1 female (AMNH); 3 mi. SE El Coco [10.55, -85.70], Altitude 400 ft, 15.viii.1967 (R.H. & E.M. Painter colls.), 1 female (USNM) [R.H. Painter 1975]; Alajuela: San Mateo, Higuito [9.95, -84.55], (Pablo Schild coll.), 4 females (USNM) [R.H. Painter Collection 1975].

Diagnosis. Lepidophora vetusta is similar to L. cuneata (see diagnosis of L. cuneata) but differs in having frons with spatulate white and light brown scales; pedicel brown with orange base; light brown scales around the occipital foramen; wings with brown and white scales on both surfaces; femur I with white scales on anterior surface, brown on posterior surface; tibia I with longitudinal row of bristles on anterodorsal, posterodorsal, posterior and posteroventral surfaces; tibia II with white scales on anterior and ventral surfaces; abdominal segments VI and VII with short dark brown scales laterally; and abdominal segment VIII with tuft of long and pedunculate dark brown scales laterally.

Redescription of maleLengths. Body 12.14–20.57 mm; wing 7.86–15.00 mm.

Head. As wide as thorax; ocellar tubercle dark brown with gray pruinescence, long and thin proclinate dark brown bristles; frons dark brown on upper ½, light brown with gray pruinescence on lower ½, short and thin dark brown bristles and short and spatulate light brown scales around the antennal base, slightly longer laterally; antenna (Fig. 2I) polished light brown with gray pruinescence, base of scape and pedicel orange; scape 2.6 times longer than pedicel; pedicel shorter than postpedicel; scape and pedicel with short and spatulate dark brown scales, long on ventral surface, medium dark brown bristles; scape with long and thin dark brown bristles on proximal 1/3; pedicel with short and strong dark brown bristles on apex; postpedicel dark brown with gray pruinescence, sparse short and strong dark brown bristles on base, row of long and thin dark brown scales on proximal 2/3 of outer surface, numerous and long on proximal ½, whose length does not exceed the length of postpedicel; face polished light brown, gray pruinescence on lower 1/3, upper 1/3 with short, medium and long spatulate dark brown scales more concentrated on clypeus margin, short light brown scales above the uppermost portion of clypeus margin, long dark brown bristles bordering the eyes; proboscis polished dark brown, 2/3 length of antenna, short and sparse medium dark brown bristles except on apex of labellum, where they are short and light brown, labellum 2/5 length of proboscis; palpus brown, 3/5 length of proboscis, palpomere I longer than palpomere II; occiput dark brown with gray pruinescence, short and spatulate dark brown scales on upper and lower thirds along the margin of eyes, short and spatulate light brown scales on medial 1/3 along the margin of eyes, long posteriorly around the occipital foramen; lower surface of head with long brown hairs.

Thorax. Pronotum polished dark brown with gray pruinescence, long and thin dark brown bristles, anterior margin with short and spatulate light brown and white scales, long and thin white scales laterally; scutum polished dark brown with gray pruinescence, margins orange brown especially on anterior 1/3, short brown hairs, short white covering scales; postfrontal lobe with sparse long dark brown bristles, long and thin white scales on lower 1/3, long and strong dark brown bristles on anterior 1/3, medium on medial 1/3, short and thin on posterior 1/3; supra alar callus with long and thin white scales, long and strong dark brown bristles, sparse medium dark brown bristles around; postalar callus with long and strong and medium dark brown bristles, long and thin white scales, long light brown and white hairs; pleurae orange brown with gray pruinescence; anepisternum and notopleuron with short and spatulate and long and thin white scales, long brown hairs; anepisternum with dark brown spot on anterior 2/3, posteroinferior corner with long yellow hairs; proepimeron with long and thin white scales on posterosuperior corner; proepisternum with sparse medium dark brown bristles, long and thin white scales; katepisternum with polished brown spot on lower ½, long and thin white scales, long brown hairs on upper ½; meropleurite with long brown hairs on posterior 1/3; metepimeron with short white and brown hairs; anepimeron, metepisternum, laterotergite and metanotum bare; tuft of short white and light brown hairs near the posterior spiracle; halter with stem yellowish brown and short and thin white scales, knob yellowish white with sparse short and thin dark brown scales; scutellum polished brown with short and spatulate brown scales, posterior margin with long and spatulate and long and thin white scales.

Wings (Fig. 3H). Dark brown, lighter areas on center of r2+3, discal and m2 cells, hyaline areas along the

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posterior margin and apex; alula hyaline; short and spatulate dark brown and white scales on both surfaces of wing following the dark brown area; costal vein with two rows of toothed bristles that extend from the ½ of wing at height of beginning of discal cell to the R4 vein; costal vein and R1 with short dark brown bristles, short and spatulate white and dark brown scales on base; stem vein with short and spatulate white and dark brown scales on base; basicosta with tuft of long and spatulate brown scales; anal cell closed, CuA2 and A1 meeting just before reaching the margin of wing; calypter whitish, small dark brown spot on proximal 1/3, margin with long white and brown hairs on proximal 2/3, short white scales on distal 1/3; r-m crossvein positioned just beyond the ½ of discal cell.

Legs. Orange brown with gray pruinescence. Leg I. Coxa with long and thin light brown scales and long brown hairs, long and strong dark brown bristles on distal 1/3 of anterodorsal surface; trochanter with short and thin light brown scales, sparse short brown hairs; femur with short and spatulate scales, brown on posterior and dorsal surfaces, white on anterior and ventral surfaces, long brown hairs on posterior and ventral surfaces, longitudinal row of short and strong dark brown bristles on posterodorsal surface, strong apical dark brown bristles on anteroventral and anterodorsal surfaces; tibia with short dark brown bristles more concentrated ventrally, short and spatulate white scales on posterior surface, longitudinal row of short and strong dark brown bristles on anterodorsal, posterodorsal, posterior and posteroventral surfaces, strong apical dark brown bristles on dorsal (2), ventral (2), posteroventral (1) and posterior (1) surfaces. Leg II. Coxa with long and thin light brown and white scales and long brown hairs, long and strong dark brown bristles on distal 1/3 of anterodorsal surface; trochanter with short and thin light brown scales, sparse short brown hairs; femur with short and spatulate scales, brown on anterior and dorsal surfaces, white on posterior and ventral surfaces, long brown hairs on posterior and ventral surfaces, longitudinal row of short and strong dark brown bristles on distal ½ of anteroventral surface, strong apical dark brown bristle on posteroventral surface; tibia with short dark brown bristles, short and spatulate white scales, longitudinal row of short and strong dark brown bristles on anterodorsal, posterodorsal, posterior, posteroventral and anteroventral surfaces, strong apical dark brown bristles on dorsal (3), ventral (2), posterior (1), posteroventral (1), anteroventral (1) and anterior (1) surfaces. Leg III. Coxa with long and thin light brown scales and long brown hairs, long and strong dark brown bristles on distal 1/3 of anterodorsal and posterodorsal surfaces; trochanter with short and thin light brown scales, sparse short brown hairs; femur with short and spatulate brown scales, longitudinal row of short and strong dark brown bristles on distal ½ of anteroventral and ventral surfaces, apical crown of strong dark brown bristles; tibia with short dark brown bristles, short and spatulate brown scales, longitudinal row of short and strong dark brown bristles on anterodorsal, posterior, posterodorsal, posteroventral and anteroventral surfaces, strong apical dark brown bristles on ventral (2), posteroventral (1), posterior (2), anteroventral (1), anterior (1), anterodorsal (1) and posterodorsal (1) surfaces; tarsi with short and spatulate dark brown scales.

Abdomen. Dark brown, margins of tergites light brown, slightly clavate, segments III and IV narrower than others; segment I with long brown hairs on anterior margin followed by long dark brown hairs until the anterior ½; segments I–IV with short and spatulate white scales laterally; segment VIII with tuft of long and pedunculate dark brown scales on sternite and laterally. Male terminalia (Figs. 14A–C). Gonocoxites 2.5 times longer than wide; gonocoxal apodeme long, apex rounded in dorsal view and dilated in lateral view; hypandrium subtriangular, anterior margin slightly concave; basiphallus rounded in lateral view; distiphallus not exceeding the apex of epiphallus and reaching the posterior margin of gonocoxites in lateral view; epiphallus with apex dilated dorsally surpassing the posterior margin of gonocoxites in lateral view; lateral aedeagal apodeme not exceeding the lateral margin of gonocoxites in dorsal view; ejaculatory apodeme long, spatulate, exceeding the anterior margin of gonocoxites in dorsal view.

FemaleLengths. Body 11.71–19.57 mm; wing 7.71–14.00 mm.

Similar to male, except: frons with tuft of short and spatulate white scales near the margin of eyes at height of antennal base, lower ½ with medium dark brown bristles and short and spatulate white and light brown scales, slightly longer laterally at height of antennal base; antenna dark brown with gray pruinescence; short and medium spatulate white scales above the uppermost portion of clypeus margin. Female genitalia (Figs. 14D–E). Sperm pump about 1/5 length of spermathecal duct; tubules on proximal ½ of spermathecal bulb; tubules on apex and proximal 2/3 of apical spermathecal duct; furca with bars extended anteriorly and laterally, slightly extended posteriorly.

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FIGURE 14A–E. L. vetusta Walker, 1857: A–C male terminalia: (A) lateral view; (B) dorsal view; (C) ventral view; D–Efemale genitalia: (D) spermathecae; (E) furca.

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Geographical Records (Fig. 4)

Nearctic. United States (Texas), Mexico (Colima, Guerrero, Jalisco, Morelos, Nayarit, San Luis Potosi, Sinaloa, Sonora, Veracruz-Llave). Neotropical. Costa Rica (Alajuela, Guanacaste), Guatemala (Chiquimula, Escuintla, Esteli, Guatemala, Managua, Retalhuneu), Mexico (Chiapas, Oaxaca), Nicaragua.

Remarks. The following variations were observed in the sample series (males and females): frons with long and thin bristles; proboscis orange brown on proximal ½ and polished dark brown on distal ½; basicosta with some transparent scales, like faded; anal cell open.

The type locality was erroneously given by Walker (1857) as “Valley of Amazon”. Painter and Painter (1962), upon examination of type material and designation of the lectotype, corrected the location to Veracruz-Llave (Mexico), following the single label affixed to the type with a handwritten ‘V. Cruz’. Evenhuis and Greathead (1999) stated, based on the original description, the “Valley of Amazon” as the type locality, considering Veracruz-Llave proposed by Painter and Painter (1962), an error. Based on our observation of the lectotype deposited in BMNH, we consider the type locality of L. vetusta to be correct as Veracruz-Llave (Mexico), and “Valley of the Amazon” an error, corroborating Painter and Painter (1962).

Acknowledgments

The authors thank Dr. David Grimaldi and Dr. Erica McAlister for the kind assistance during the visits of CJEL respectively to the American Museum of Natural History (New York) and to the Natural History Museum (London). Dr. Neal Evenhuis (USNM [at Bishop Museum]), Keith Arakaki (USNM [at Bishop Museum]), Dr. Augusto Henriques (INPA), Dr. Peter Haase (SMFD), Irene Rademacher (SMFD), Dr. Claudio Carvalho (DZUP), Dr. Joachim Ziegler (ZMHB), Dr. David Grimaldi (AMNH) and Dr. Charles Griswold (CAS) for loaning us specimens of Lepidophora. Thanks go to the reviewers, Dr. Christine Lambkin (Queensland Museum, Australia) and Dr. Neal Evenhuis (Bishop Museum, USA). Thanks also to the Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq) (Proc. No. 563256/2010-9) and Fundação de Amparo à Pesquisa do Estado de São Paulo (FAPESP) (Proc. No. 2010/52314-0) for the support to the SISBIOTA Project, CNPq (Proc. No. 481024/2008-5) for the grant to CJEL and finally to Coordenação de Aperfeiçoamento de Pessoal de Nível Superior (CAPES) for the financial support to PFMR.

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