Taxonomic revision of the Neotropical genus ...

Zootaxa 3827 (2): 231–257 www.mapress.com /zootaxa / Copyright © 2014 Magnolia Press

Article

ISSN 1175-5326 (print edition)

ZOOTAXA

ISSN 1175-5334 (online edition)

http://dx.doi.org/10.11646/zootaxa.3827.2.6 http://zoobank.org/urn:lsid:zoobank.org:pub:D6C800AA-8127-41D1-ACEA-52254F7CE89D

Taxonomic revision of the Neotropical genus Arthropeina Lindner, 1949 (Diptera: Xylomyidae) DIEGO AGUILAR FACHIN1,3 & DALTON DE SOUZA AMORIM2 1

Graduate Course on Entomology, Departamento de Biologia, Faculdade de Filosofia, Ciências e Letras de Ribeirão Preto, Universidade de São Paulo, Av. Bandeirantes 3900, 14040–901, Ribeirão Preto, SP, Brasil. 2 Departamento de Biologia, Faculdade de Filosofia, Ciências e Letras de Ribeirão Preto, Universidade de São Paulo. 3 Corresponding author, e-mail: [email protected]

Abstract Five new species are herein described for the Neotropical genus Arthropeina previously known only from the type–species, A. fulva Lindner: A. colombiana, sp. nov., A. diadelothorax, sp. nov., A. lindneri, sp. nov., A. melanochroma, sp. nov. and A. pseudofulva, sp. nov. The new species are described and habitus, antenna, palpus, thorax, wing, and male and female genitalia, including genital fork and spermathecae, are illustrated. A key to the species of the genus is provided. The diagnosis of Arthropeina is emended to include these new species. Additionally, the genus is recorded for the first time for Guyana, Colombia, Ecuador, Peru and Bolivia. Key words: Arthropeina, biodiversity, Neotropics, taxonomy, Xylomyidae

Introduction Xylomyidae is a family of the Stratiomyomorpha and sister to the Stratiomyidae (Woodley, 1989; Woodley et al., 2009). The Stratiomyomorpha also includes the exclusively Neotropical family Pantophthalmidae (see Papavero, 2008, 2009). Xylomyids have a body length ranging from 4 to 14 mm and have quite varied coloration. They are easily recognized by the presence of spurs on the mid and hind tibiae, by the conical antennae, with most flagellomeres uniform in shape and color, by the elongated discal cell and by the cell m3 closed before the wing margin. Little is known about the biology of these flies, but frequently the males are associated with forest environments and the immature stages may occur under the bark of fallen trees (Webb, 1984; Woodley, 2009, 2011). A total of 138 species of Xylomyidae have been described for the world, placed in four genera (Woodley, 2011). From these, 11 species in three genera are recorded for the Neotropics: one species in Arthropeina Lindner, 1949; seven species in Solva Walker, 1859; and three species in Xylomya Rondani, 1861. Most of the other species of the family are Palearctic and Oriental, respectively with 46 and 62 described species (Woodley, 2011). Papavero & Artigas (1991) were the first to discuss the relationships between the genera of Xylomyidae in a phylogenetic perspective, using mainly female genitalia characters. More recently, Woodley (2011) proposed a new phylogenetic hypothesis for the Xylomyidae at the generic level, reinterpreting some of the characters available in the literature. In his proposal, the pair of clades (Solva + Arthropeina) and (Coenomyiodes + Xylomya) would be sister-groups.

The genus Arthropeina The genus Arthropeina is endemic to the Neotropical Region and before now included a single described species, Arthropeina fulva Lindner, 1949. The real diversity of the genus, however, is pretty large, as predicted by Woodley (2009, 2011). Accepted by N. Evenhuis: 12 Jun. 2014; published: 3 Jul. 2014

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Besides the original description of A. fulva by Lindner (1949), subsequent papers have addressed the genus. Papavero & Artigas (1991) illustrated the female genitalia, including the spermatheca morphology. Woodley (1999) made lectotype and paralectotype designations for the type material, while Woodley (2009, 2011) commented on the diagnosis of the genus within a phylogenetic discussion of the family (Woodley, 2011). The recognition of the genus was largely based on body color—primarily yellow, sometimes with shining black maculae—and the shape of the two apical flagellomeres, which form a stylus. The shape of the stylus was considered autapomorphic for the genus (Papavero & Artigas, 1991; Woodley, 2011). Furthermore, Papavero & Artigas (1991) proposed that a helicoid spermathecae with three “layers” or coils was autapomorphic for Arthropeina. This, however, was criticized by Woodley (2011), who commented that the blind duct illustrated by the former authors could be a sac ruptured during the preparation, not a derived condition, as proposed. A possible additional apomorphy mentioned by Woodley (2011) are the valves of the spermathecal ducts, illustrated by Papavero & Artigas (1991). This paper reviews the genus Arthropeina, describing five new Neotropical species and redescribing A. fulva, including illustrations of the antenna, palpus, thorax, spermatheca, genital fork, and male genitalia. The paper also adds some comments on the issue of the relationships within the genus and provides an identification key for the known species of Arthropeina.

Material & methods The material studied in this paper belongs to the following institutions: CSCA CNC DZUP IAvH MZUSP USNM

California State Collection of Arthropods, Sacramento, California, United States. Canadian National Collection of Insects, Arachnids and Nematodes, Ottawa, Canada. Departamento de Zoologia, Universidade Federal do Paraná, Curitiba, Brazil. Instituto de Investigación de Recursos Biológicos Alexander von Humboldt, Bogotá, Colombia. Museu de Zoologia da Universidade de São Paulo, São Paulo, Brazil. National Museum of Natural History, Washington, D.C., United States.

Terminology mainly follows Cumming & Wood (2009), except for some details of the genital fork, that follow Woodley (1981), and details of the spermatheca morphology and female reproductive tracts, that in some extent follow Artigas (1971). The body length is measured without the antennae. A previously undescribed structure of the female genitalia recognized in this study (e.g., Fig. 21) is referred to here as the membranous lobes (ml). The male and female genitalia were treated with 10% KOH at 40ºC for 50 minutes and the spermathecal preparations were treated at same temperature for 15–20 minutes, and both were temporarily mounted on cavity slides with glycerin. Antennae, wings and thorax were mounted on permanent slides, after dehydration procedure, and photographed using a Leica M16 stereomicroscope and a Leica DC 500 camera. The photos were assembled using the Auto-Montage Pro v5.02.0096 software, with some further editing with Adobe Photoshop CS4. Drawings (e.g., antenna, thorax, male genitalia) were made under camera lucida and redrawn using Adobe Illustrator CS5. Illustrations of the male genitalia were made with both dorsal and ventral views, including the epandrium in dorsal view, and the gonostylus in ventral and inner views. Additional illustrations include the female genitalia in dorsal view, the genital fork in ventral, dorsal and lateral views, and the female reproductive tracts with spermathecae in dorsal view. The dissections of the female reproductive tracts were made with a pin vise under a Leica M16 stereomicroscope, to remove fat and remaining eggs. The male and female genitalia are preserved in a microvial pinned with the each specimen. The map of the geographic distribution of the species was prepared using Google Earth 7.1.1.1580 (beta) and Quantum Gis 2.0.1-Dufour.

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FACHIN & AMORIM

FIGURES 1–7. Antennae of Xylomyidae species. 1. Solva sp., female. 2. Arthropeina fulva. 3. A. colombiana, sp. nov., female holotype. 4. A. diadelothorax, sp. nov., holotype. 5. A. lindneri, sp. nov., paratype. 6. A. melanochroma, sp. nov., female paratype. 7. A. pseudofulva, sp. nov., paratype. Abbreviations: flgm, flagellomere; ped, pedicel; scp, scape.

Arthropeina Lindner, 1949 (Figs. 2–89) Arthropeina Lindner, 1949: 789. Type-species: Arthropeina fulva Lindner, 1949 (mon.).

Diagnosis. Body color mostly yellow to reddish yellow. Antennae conically tapered towards apex; apical two flagellomeres (7–8) forming a stylus (e.g., Fig. 2), as long as or longer than the remaining of the flagellomeres. Hind femur without row of tubercles, frequently slender, not differing from the other femora. Membranous lobes present in the female genital fork (e.g., Fig. 15). Redescription. Male. Length: body, 6.5–7.0 mm; wing, 6.5–7.0 mm. Head. Ocellar tubercle black, only slightly prominent, short setae at vertex. Eyes black, bare. Upper frons concave, gradually widening ventrally. Upper part of face silver tomentose. Pilosity short on frons, erect on genal region, and on posterior part of head. Antenna narrowing towards apex, conical (e.g., Fig 2); scape slightly shorter than pedicel, reddish yellow with yellow setae; first flagellomere longer than subsequent flagellomeres except last; eighth flagelomere considerably longer than seventh, length ratio ranging from 2:1 to 5:1; flagellomeres 7–8 forming a stylus as long as or longer than rest of flagellomeres combined. Palpus elongate, basal (first) segment cylindrical, longer than wide; apical (second) segment oval, at least two times length of basal segment; pale hairs on both segments. Proboscis yellowish with pale hairs. Thorax (Fig. 8). Postpronotal lobes and notopleural strip frequently contrasting with color of scutum. Scutum and scutellum unicolorous or with dark bands or spots; pleura frequently with dark spots, pilosity short. Legs. Frequently yellow with some dark marks, or almost completely dark; pilosity on legs short, dense, yellowish, conforming to color of surface, except tarsomeres, mostly with brownish hairs. Wing. Venation relatively uniform among the species (e.g., Fig. 66). Veins brown to dark brown; C extending to M1. M2 and cell m3 not reaching wing margin. Cells bm, br and cup well defined. Cell cup not reaching cubital fork. Alula narrow, gradually widening towards apex. Abdomen. Pale to reddish yellow, with dark transverse bands on tergites; sternites weakly sclerotized; six to eight segments visible; pilosity conforming to color of surface, frequently FACHIN & AMORIM—REVISION OF ARTHROPEINA

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yellowish. Male genitalia. Gonocoxite rounded laterally, distal margin strongly projected; inner plate with acuminate projections ventrally to phallus (aedeagus + parameres) (e.g., Fig 10); gonocoxal apodeme absent. Gonostylus long and wide, varying in shape, but frequently with inner projections. Aedeagus elongate, completely enclosed by the parameres (e.g., Fig 10); ejaculatory apodeme short, not exceeding basal margin of gonocoxite; a pair of long, acuminate projections lateral to phallus (e.g., Fig 10). Epandrium quadrangular, wide, without lateral projections (surstylus) (e.g., Fig. 14). Hypoproct (S10) strongly developed, epiproct (T10) limited to small sclerotized plate dorsally; cercus well developed, wider on distal half. Female. Differs from male as follow. Length: body, 6.0–7.5 mm; wing, 6.0–8.0 mm. Female genitalia. Tergite 8 wide, rounded anteriorly. Sternite 8 more or less quadrate, weakly sclerotized distally; gonapophyses unsclerotized, with short lobes. Tergite 9 laterally more sclerotized, completely fused to sternite 9 (genital fork) anterolaterally (e.g., Figs. 15–17). Genital fork strongly sclerotized, anterior arm rounded basally; posterolateral processes clearly membranous, covering the genital opening distally; membranous lobes covering the genital fork ventrally (e.g., Fig. 15), with some variation in shape; projection of posterior margin frequently prominent; margin between the posterior process with a medial incision; genital opening varying from small to quite large, placed between projections of posterior margin of genital fork, with short microtrichia on bursa. Three spermathecae present (Figs. 56–59), lateral ones with moderately long ducts, median one large, with a short duct; common duct membranous, unpigmented; posterior part of lateral ducts more sclerotized, with a valve medially; spermathecal capsules circular, more or less ovoid, strongly sclerotized. Tergite 10 not well sclerotized. Sternite 10 well developed, more or less triangular, distal margin rounded. First segment of cercus longer than second segment. Geographic distribution. The genus Arthropeina is known only from the Neotropical region. Up to now, the genus was known only from the type-locality, Nova Teutônia, Santa Catarina, in southern Brazil. The geographic distribution of this genus is now greatly expanded, with species known from northern Brazil and north and northwest South America, reaching Guyana, Colombia, Ecuador, Peru and Bolivia (Fig. 92). These are the first records of the genus for these last five listed countries. Comments. The genus can be easily separated from the remaining xylomyid genera by having the antenna conically tapered to the apex, with the apical two flagellomeres (7–8) forming a stylus (e.g., Fig. 2), which is as long as or longer than the basal six flagellomeres—an autapomorphy of the genus. This condition is clearly distinct from that of Solva species (Fig. 1). Additionally, Arthropeina species do not present the row of tubercles ventrally on hind femora found in Solva (Figs. 90–91). The male genitalia of Arthropeina, illustrated for the first time in this paper, show great resemblances to the genitalia of Solva and Xylomya (Webb, 1984). The gonocoxites are well developed, distally projected, the gonostylus is wide and frequently elongate, and the aedeagus is entirely enclosed by the parameres, forming a triangular structure that narrows towards the apex. Some inner gonocoxite projections in Solva and Xylomya,—usually named as lateral (“interbasis” – Nagatomi & Tanaka, 1971), median and ventral parameres (Webb, 1984),—have uncertain homology in the genitalia of Arthropeina. Apparently, the “lateral paramere” may be the lateral projections of the aedeagus (e.g., Figs. 10, 39), while the “ventral paramere” may be homologous to the inner plate of the gonocoxite (e.g., Figs. 10, 39) that in Arthropeina has distal projections. A more rigorous understanding of homology in xylomyid male genitalia is still necessary to properly understand their correspondence to structures illustrated in detail by Webb (1984) and Nagatomi & Tanaka (1971). There is apparently some confusion on structures in Xylomyidae, especially on the inner projections of the gonocoxite, resulting in some difficulty to establish homology and character polarity in male genitalia of Arthropeina.

Arthropeina fulva Lindner (Figs. 2, 8, 9–20, 56, 60, 66–67, 78, 84) Arthropeina fulva Lindner, 1949: 790. Type-locality: Brazil, Santa Catarina, Nova Teutônia. Distr – Brazil (Santa Catarina). Ref.: Woodley, 2011: 498 (cat.).

Diagnosis. Body color mostly yellow to reddish yellow (Figs. 78, 84). Scutum homogeneously reddish yellow, no maculation (Fig. 60). Projection of posterior margin of genital fork short, inner margin oblique (Figs. 15, 17). Spermathecal capsule helicoid, with two coils (Fig. 18).


FACHIN & AMORIM

FIGURE 8. Arthropeina fulva, female. 8. Thorax, lateral view. Abbreviations: anepm, anepimeron; anepsta, anepisternum anterior; anepstp, anepisternum posterior; aprn, antepronotum; a spr, anterior spiracule; cx, coxa; hlt, halter; kepm, katepimeron; kepst, katepisternum; ltg, laterotergite; mr, meron; mtepm, metepimeron; mtg, mediotergite; mtkepst, metakatepisternum; mtn, metanotum; npl, notopleural suture; pprn, postpronotum; pprn lb, postpronotal lobe; p spr, posterior spiracule; prepm, proepimeron; prepst, proepisternum; presct, prescutum; sct, scutum; sctl, scutellum; st, sternite; tg, tergite; trn sut, transverse suture. Scale bar, 1 mm.

Material examined. BRAZIL, 1 ♀ (left antenna and left wing slide–mounted), Mato Grosso, Chapada dos Guimarães, 27.xi–05.xii.1983, Alice Yamamoto et al. (DZUP). 1 ♀, Minas Gerais, Botelhos, Córrego da Onça, 21º40’90”S 46º22’05”W, Malaise trap-mata, 07.xii.2008–06.i.2009, João Basso col. (MZUSP). 1 ♀ (slide–mounted), Cabo Verde, Fazenda da Cata, 21º27’11.04”S 46º20’52.8”W, 598m, Malaise trap 2-mata, 29.xi.2008–22.ii.2009, Amorim, Ribeiro, Falaschi & Oliveira col (MZUSP); 1 ♀, 21º27’11”S 46º20’52”W, 598m, Malaise, 03.xi.2010–19.ii.2011, D.S. Amorim & S.S. Oliveira col. (MZUSP). 1 ♀, Presidente Olegário, Fazenda Gigante, 18º31’S 46º18’W 1000m, Shannon, 02–09.i.2010, Ribeiro, Amorim, Silva & Berbert. (MZUSP). 1 ♀, Rio de Janeiro, Nova Iguaçu, Reserva Biológica do Tinguá, 22º34’37”S 43º26’05”W, Malaise-Trilha Ponto 1, 5–8.iii.2002, Amarante, S.T.P. & eq. Col (MZUSP); 1 ♀, Nova Iguaçu, Reserva Biológica do Tinguá, 22º34’37”S

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43º26’05”W, Malaise-Bosque Ponto 1, 5–8.iii.2002, Amarante, S.T.P. & eq. Col (MZUSP); 2 ♀, 22º34’30”S 43º26’07”W, Malaise-Trilha Ponto 4, 5–8.iii.2002, Amarante, S.T.P. & eq. Col (MZUSP); 2 ♀, 22º34’30”S 43º26’08.4”W, Malaise-Bosque Ponto 4, 8–11.iii.2002, Amarante, S.T.P. & eq. Col (MZUSP); 1 ♀, 22º34’28”S 43º26’10.7”W, Malaise-Bosque Ponto 5, 5–8.iii.2002, Amarante, S.T.P. & eq. col. (MZUSP). 1 ♀, Nova Iguaçu, Reserva Biológica do Tinguá, 22º34’37”S 43º26’05”W, Malaise-Bosque Ponto 1, 8–11.iii.2002, Amarante, S.T.P. & eq. Col. (MZUSP). 1 ♀, Santa Catarina, Chapecó, Linha Monte Belo, 27º7’74”S 52º32’20.22”W, 580 m, 01–31.x.2012, M. Savaris, S. Lampert cols. (MZUSP). 1 ♀, Nova Teutônia, 27º11’S 52º23’W, 300-500 m, i.1959, Fritz Plaumann (MZUSP); 1 ♂, 3.i.1949 (CNC); 1 ♀, 4.iii.1956 (CNC); 1 ♀, 11 ♀, 19.i.1957 (CNC); 1 ♀, 29.i.1957 (CNC); 1 ♀, 4.ii.1957 (CNC); 1 ♀, 23.iv.1957 (CNC); 1 ♂, 28.xii.1968 (USNM); 1 ♀, 29.x.1958 (USNM); 2 ♀, 17.xii.1959 (CNC); 1 ♀, 18.xi.1960 (CNC); 1 ♀, 17.i.1961 (CNC); 1 ♀, 2.iii.1961 (USNM); 1 ♀, 26.x.1961 (CNC); 1 ♀, 18.xi.1961 (CNC); 1 ♀, 28.xii.1961 (CNC); 1 ♀, 6.ii.1962 (CNC); 1 ♂, xi.1962 (CNC); 1 ♀, xii.1963 (CNC); 1 ♀, xi.1964 (CNC); 1 ♀, ii.65 [1965] (CNC); 1 ♀, xii.1965 (CNC); 1 ♀, x.1966 (CNC). 1 ♀, São Paulo, Bertioga, Parque Estadual Restinga de Bertioga, Rio Perequê Mirim, 23º43’35”S 45º56’10”W, 21.xi.2012–5.i.2013, Malaise, Biffi, Cezar & Fuhrmann col. (MZUSP); 1 ♀, Parque Estadual Restinga de Bertioga, Rio Itapanhaú, 23º45’07”S 46º03’13”W, 248m, 23.xi.2012–5.i.2013, Malaise, Biffi, Cezar & Fuhrmann col. (MZUSP). 1 ♀, Jundiaí, Serra do Japi, Trilha da Cachoeira do Paraíso, riacho após 2ª represa, 23º14’S 46º57’W, 1050m, Luz UV/branca, 12.iii.2008, Lecci, L.S. & Nascimento, E.A. (MZUSP). 1 ♀, Ribeirão Preto, E.E.R.P, Mata de Sta. Teresa, Malaise Branca, Rio, 21°13’30”S 47°51’01”W, 19.xii.2005, Polegatto, C.M. et al. (MZUSP); 2 ♀, Malaise Preta, 21.xi.2005 (MZUSP); 1 ♀, Malaise Branca, interior, 21.xi.2005 (MZUSP); 1 ♀, 20.i.2006 (MZUSP); 3 ♀, 17.i.2007 (MZUSP); 2 ♀, iv.2008 (MZUSP). 8 ♀ (1 ♀, antennae and left wing slide–mounted), Teodoro Sampaio, P.E. Morro do Diabo, 22º36’18.4”S 52º18’10.2”W, 16.xii.2010, Malaise-pto 5, N.W. Perioto & eq. col. (MZUSP). 1 ♀, São Luís do Paraitinga, P.E.S.M. – Núcleo Santa Virgínia, 23º19’27.2”S 45º05’38.5”W, 22.x.2010, Malaise Ponto 7 (teste), N.W. Perioto & eq. cols. (MZUSP). 1 ♀, Ubatuba, Pq. Est. Serra do Mar, Núcleo Picinguaba, Bacia amarela – 600 m, 21.ii–02.iii2007, E.F. Santos, & C.P. Scott-Santos cols. (MZUSP). Redescription. Male. Length: body, 6.5–7.0 mm; wing, 6.5–7.0 mm. Head. Vertex yellow, setation yellow. Upper frons reddish yellow. Flagellomeres 1–3 brownish on outer margin, paler ventrally on inner margin; flagellomeres 4–8 dark brown to black; eighth flagellomere five times length of seventh flagellomere (Fig. 2). Palpus (Fig. 67) reddish yellow, elongate; basal segment cylindrical, as long as wide; apical segment oval, 1.8–2.3 its width, 1.6–4.0 length of basal segment, both with pale hairs. Thorax (Figs. 8, 60). Entirely reddish yellow, postpronotal lobes and notopleural strip slightly contrasting with color of scutum. Scutum and scutellum orange, with brownish pilosity. Legs. Mostly reddish yellow, except for brownish tarsomeres (basitarsus only distally), mid and hind trochanters, hind coxae and hind tibiae dark brown to black. Wing as in Fig. 66. Halter yellow. Abdomen. Reddish yellow, with black transverse bands on tergites; tergites 2–4 with one small macula, mostly yellow; sternites weakly sclerotized. Male genitalia (Figs. 9–14). Distal margin of gonocoxite rounded (Fig. 9); inner plate with two acuminate projections distally (Fig. 10). Gonostylus wider on basal half, with a small rounded inner projection (Figs 12–13). Ejaculatory apodeme truncate basally (Fig. 9). Distal margin of lateral projections of phallus rounded (Figs. 9–10). Cercus more or less oval (Fig. 14). Female. Differs from male as follow. Length: body, 6.0–7.5 mm; wing, 6.0–8.0 mm. Abdomen. Tergites mostly black, except for tergites 7–8; sternites weakly sclerotized, 1–4 with a small black macula. Female genitalia (Figs. 15–20). Tergite 8 wide, more or less rectangular, rounded anteriorly. Sternite 8 quadrate, narrowing towards apex (Fig. 19); gonapophyses slightly sclerotized, strongly bilobed, rounded apically. Tergite 9 more or less rounded laterally (Fig. 16). Genital fork with anterior arm rounded at extreme base (Figs. 15–17); membranous lobes of genital fork wide, covering nearly entire distal half; projection of posterior margin short, inner margin oblique; margin between posterior projection with a more or less oval medial incision; genital opening large, circular, a few microtrichia on bursa. Three spermathecae present (Fig. 56), lateral ones with short ducts, more sclerotized posteriorly, with a distinct rounded valve near middle of duct; spermathecal capsules circular, helicoid, with two coils, strongly sclerotized (Fig. 18). Cercus with first segment moderately compressed, basally wider than distally; first segment of cercus more than 1.5 times length of second segment. Geographic distribution. Brazil (states of Mato Grosso, Minas Gerais, Rio de Janeiro, Santa Catarina and São Paulo) (Fig. 92).


FACHIN & AMORIM

FIGURES 9–14. Arthropeina fulva, female. 9. Male genitalia, dorsal view. 10. Male genitalia, ventral view. 11. Male genitalia, lateral view. 12. Gonostylus, ventral view. 13. Gonostylus, inner view. 14. Epandrium, dorsal view. Black seta, region of insertion of the gonostylus. Abbreviations: aed, aedeagus; cerc, cercus; ej apod, ejaculatory apodeme; epand, epandrium; epiprct, epiproct; goncx, gonocoxite; gonst, gonostylus; hyprct, hypoproct; pm, paramere.



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FIGURES 15–20. Arthropeina fulva. 15. Genital fork, ventral view. 16. Genital fork, lateral view. 17. Genital fork, dorsal view. 18. Spermatheca. 19. Sternite 8, dorsal view. 20. Female genitalia, dorsal view. Abbreviations: cerc, cercus; go, genital opening; ml, membranous lobe; pp, posterolateral process; ppm, projection of posterior margin; st, sternite; tg, tergite. Scale bar, 0.1 mm.


FACHIN & AMORIM

FIGURES 21–26. Arthropeina colombiana, sp. nov., holotype. 21. Genital fork, ventral view. 22. Genital fork, lateral view. 23. Genital fork, dorsal view. 24. Spermatheca. 25. Sternite 8, dorsal view. 26. Female genitalia, dorsal view. Abbreviations: cerc, cercus; go, genital opening; ml, membranous lobe; pp, posterolateral process; ppm, projection of posterior margin; st, sternite; tg, tergite. Scale bar, 0.1 mm.



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FIGURES 27–31. Arthropeina diadelothorax, sp. nov., holotype. 27. Genital fork, ventral view. 28. Genital fork, lateral view. 29. Genital fork, dorsal view. 30. Sternite 8, dorsal view. 31. Female genitalia, dorsal view. Abbreviations: cerc, cercus; go, genital opening; ml, membranous lobe; pp, posterolateral process; ppm, projection of posterior margin; st, sternite; tg, tergite.


FACHIN & AMORIM

FIGURES 32–37. Arthropeina lindneri, sp. nov., paratype. 32. Genital fork, ventral view. 33. Genital fork, lateral view. 34. Genital fork, dorsal view. 35. Spermatheca. 36. Sternite 8, dorsal view. 37. Female genitalia, dorsal view. Abbreviations: cerc, cercus; go, genital opening; ml, membranous lobe; pp, posterolateral process; ppm, projection of posterior margin; st, sternite; tg, tergite. Scale bar, 0.1 mm.



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FIGURES 38–42. Arthropeina melanochroma, sp. nov., holotype. 38. Male genitalia, dorsal view. 39. Male genitalia, ventral view. 40. Gonostylus, ventral view. 41. Gonostylus, inner view. 42. Epandrium, dorsal view. Black seta, region of insertion of the gonostylus. Abbreviations: aed, aedeagus; cerc, cercus; ej apod, ejaculatory apodeme; epand, epandrium; epiprct, epiproct; goncx, gonocoxite; gonst, gonostylus; hyprct, hypoproct; pm, paramere.


FACHIN & AMORIM

FIGURES 43–49. Arthropeina melanochroma, sp. nov., paratype. 43. Genital fork, ventral view. 44. Genital fork, lateral view. 45. Genital fork, dorsal view. 46. Spermatheca. 47. Spermatheca, lateral view. 48. Sternite 8, dorsal view. 49. Female genitalia, dorsal view. Abbreviations: cerc, cercus; go, genital opening; ml, membranous lobe; pp, posterolateral process; ppm, projection of posterior margin; st, sternite; tg, tergite. Scale bar, 0.1 mm.



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Comments. This species shows variation in the hind leg color. The specimens of the type series of Arthropeina fulva, according to the original description, and some of the identified specimens have dark brown to black hind coxa, while specimens from Teodoro Sampaio (State of São Paulo), Presidente Olegário (State of Minas Gerais) and Chapada dos Guimarães (State of Mato Grosso) have the hind coxae yellow to reddish yellow. This difference does not seem significant to indicate that this set of specimens are not conspecific. Structures of female genitalia and spermatheca do not show substantial differences. Most specimens from this species studied here, however, are females. Maybe males from these areas may show more significant differences. At present, we decided to treat these specimens as conspecific and the differences in color as intraspecific variation.

Arthropeina colombiana, sp. nov. (Figs. 3, 21–26, 61, 68–69, 79, 85) Diagnosis (female). Length of eighth flagellomere two times length of seventh (Fig. 3). Prescutum anteriorly and laterally dark brown, postscutum basally dark brown (Fig. 61). Projection of posterior margin of genital fork rounded basally (Figs. 21, 23). Tergite 10 projected between the cerci in females (Fig. 26). Material examined. HOLOTYPE, ♀ (right antenna and right wing slide–mounted), COLOMBIA, Valle del Cauca, PNN Farallone de Cali, Cgto. La Meseta, 03º34’N 76º40’W 1960 m, Malaise 10–25.ix.2003, S. Sarria & M. Losso Leg. M. 4547 (IAvH). Description. Female. Length: body, 8.1 mm; wing, 9.0 mm. Head. Vertex pale, setation yellow. Upper frons slightly reddish yellow. Flagellomeres l–3 brownish on outer margin, pale on inner margin; flagellomeres 4–8 dark brown to black. Eighth flagellomere twice the length of seventh (Fig. 3). Palpus (Fig. 69) whitish yellow, elongate; basal segment three times its width; apical segment oval, 2.75 times as long as wide, 1.5 times length of basal segment, both with pale hairs. Thorax (Fig. 61). Mostly yellow. Postpronotal lobes whitish yellow, dark brown anteriorly, notopleural strip dark brown, both contrasting with scutum color. Prescutum anteriorly and laterally dark brown, postscutum basally and its connection with the scutellum dark brown; scutellum mostly whitish yellow, except dark brown on outer margin; brownish pilosity. Legs. Mostly reddish yellow, except for tarsomeres slightly brownish, all trochanters brown to dark brown, hind coxae dorsally, and mid and hind tibiae on basal two-thirds dark brown. Wing as in Fig. 68. Halter pale, knob whitish yellow. Abdomen. Reddish yellow, with black transverse bands on tergites; sternites weakly sclerotized, 1–4 with small brown maculae; first tergite entirely black. Female genitalia (Figs. 21–26). Tergite 8 wide, more or less rectangular, rounded anteriorly. Sternite 8 quadrate (Fig. 25), narrowing towards distal half; gonapophyses sclerotized, lobes divergent, rounded apically. Tergite 9 with distal half wider than basal half (Fig. 22). Genital fork with anterior arm rounded (Figs. 21–23), as wide as posterior arm; membranous lobes of fork wide; projection of posterior margin rounded basally; margin between posterior projection with a moderately circular incision medially; genital opening small, circular, a few microtrichia on bursa. Three spermathecae present (Fig. 24), lateral ones with long ducts; spermathecal capsules more or less ovoid, strongly sclerotized, not clearly helicoid. Tergite 10 projected between the cerci (Fig. 26). Cercus of first segment moderately compressed, basally wider than distally; segment 1 of cercus two times length of segment 2. Male. Unknown. Etymology. The name is feminine and the specific epithet is a reference to the type–locality. Geographic distribution. Known only from the type-locality in Valle del Cauca, Colombia (Fig. 92).

Arthropeina diadelothorax, sp. nov. (Figs. 4, 27–31, 62, 70–71, 80, 86) Diagnosis (female). Body color yellowish (Figs. 80, 86). Scutum mostly reddish yellow, with two quadrangular black spots on prescutum, two triangular black spots on postscutum, and a medial, wider black band extending from anterior margin of scutum to scutellum basally (Fig. 62); anepisternum anteriorly with a black spot; laterotergite and mediotergite dark brown to black. Projection of posterior margin of genital fork long, abruptly narrowing towards apex (Figs. 27–29).


FACHIN & AMORIM

FIGURES 50–55. Arthropeina pseudofulva, sp. nov., paratype. 50. Genital fork, ventral view. 51. Genital fork, lateral view. 52. Genital fork, dorsal view. 53. Spermatheca. 54. Sternite 8, dorsal view. 55. Female genitalia, dorsal view. Abbreviations: cerc, cercus; go, genital opening; ml, membranous lobe; pp, posterolateral process; ppm, projection of posterior margin; st, sternite; tg, tergite. Scale bar, 0.1 mm.



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FIGURES 56–59. Female reproductive tracts, including spermathecae of Arthropeina spp. 56. Arthropeina fulva. 57. A. lindneri, sp. nov., paratype. 58. A. melanochroma, sp. nov., paratype. 59. A. pseudofulva, sp. nov., holotype. Scale bar, 0.1 mm.


FACHIN & AMORIM

FIGURES 60–65. Arthropeina spp., thorax in dorsal view. 60. Arthropeina fulva, male. 61. A. colombiana, sp. nov., holotype. 62. A. diadelothorax, sp. nov., holotype. 63. A. lindneri, sp. nov., paratype. 64. A. melanochroma, sp. nov., holotype. 65. A. pseudofulva, sp. nov., paratype. Scale bar, 1 mm.

Material examined. HOLOTYPE, ♀, BRAZIL, Rondônia, Cacaulândia, Trilha da Cachoeira Jamari, 10º13’28,8”S 63º13’49”W, Malaise 27, 10.ii–20.v.2012, Lamas, Nihei & eq. col. (MZUSP). PARATYPES: 1 ♀, (antennae and left wing slide–mounted), 04.xi.2011–10.ii.2012, same locality as holotype (MZUSP). Description. Female. Length: body, 5.0–6.0 mm; wing, 5.0–6.0 mm. Head. Vertex yellowish, setation yellow. Upper frons yellowish. Flagellomeres l–3 dark brown on outer margin, pale ventrally on inner margin; flagellomeres 4–8 dark brown to black; eighth flagellomere more than three times the length of seventh (Fig. 4). Palpus (Fig. 71) pale, elongate; basal segment cylindrical, two times its width; apical segment oval, 1.6 times its width, twice length of basal segment; both with pale hairs. Thorax (Fig. 62). Postpronotal lobes pale, with a narrow blackish notopleural strip. Scutum reddish yellow, with two more or less quadrate black spots on prescutum, two triangular black spots on postscutum, and a medial wider black band (wider at anterior and posterior ends) extending from anterior margin of scutum to base of scutellum; scutellum black basally, with short yellow pilosity. Pleura pale, except anepisternum anteriorly with a large medial black spot; laterotergite and mediotergite dark brown to black. Legs. Mostly yellow, except for tarsomeres vaguely brownish, hind coxae and trochanters dark brown, mid femora at extreme base and ventrally at apex brownish, hind femora at extreme base as well as two-thirds of hind tibiae dark brown. Wing as in Fig. 70. Halter reddish yellow. Abdomen. Yellowish with black transverse bands on tergites except on tergite 1; black color slightly decreasing in width from tergites 2 to 4; tergites 5–7 entirely black; sternites weakly sclerotized. Female genitalia (Figs. 27–31). Tergite 8 wide, more or less quadrate, rounded anteriorly, slightly flattened medially at basal margin. Sternite 8 roughly quadrate (Fig.



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30), weakly sclerotized, narrowing towards apex; gonapophyses unsclerotized, with short and wide lobes. Tergite 9 wider distally (Fig. 28). Genital fork with anterior arm rounded (Figs. 27–29), clearly more slender than posterior arm; posterolateral processes rounded; membranous lobes of genital fork more rounded distally, with basal half more slender than distal half; projection of posterior margin long, narrow, acuminate; margin between posterior projection with a moderately profound medial incision; genital opening small, oval, placed basally on posterior margin, bursa densely covered with microtrichia. Cercus with segment 1 more than twice length of segment 2. Male. Unknown. Etymology. The species name is feminine, and the specific epithet comes from the Greek roots diadelos, meaning distinctive, distinguishable, and thorax, as a reference to the conspicuous color pattern of the scutum of the species. Geographic distribution. Northern Brazil (State of Rondônia) (Fig. 92). Comments. The laterotergite of this species may be entirely or only partly black. In addition, on the abdomen, the black color of tergites 5–7 sometimes does not cover the entire surface.

Arthropeina lindneri, sp. nov. (Figs. 5, 32–37, 57, 63, 72–73, 81, 87) Diagnosis (female). Body color mostly whitish yellow to yellow (Figs. 81, 87), except scutum reddish yellow (Fig. 63). Scutum with two black spots anteriorly, a pair of quadrate black spots between posterior margin of postpronotal lobes and transverse suture, and a medial black band gradually widening from anterior margin of scutum to base of scutellum (Fig. 63). Projection of posterior margin of genital fork long, rounded (Figs. 32–34). Spermathecal capsule simple, with no coils (Fig. 35). Material examined. HOLOTYPE, ♀, BRAZIL, Rondônia, Porto Velho, Área Mutum, M5P4, 09º35’29.5”S 65º02’57.6”W, 29.iii–05.iv.2012, Silva, R.R. & Albuquerque, E.Z. cols. (MZUSP). PARATYPES: 1 ♀, BRAZIL, Acre, Mancio Lima, Pq. Nac. Serra do Divisor, Malaise, 5.ix.2007, L. Menezes (MZUSP); 1 ♀, Igarapé do Amor, 5.xi.2007 (MZUSP). 1 ♀, Roraima, Caracaraí (Vila Caicubi, Trilha do Bacaba), 00°58’36.5”S 62°06’08.7”W, 28.viii–10.ix.2011, Malaise 1, Biffi, G. & Prado, L.R. cols. (MZUSP). 1 ♀, Rondônia, Cacaulândia, Trilha Cachoeira Jamari, 131 m, 10º13’26.8”S 63º13’49”W, 04.xi2011–10.ii.2012, Malaise 27, Lamas, Nihei & eq. col. (MZUSP). 1 ♀, Monte Negro, Fazenda Amorim, 10°40’6”S 63°29’0”W, 248m., Malaise trap 6., 03–15.xii.2011, Amorim, Ament & Riccardi col. 1 ♀, Porto Velho, Área Mutum, M7P3, 09°35’41.6”S 65°03’54.2”W, 29.iii–05.iv.2012, Silva, R.R. & Albuquerque, E.Z. cols. (MZUSP). 1 ♀ (antennae and left wing slide–mounted), Porto Velho, Área Caiçara, C3P3, 09º26’46.8”S 64º49’31.1”W, 29.iii–05.iv.2012, Silva, R.R. & Albuquerque, E.Z. cols. (MZUSP). 1 ♀, GUYANA, Mazaruni-Potaro District, Takutu Montains, 6°15’N 59°5’W, 14.xii.1983, Malaise trap near stream in montane rainforest, P.J. Spangler & W.E. Steiner (USNM). Description. Female. Length: body, 5.0–7.5 mm; wing, 5.5–8.0 mm. Head. Vertex pale, setation yellow. Upper frons yellow to reddish yellow. Flagellomeres l–3 dark brown to black on outer margin, pale ventrally on inner margin; flagellomeres 4–8 dark brown to black; length of flagellomere 8 more than three times length of seventh (Fig. 5). Palpus (Fig. 73) pale, elongate; basal segment cylindrical, as long as wide; apical segment oval, length 2.25–2.0 times its width, 2.0–3.75 times length of basal segment; pale hairs. Thorax (Fig. 63). Postpronotal lobes pale, slightly contrasting with scutum color, and with a very narrow notopleural strip light brownish. Scutum reddish yellow, with two black spots anteriorly, two more or less quadrangular black spots between the posterior margin of postpronotal lobes and the transverse suture, and a medial black band gradually widening from anterior margin of scutum to base of scutellum; scutellum pale distally, with short brownish pilosity. Pleura pale, except anepisternum, with a medial black spot; laterotergite partly black distally. Legs. Mostly yellow, except for tarsomeres slightly brownish; coxae, trochanters and hind femora dark brown apically and hind tibiae entirely dark brown, except at extreme base. Wing as in Fig. 72. Halter reddish yellow. Abdomen. Pale with black transverse bands on tergites, except on tergite 1; black color decreasing in width from tergite 2 to 4, tergites 5–7 entirely black, also part of sternites; sternites weakly sclerotized. Female genitalia (Figs. 32–37). Tergite 8 more or less quadrate, slightly rounded anteriorly. Sternite 8 almost quadrate (Fig. 36), weakly sclerotized distally, narrowing towards apex; gonapophyses unsclerotized, with short lobes separated by a shallow medial incision. Tergite 9 more or less straight laterally (Fig. 33). Genital fork with anterior arm rounded (Figs. 32–34), distinctly narrower than posterior arm; posterolateral process wider, distally narrower than basally; membranous lobes of genital fork more rounded


FACHIN & AMORIM

distally; projection of posterior margin long, rounded, with a deep medial incision; genital opening small, circular. Three spermathecae present (Fig. 57), lateral ones with long ducts; valves only distinguishable as a widening between posterior and anterior part of ducts; spermathecal capsules circular, strongly sclerotized (Fig. 35). Cercus with segment 1 elongate, length more than three times length of segment 2. Male. Unknown. Etymology. The name given is feminine and the specific epithet is named after Erwin Lindner (1888–1988), a Stuttgart-based German entomologist with a long career and a major contribution to world dipterology. He described, among his contributions to many Diptera families, the genus Arthropeina and its type-species. Geographic distribution. Guyana and northern of Brazil (States of Acre, Rondônia and Roraima) (Fig. 92).

Arthropeina melanochroma, sp. nov. (Figs. 6, 38–49, 58, 64, 74–75, 82, 88) Diagnosis. Body color including the scutum, mostly black (Figs. 82, 88). Postpronotal lobes and notopleural strip whitish yellow, strongly contrasting with scutum color (Fig. 64). Spermathecal capsule biconcave (Figs. 46–47). Material examined. HOLOTYPE, ♂, ECUADOR, Napo, Res. Ethnica, Waorani, 1 km S. Okone Gare, Camp. Trans. Ent. 00º39’25.7”S 076°27’10.8”W, 261m, Insectidal fogging, terre firme forest, Lot. 964, Transect 4, Station 8, 10.x.1994, T.L. Erwin et al. (USNM). PARATYPES: 1 ♀ (antennae and right wing slide–mounted), BRAZIL, Rondônia, Monte Negro, Reserva Legal/Loteamento, 10º16’35”S 63º20’40”W, 187m, Malaise, 03–15.xii.2011, Amorim, Ament & Riccardi col. (MZUSP). 1 ♀, ECUADOR, Napo, Tiputini, Biodiversity Station, nr Yasuni National Park, 00º37’55”S 076°08’39”W, 220–250m, Insectidal fogging, terre firme forest, Erwin Transect T/2, Lot. 2017, Transect 2, Station 8, 09.ii.1999, T.L. Erwin et al. (USNM). 1 ♀, PERU, Madre de Dios, Avispas, 1-15.x.1962, L. Pena, 400m (CNC). Description. Male. Length: body, 6.5 mm; wing, 6.0 mm. Head. Vertex reddish yellow, setation yellow. Upper frons yellow to reddish yellow. Antenna with scape brown, whitish yellow distally; pedicel whitish yellow, setation light yellow; flagellomeres l–3 brownish yellow on outer margin, pale ventrally on inner margin; flagellomeres 4–8 dark brown; eighth flagellomere more than three times length of seventh (Fig. 6). Palpus (Fig. 75) whitish yellow, elongate; basal segment cylindrical, sclerotized distally, length 1.8 times its width; apical segment oval, sclerotized basally, length three times its width, twice length of basal segment; pale hairs. Proboscis whitish yellow, at least dark brown on distal half; pale hairs. Thorax (Fig. 64). Black, with postpronotal lobes and notopleural strip whitish yellow, strongly contrasting with the color of scutum. Scutum and scutellum entirely black, except subalar callus and postpronotal lobes; pilosity yellowish. Pleura black except for posterior half of anepisternum whitish. Legs. Mostly whitish yellow, except for coxae, trochanters, fore and mid femora at extreme base and apex, hind femora on basal two thirds and apex, hind femora almost entirely (except at extreme basal) and all tarsomeres, dark brown to black. Wing as in Fig. 74. Halter pale, knob yellow whitish. Abdomen. Whitish yellow, with black transverse bands on segments; black color covering nearly all segments. Male genitalia. (Figs. 38–42) Distal margin of gonocoxite strongly pointed (Fig. 38); inner plate with three projections distally (Fig. 39), medial one narrower and longer than lateral ones. Gonostylus with three projections (Figs. 40–41), distal one clearly longer than others. Ejaculatory apodeme rounded basally (Fig. 38). Distal margin of lateral projections of phallus gradually narrowing towards apex (Figs. 38–39). Cercus wider on distal half (Fig. 42). Female. Differs from male as follow. Length: body, 7.0 mm; wing 7.0–7.5 mm. Legs. Fore femora entirely whitish yellow or with extreme base and apex, dark brown to black. Female genitalia (Figs. 43–49). Tergite 8 wide, more or less rectangular, rounded anteriorly. Sternite 8 elongate (Fig. 48), more slender basally than distally; gonapophyses sclerotized, strongly bilobed, gradually tapering towards apex. Tergite 9 wider basally than distally (Fig. 44). Genital fork with anterior arm rounded basally (Figs. 43–45), as wide as posterior arm; membranous lobes of genital fork rounded, wider distally; projection of posterior margin slightly pointed distally; margin between posterior projection with a moderately circular incision medially; genital opening more or less oval, gradually narrowing towards apex, with numerous microtrichia on bursa. Three spermathecae present (Fig. 58), lateral ones with long ducts, strongly sclerotized posteriorly, with distinct circular valve medially; spermathecal capsules biconcave, more sclerotized basally (Figs. 46–47). Tergite 10 limited to two lateral triangular plates in dorsal view. Cercus with first segment compressed, wider basally than distally; segment 1 of cercus about two times length of segment 2. FACHIN & AMORIM—REVISION OF ARTHROPEINA


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FIGURES 66–77. Arthropeina spp, wing and palpus. 66–67. Arthropeina fulva, female. 68–69. A. colombiana, sp. nov., holotype. 70–71. A. diadelothorax, sp. nov., paratype. 72–73. A. lindneri, sp. nov., paratype. 74–75. A. melanochroma, sp. nov., female paratype. 76–77. A. pseudofulva, sp. nov., paratype. Scale bar, 1 mm.


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FIGURES 78–83. Arthropeina spp., dorsal view. 78. Arthropeina fulva, female. 79. A. colombiana, sp. nov., holotype. 80. A. diadelothorax, sp. nov., paratype. 81. A. lindneri, sp. nov., paratype. 82. A. melanochroma, sp. nov., female paratype. 83. A. pseudofulva, sp. nov., paratype. Scale bar, 1 mm. FACHIN & AMORIM—REVISION OF ARTHROPEINA


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FIGURES 84–89. Arthropeina spp., lateral view. 84. Arthropeina fulva, female. 85. A. colombiana, sp. nov., holotype. 86. A. diadelothorax, sp. nov., holotype. 87. A. lindneri, sp. nov., paratype. 88. A. melanochroma, sp. nov., female paratype. 89. A. pseudofulva, sp. nov., holotype. Scale bar, 1 mm.


FACHIN & AMORIM

FIGURES 90–91. Solva sp., female. 90. Hind femur, ventral view. 91. Hind femur in detail, ventral view. Black seta, socket of setae modified in small black tubercle. Scale bar, 0.1 mm.

Etymology. The species name is feminine, and the specific epithet comes from the Greek roots melanos, meaning black, dark, and chroma, color, as a reference to dark body surface of the species. Geographic distribution. Ecuador, Peru and northern Brazil (State of Rondônia) (Fig. 92). Comments. This species presents variation on intensity leg as well as areas that in some specimens are dark brown to black. Occasionally, the hind femur may be almost entire black, except for distal whitish annulus, and the mid tibia may have the distal two-thirds dark brown to black, as well as the fore tibia may be brownish apically. In the other hand, the dark color of the hind femur may be restricted to the basal and distal extremes.

Arthropeina pseudofulva, sp. nov. (Figs. 7, 50–55, 59, 65, 76–77, 83, 89) Diagnosis (female). Body color mostly whitish yellow to reddish yellow (Figs. 83, 89). Thorax mostly whitish yellow except for scutum reddish yellow (Fig. 65). Legs mostly whitish yellow, including hind coxa (Fig. 89). Projection of posterior margin of genital fork very short, nearly indistinguishable (Figs. 50–52). Spermathecal capsule helicoid, with three coils (Fig. 53). Material examined. HOLOTYPE, ♀, BRAZIL, Rondônia, Campo Novo, Fazenda Amorim, 10º40.6’S 63º29’0”W, Malaise trap 6m, 06–10.xii.2011, Amorim, Ament e Riccardi col. (MZUSP). PARATYPES: 1 ♀, BOLIVIA, Santa Cruz Dist., 4 km N Bermejo, Refugio Los Volcanes, 18°06’S 63°36’W, AR Cline, 1000m, 25–30.x.2007, Malaise trap. (CSCA). 1 ♀ (left antenna and left wing slide–mounted), BRAZIL, Monte Negro, Fazenda Amorim, 10°40.6’S 63°29’0”W, 248m, Malaise trap, 03–15.xii.2011, Amorim, Ament e Riccardi col. (MZUSP). Description. Female. Length: body, 6.5–9.0 mm; wing, 6.0–7.5 mm. Head. Vertex pale, setation whitish yellow. Upper frons reddish yellow. Flagellomeres 1–3 brownish on outer margin, paler ventrally on inner margin; flagellomeres 4–8 dark brown to black; length of eighth flagellomere five times length of seventh (Fig. 7). Palpus (Fig. 77) whitish yellow, elongate; basal segment cylindrical, length two times its width; apical segment oval, length two times its width, 1.7 times length of basal segment, both with pale hairs. Thorax (Fig. 65). Entirely whitish yellow to yellow, except scutum reddish yellow; postpronotal lobes whitish yellow, notopleural strip brownish, moderately contrasting with scutum color. Scutellum whitish yellow. Legs. Mostly reddish yellow, except for tarsomeres brownish, hind trochanters, hind femora apically and hind tibiae brown to black. Wing as in Fig. 76. Halter pale, knob yellow whitish. Abdomen. Reddish yellow, with black transverse bands on tergites; sternites unsclerotized; tergites mostly black, except tergite 1 with a thin transverse black distal line; tergites 7–8 weakly sclerotized. Female genitalia (Figs. 50–55). Tergite 8 wide, more or less rectangular, slightly rounded



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anteriorly. Sternite 8 more or less oval (Fig. 54), narrowing towards apex; gonapophyses slightly sclerotized, strongly bilobed, rounded apically. Tergite 9 wider distally (Fig. 51). Genital fork with anterior arm rounded, clearly narrower than posterior arm (Figs. 50, 52); membranous lobes of genital fork wide, rounded, covering the posterolateral process; projection of posterior margin very short; margin between posterior projections with a moderate medial incision; genital opening more or less oval, widest medially; lateral membrane of bursa sclerotized with a few microtrichia. Three spermathecae present (Fig. 59), lateral ones with short ducts, more sclerotized posteriorly, with a distinct rounded valve near middle; spermathecal capsules more or less ovoid, helicoid, with three coils, strongly sclerotized (Fig. 53). Cercus of first segment moderately compressed, wider basally than distally; length of segment 1 of cercus more than twice that of segment 2. Male. Unknown. Etymology. The species name is feminine and the specific epithet adds to fulva the Greek root pseudo, for false, considering its similarity with A. fulva, but actually being a different species. Geographic distribution. Northern Brazil (State of Rondônia) and Bolivia (Fig. 92).

FIGURE 92. Geographic distribution of Arthropeina species. Squares represent holotypes, and circles paratypes and additional material.


FACHIN & AMORIM

Comments. This species closely resembles A. fulva, mainly due to the general body color. The hind coxa color may be misleading because some specimens of A. fulva can have a yellow hind coxa, as in A. pseudofulva (see comments on A. fulva). The antenna is also similar in both species, with the same relative lengths of flagellomeres 7–8. Nevertheless, A. pseudofulva can be easily discerned from A. fulva based on the shape of the genital fork—with the projection of the posterior margin very short, nearly indistinguishable—the shape of the genital opening, and the helicoid spermathecal capsules. The distributions of the two species (Fig. 92) also differ. Arthropeina fulva is restricted to southern Brazil, except for a single record for the State of Mato Grosso, in western Brazil. Arthropeina pseudofulva is restricted to western Amazonia and Bolivia.

Key to the adult species of Arthropeina 1. -. 2. -. 3.

-. 4. -. 5.

-.

Body predominantly pale yellow to reddish yellow (e.g., Fig. 78); postpronotal lobes and notopleural strip slightly contrasting with scutum color (e.g., Fig. 60. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 Body predominantly black (Figs. 82, 88); postpronotal lobes and notopleural strip white, strongly contrasting with scutum color (Fig. 64) (Brazil, Ecuador and Peru) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. melanochroma, sp. nov. Scutum and scutellum with no black markings or bands (e.g., Fig. 60) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 Scutum and scutellum with black markings or bands (e.g., Fig. 62) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4 Hind coxa predominantly dark brown to black (Fig. 84); projection of posterior margin of genital fork short (e.g., Fig. 15); spermathecal capsule helicoid, with two coils (Fig. 18) (Brazil, States of Mato Grosso, Minas Gerais, Rio de Janeiro, Santa Catarina and São Paulo) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. fulva Lindner Hind coxa yellow (Fig. 89); projection of posterior margin of genital fork nearly indistinguishable (e.g., Fig. 52); spermatheca capsule helicoid with three coils (Fig. 53) (Brazil, State of Rondônia; Bolivia) . . . . . . . . . . . . . . . . . . .A. pseudofulva, sp. nov. Length of flagellomere 8 more than three times length of flagellomere 7 (e.g., Figs. 4); scutum with distinct black markings (e.g., Fig. 62); anepisternum anteriorly with black markings (e.g., Figs. 86) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5 Length of flagellomere 8 twice length of flagellomere 7 (Fig. 3); scutum with black markings only laterally (Fig. 61); anepisternum anteriorly without markings (Fig. 85) (Colombia) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. colombiana, sp. nov. Medial band of scutum wide, nearly parallel-sided anteriorly, extending from prescutum to scutellum (Fig. 62); laterotergite with or without black maculae; projection of posterior margin of genital fork very long and acuminate (e.g., Fig. 27) (Brazil) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. diadelothorax, sp. nov. Medial band of scutum very slender anteriorly, nearly absent at anterior margin, widening abruptly in posterior half (Fig. 63); laterotergite without black maculae; projection of posterior margin of genital fork long, rounded (e.g., Fig. 34) (Brazil, Guyana) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A. lindneri, sp. nov.

Discussion The species of Arthropeina show a considerable diverse non-genitalic morphology that allows species level recognition. This mostly concerns body color and differences in antennal morphology. Occasionally, however, some color features may be variable and misleading in species identification, as in A. fulva and A. pseudofulva. On the other hand, male and female genitalia morphology, including the spermathecae, allows clear species recognition. The body color, mainly of the scutum, is helpful for species identification. A. fulva and A. pseudofulva have a rather similar general color, without black maculae, but the remaining species are easily recognized by the conspicuous scutum color pattern (e.g., Fig. 62) or the body color as a whole, e.g., in A. melanochroma (e.g., Fig. 82). There is also variation between species of Arthropeina in the length of flagellomeres 7 and 8. A. fulva and A. pseudofulva have a long flagellomere 8, about five times the length of flagellomere 7 (e.g., Fig. 2). A. diadelothorax, A. lindneri and A. melonochroma have flagellomere 8 only moderately long, about three times the length of the seventh (e.g., Fig. 4). In A. colombiana, unlike the remainder of the species of the genus, flagellomere 8 is relatively short, twice the length of flagellomere 7 (Fig. 3), a condition similar to that seen in species of Solva (Fig. 1). Maxillary palpus features also can be used to discriminate between species. There are different ratios between the apical and basal segments, as well as different width of apical segments (e.g., Figs. 67, 75). Among the genera of Xylomyidae, including Arthropeina, the female genitalia morphology also allows



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separation between species. These differences mostly concern the genital fork and spermatheca morphology, which may also be characters that are useful for a phylogenetic study of the genus. The species of Arthropeina present great variation in different aspects of the genital fork and the spermatheca (e.g., Figs. 15, 21). This is the case, e.g., of the shape of the posterolateral process, the shape and length of the posterior projection of the genital fork, the shape and position of the genital opening in the bursa (e.g., Figs. 15, 21), as well as the shape of the anterior part of the genital fork. Furthermore, structures of the female genital tract also show variation, mostly concerning the length of the spermathecal ducts and the shape of the spermathecal capsule (Figs. 56–59). This kind of information also has great importance to discriminate genera and species in other brachyceran families, as seen, for example, in Stratiomyidae (e.g., Woodley, 1981), Asilidae (e.g., Artigas, 1971), and also in Muscidae (e.g., Couri, 1998).

Arthropeina apomorphies The conical shape of the antenna, with flagellomeres 7 and 8 together, forming a stylus (Figs. 2–7) that is as long as or longer than the remainder of the flagellum has been proposed as synapomorphic for the genus (Artigas & Papavero, 1991; Woodley, 2011). This differs from the combined length of flagellomeres 7 and 8, which are shorter than the remaining flagellomeres in Solva and do not form a stylus (Fig. 1). In Arthropeina, the flagellomeres differ from each other in size and shape (Figs. 2–7), while in Solva (Fig. 1; Daniels, 1976, fig. 3) most flagellomeres are similar in size and shape (except for flagellomere 8 which is slightly narrowed towards apex). This is another feature that may be synapomorphic for Arthropeina. A more detailed comparison between additional species of Solva and other xylomyids would make this statement more robust. Papavero & Artigas (1991) proposed that the spermathecal capsule formed by three “layers” or coils—based only on their examination of Arthropeina fulva—is apormophic in species of the genus. This condition is actually shown here to be autapomorphic for A. pseudofulva (Fig. 53), while A. fulva has the spermatheca formed by two coils (Fig. 18). The coiled spermathecal capsule, as described by Papavero & Artigas (1971) (independently of the number of coils), seems to be a synapomorphy for a small clade within Arthropeina including A. fulva, A. colombiana and A. pseudofulva. The “remnant of middle spermathecal duct” (Papavero & Artigas, 1991) certainly corresponds to the spermathecal duct (e.g., Fig. 56) cut during preparation, as already stated by Woodley (2011: 422). The statement that the valves of the spermathecal ducts (Papavero & Artigas, 1991) are an apomorphic condition in the genus, hence, as commented by Woodley (2011), is not confirmed. Other dipteran families also have this condition (see e.g., Artigas, 1971). A wider comparison with other groups of lower brachycerans would be necessary to properly establish the level of origin of this feature—a single origin with losses or independent origins in different families. Additionally, the genital fork in Arthropeina presents a unique condition in Xylomyidae at the level of current knowledge, the membranous lobes (e.g., Fig. 21), not found in other xylomyid genera. A careful morphological study of Solva species would certainly help to better understand of the evolution of features varying within Arthropeina and furnish important evidence towards the construction of a detailed phylogenetic study of the family.

Acknowledgments The authors thank Carlos J. E. Lamas, Diptera Curator of the Museu de Zoologia da USP (MZUSP) and Martin Hauser (CSCA), for the loan of part of the material used in this study. This work includes material of the SisBiota Project from western Brazil (FAPESP Grant 2010/52.314-0; CNPq Grant 563256/2010-9). Additional number of specimens was provided by a Hymenoptera diversity survey along the Atlantic Forest, kindly made available by Dr. Carlos Roberto Ferreira Brandão and Eliana M. Cancello (FAPESP Grant 1998/05.083-0). We would to like to thank Norman E. Woodley (USNM) for providing a large number of specimens, including some males, and he was very helpful with interesting discussions on the genus. Maria Isabel P.A. Balbi (Universidade de São Paulo) has always been of special help in sorting and preparing specimens for examination, as well as providing other laboratory support. This study benefited from FAPESP Grants 2011/14472-6 and 2013/02824-0.


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