Aspistomella, Hendel, 1909

Kovac, Damir, Kameneva, Elena P., Korneyev, Severyn V., Araújo, Alexandre Santos, Savaris, Marcoandre, Smit, John T., Schneider, Alexander, Schreiber, Robert & Korneyev, Valery A., 2024, Revision of the Aspistomella group of genera (Diptera: Ulidiidae: Pterocallinae: Lipsanini), Zootaxa 5530 (1), pp. 1-117 : 9-21

publication ID

https://doi.org/ 10.11646/zootaxa.5530.1.1

publication LSID

lsid:zoobank.org:pub:41376D87-B401-4301-9DDC-54606653881F

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https://treatment.plazi.org/id/03F02928-1E68-FFE0-FF21-52E2FD57FB29

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scientific name

Aspistomella
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Aspistomella View in CoL group of genera

Within the tribe, the genera Aspistomella , Polyteloptera , and Ulivellia form a group here called the Aspistomella group, which differs from other genera of Lipsanini by the combination of a head with a moderately or very large epistome (lower part of the face), strongly anteroventrally produced and conspicuously (usually 2×) higher than the low clypeus (synapomorphy) almost hidden in the peristomal cavity (synapomorphy).

The Aspistomella group of genera belongs to a broader monophyletic lineage (here called the Amethysa lineage) together with the taxa sharing frontal setulae modified to form long and strong frontal and interfrontal setae (synapomorphy); row of fine and short parafrontal setulae 3–5× shorter than frontal setae (synapomorphy); very long ocellar seta (usually 2× or more as long as the postpedicel); very long inner vertical seta (slightly longer than the ocellar seta); short outer vertical seta (at most half as long as inner vertical seta or vestigial) (apomorphies), often setulose vein R 1 (plesiomorphy; secondarily bare in some species); unmodified, relatively short aculeus with tergosternite 8 moderately wide and oval in section (plesiomorphy) and cercal unit oval, not flattened and bearing long setulae (plesiomorphy) (Kameneva & Korneyev, in press).

Other taxa of the Amethysa lineage are distinguished by the low, less produced epistome combined with a high clypeus almost as high as the epistome (except those either without epistome of a receding face ( Eumetopiella , Stenomyia ) or, if the epistome is antero-ventrally produced ( Steneretma , Zacompsia ), then with a high clypeus (Kameneva & Korneyev, in press).

The genera of Lipsanini , except those of the Amethysa lineage, are easily distinguished by the uniformly short, undifferentiated (plesiomorphy), subequal inner and outer vertical setae (plesiomorphy).Among them, many species assigned to Acrosticta , Acrostictomyia , Axiologina , Euxesta , and Pareuxesta share a very narrow and long aculeus, 1.5× longer than the oviscape (synapomorphy), and apparently form another monophyletic lineage (Kameneva & Korneyev, in press).

The species assigned here to the genera of the Aspistomella group often have dorsoapically setulose vein R 1 (possible plesiomorphy, occurring occasionally in different lineages of Lipsanini and Pterocallini, but very common in the Otitini and Myennidini , black halteres (possible apomorphy, not found in most other Ulidiidae ), and sinuate tremae of the larval posterior spiracle (possible apomorphy of the few Aspistomella and Ulivellia whose larvae are known, but not found in the other Ulidiidae ), further supporting the inclusion of the genera Aspistomella and Ulivellia in a monophyletic lineage.

The species assigned here to the genera of the Aspistomella group have male genital structures so diverse that they would merit numerous separate genera, but we prefer to include most of them in the genera Aspistomella and Ulivellia . Morphology-based phylogenetic analysis does not provide a reliable resolution of their relationships, and the subdivision of this group into genera is tentative.

Many of the species discussed below are known from single specimens or only a few female specimens, so their descriptions lack male genitalia and can only be provisionally assigned to genera. Further study of additional material is required.

Key to adult Aspistomella View in CoL and related genera

1 Vein r-m far proximal to R 1 apex, at level with Sc apex; postero-apical lobe of cell cua shorter or at most as long as cell width; wing <3.5× longer than wide ( Figs 4 A–G, I–K View FIGURE 4 )............................................................. 2

- Vein r-m distal to R 1 apex ( Figs 5 View FIGURE 5 , 6 A, C–H View FIGURE 6 ), and if slightly proximal in Aspistomella garleppi ( Fig. 6 D View FIGURE 6 ), then distal to Sc apex level and wing> 3.5× longer than wide.................................................................... 9

2 Epistome low, clypeus as high as epistome ( Fig. 3 D View FIGURE 3 )......................................................... 3

- Epistome high, at least 2× higher than clypeus ( Figs 3 A, C View FIGURE 3 )........................................ Ulivellia View in CoL …… 4

3 Frontal and interfrontal setae 2–3 longer than fine parafrontal setulae ( Fig. 3 D View FIGURE 3 ). Wing pattern as in Figs 4 I, J View FIGURE 4 : subcostal crossband separated from preapical crossband in cell R 1; basicostal cell black or yellowish................................................................................... “ Euxesta” insolita Hendel View in CoL and related undescribed taxa

- Frontal vitta with uniformly short, undifferentiated setulae (as on Fig. 3 B View FIGURE 3 ). Wing pattern as on Fig. 4 K View FIGURE 4 : subcostal and preapical crossbands connected in pterostigma; basicostal cell black............................................. Axiologina View in CoL

4 Vein R 1 at pterostigma slightly dipping downwards, thickened and setulose; basal 0.8 of costal cell brown; vein R 2+3 at apex close and subparallel to costa, turning towards it before very apex. Wing sexually dimorphic: male with broad brown pattern, costa strongly bent at middle; female with 4 partly fused brown bands and moderately bent costa ( Figs 4 D, E View FIGURE 4 )................................................................................................ U. arcuata sp. nov.

- Vein R 1 at pterostigma straight, neither thickened nor setulose; at most basal half costal cell brown; vein R 2+3 variable. Wing either equal in both sexes, narrowly banded, or only one sex is known, costa only slightly bent anteriorly................ 5

5 Discal crossband (DB) oblique, ending just at base of vein CuA+CuP ( Figs 4 F, G View FIGURE 4 )................................. 6

- Discal crossband (DB) angulate, <-shaped, with posterior part directed posteroapically, terminating just at the apex of vein CuA+CuP ( Figs 4 A–C View FIGURE 4 )................................................................................ 7

6 Wing narrow,> 3.1× longer than wide, hyaline interspace between dark discal and preapical crossbands> 2.5× broader than discal crossband in discal cell ( Fig. 4 F View FIGURE 4 ). Legs and halter at least partly yellow. Prescutellar acrostichal setae absent............................................................................................. U. tenoris sp. nov.

- Wing wider, <3.1× longer than wide, hyaline interspace between dark discal and preapical crossbands <2× broader than discal crossband in discal cell ( Fig. 4 G View FIGURE 4 ). Legs and halter entirely black. Prescutellar acrostichal setae present.................................................................................................. U. pseudinsolita sp. nov.

7 Cell br entirely brown, connecting subbasal and discal crossbands ( Fig. 4 A View FIGURE 4 )....................... U. amnoni sp. nov.

- Cell br partly hyaline, subbasal and discal crossbands separated ( Figs 4 B, C View FIGURE 4 )...................................... 8

8 Subapical part of vein R 2+3 almost straight, leaving arcuate hyaline area between it and connection of subapical and apical brown crossbands ( Fig. 4 B View FIGURE 4 )............................................................... U. inversa Speiser View in CoL

- Subapical part of vein R 2+3 conspicuously bent anteriorly, leaving no hyaline area between it and connection of subapical and apical brown crossbands ( Fig. 4 C View FIGURE 4 )........................................................ U. laetitiae sp. nov.

9 Wing conspicuously widened and shortened in apical half; apical section of vein M 1 shorter than the preapical section between crossveins r-m and dm-m; postero-apical lobe of cell cua at most as long as cell width ( Fig. 4 H View FIGURE 4 ). Vein R 1 bare. Postpedicel dorsally incised and apically pointed ( Figs 48 E, F View FIGURE 48 ).................................. Polyteloptera apotropa Hendel View in CoL

- Wing gradually narrowed and elongated in apical half; apical part of vein M 1 longer than the preapical part between crossveins r-m and dm-m ( Figs 5 View FIGURE 5 , 6 View FIGURE 6 ). Vein R 1 either setulose or bare. Postpedicel rounded apically ( Fig. 3 C View FIGURE 3 )...... Aspistomella View in CoL … 10

10 Pterostigma vestigial, conspicuously shorter than width of costal cell ( Figs 5 A–F View FIGURE 5 )................................. 11

- Pterostigma either longer than width of costal cell ( Figs 6 B–H View FIGURE 6 ) or at most slightly shorter than its width (in A. lunata , A. enderleini sp. nov. and A. sachavaca sp. nov.) ( Figs 5 G–H View FIGURE 5 , 6 A View FIGURE 6 ).............................................. 16

11 Costal vein bent and thickened anteriorly before the apex of vein Sc; pterostigma at least slightly wider than veins Sc and R 1; vein R 1 bare or setulose dorsally; wing dark with hyaline spots, incisions or bands in the posterior half ( Figs 5 B–F View FIGURE 5 )...... 12

- Costal vein neither bent nor thickened before the apex of vein Sc, pterostigma almost linear, narrower than veins Sc and R 1; vein R 1 bare; wing with hyaline or pale yellow anterior margin and gradually darkening towards posterior margin without hyaline spots or incisions ( Fig. 5 A View FIGURE 5 ).............................................................. A. pachitea sp. nov.

12 Face between antennae from lunule to epistome completely sooty black microtrichose ( Fig. 25 D View FIGURE 25 ). Wing apex with long preapical hyaline incision from cell m only into cell r 2+3, dividing brown area into two dark apical bands; cell m with 2 hyaline marks ( Fig. 5 B View FIGURE 5 )..................................................................... A. heteroptera Hendel View in CoL

- Face between antennae from lunule to epistome white or gray microtrichose ( Figs 14 D View FIGURE 14 , 22 E View FIGURE 22 ). Wing pattern different: cell m with 1 hyaline spot only ( Figs 5 C–F View FIGURE 5 ).................................................................... 13

13 Wing apex completely dark; cell m with wide basal hyaline spot ( Figs 5 C–D View FIGURE 5 ); vein R 1 with several setulae dorsally; prescutellar acrostichal setae absent; halter with black knob............................................................. 14

- Wing apex with long and narrow preapical hyaline incision from apical half of cell m into cell r 1 usually reaching anterior margin of wing and separating narrow crescent-shaped brown apical band; base of cell m completely dark ( Figs 5 E–F View FIGURE 5 ); vein R 1 bare; prescutellar acrostichal setae present; halter yellow................................................... 15

14 Wing very narrow,> 3.5× longer than wide; anterior hyaline incision (distal to pterostigma) entering only into cell r 2+3 basal to crossvein r-m, but not cells br, nor r 4+5; base of cell r 4+5 and apex of cell dm with common oblique spot ( Fig. 5 D View FIGURE 5 ).................................................................................................. A. tres sp. nov.

- Wing somewhat wider, <3.5× longer than wide; anterior hyaline incision (distal of pterostigma) entering dm through cells r 2+3 and r

4+5

distal to crossvein r-m, but apex of cell dm entirely dark ( Fig. 5 C View FIGURE 5 )............................ A. duo sp. nov.

15 Cell dm entirely dark in apical half ( Fig. 5 E View FIGURE 5 ). Male genitalia: surstyli with short lobes and slightly thickened setae; lobes of phallic guide unmodified; phallus thick and extremely long, densely trichose ( Fig. 29 View FIGURE 29 ).............. A. lobioptera Hendel View in CoL

- Cell dm apically of r-m level with hyaline spot connected to hyaline spot in cell m 4 (=cua 1) ( Fig. 5 F View FIGURE 5 ). Male genitalia: outer surstylus with elongate anteroventral lobe and wide rounded posterior lobe, inner surstylus with 3–4 thick prensisetae and posteriorly produced; lobes of phallic guide sclerotised and apically truncated; phallus non-modified, moderately developed ( Figs 41 A–F View FIGURE 41 )......................................................................... A. steyskali sp. nov.

16 Cell r 1 with 2 hyaline marks ( Figs 5 G, H View FIGURE 5 ) (if in A. sachavaca sp. nov. with unclear indentation from cell r 2+3 ( Fig. 6 A View FIGURE 6 ), then wing short oval) or almost entirely hyaline with 1–2 narrow dark marks ( Fig. 6 B View FIGURE 6 )................................. 17

- Cell r 1 with 1 hyaline mark ( Figs 6 D, G, H View FIGURE 6 )............................................................... 23

17 Wing wide oval, <2.6× (2.3–2.5×) longer than wide ( Figs 5 G, H View FIGURE 5 ).............................................. 18

- Wing long,>2.6× (2.7–4×) longer than wide ( Figs 6 B, E, F View FIGURE 6 ).................................................. 20

18 First hyaline mark in cell r 1 reaching at most vein M 1 and isolated from hyaline mark at posterior margin of the wing ( Figs 5 G, H View FIGURE 5 ). Face entirely white or grey microtrichose ( Figs 22 E View FIGURE 22 , 31 C View FIGURE 31 ). Vein R 1 bare..................................... 19

- First hyaline mark in cell r 1 complete, reaching posterior margin of the wing ( Fig. 6 A View FIGURE 6 ). Face below lunule sooty black microtrichose ( Fig. 38 C View FIGURE 38 ). Vein R 1 setulose dorsoapically..................................... A. sachavaca sp. nov.

19 First hyaline incision distal to pterostigma long, extending into cell r 4+5 distally of crossvein r-m ( Fig. 5 G View FIGURE 5 ). Male genitalia not examined.................................................................... A. lunata (Hendel) comb. nov.

- First hyaline incision distal to pterostigma short, extending at most into cell r 2+3 and isolated from round hyaline spot in cell r 4+5 distally of crossvein r-m ( Fig. 5 H View FIGURE 5 ). Male genitalia: surstyli ventrally obtuse, with short denticles on posterior margin, but without expressed lobes and prensisetae, only with slightly thickened setae on surstyli, lobes of phallic guide laterally triangular and moderately sclerotised ( Figs 23 B–E View FIGURE 23 )................................................. A. enderleini sp. nov.

20 Wing apex mostly hyaline, with two narrow oblique dark crossbands ( Fig. 39 A View FIGURE 39 ). Epistome conspicuously produced anteriorly beyond antennae ( Fig. 39 D View FIGURE 39 ). Abdominal tergites 1–4 covered with moderately long and dense microtrichia, matt ( Figs 39 E, G View FIGURE 39 )................................................................................. A. schnusei sp. nov.

- Wing apex broadly dark ( Figs 6 C, E, F View FIGURE 6 ). Epistome shorter, not produced anteriorly beyond antennae ( Figs 10 B View FIGURE 10 , 36 D View FIGURE 36 , 42 D View FIGURE 42 ). Abdominal tergites subshining.......................................................................... 21

21 Wing with crescent-shaped subapical hyaline mark; posterior margin dark, without triangular hyaline incisions or spots in cell m 4 ( Fig. 6 C View FIGURE 6 ). Femora largely black.......................................... A. angustifrons (Hendel) comb. nov.

- Wing apex with subapical hyaline marks in cells r 1 and m 1 isolated; posterior margin with hyaline marginal mark in cell m 4 ( Figs 6 E, F View FIGURE 6 ). Femora yellow................................................................................ 22

22 Distal hyaline mark in cell r 1 reaching anterior half of cell r 4+5; cell m 1 with rounded submarginal hyaline mark extending to vein M 1 ( Fig. 6 E View FIGURE 6 )............................................................... A. crucifera (Hendel) comb. nov.

- Distal hyaline mark in cell r 1 posteriorly reaching only middle of cell r 2+3; cell m 1 with cuneiform hyaline mark extending middle of cell r 4+5 ( Fig. 6 F View FIGURE 6 ).......................................................... A. quinquincisa sp. nov.

23 Cell r 1 with hyaline marginal incision extending only to cell dm ( Fig. 6 G View FIGURE 6 ). Vein R 1 bare............. A. teresensis sp. nov.

- Hyaline interspace distally to pterostigma continuous, reaching posterior margin of wing ( Figs 6 D, H View FIGURE 6 ). Vein R 1 setulose in apical part.......................................................................................... 24

24 Wing 3.2–3.3× longer than wide; dark subapical crossband uniformly narrow, cell m 1 with hyaline area in basal half ( Fig. 6 H View FIGURE 6 ). Face below lunule with medial black spot, partly silvery microtrichose ( Fig. 32 G View FIGURE 32 ). Ocellar seta very long ( Figs 32 C, D View FIGURE 32 ). Mid tibia ventroapically with 2 long spur-like setae............................................... A. obliqua sp. nov.

- Wing 3.8–4× longer than wide; wing apex from crossvein dm-m uniformly brown, including cell m 1; hyaline area between it and pterostigma wide, covering entire area between r-m and dm-m ( Fig. 6 D View FIGURE 6 ). Face entirely silvery microtrichose, without black spot. Ocellar seta short, at most 1/3 as long as orbital seta ( Figs 3 C View FIGURE 3 , 24 D View FIGURE 24 ). Mid tibia ventroapically with 1 long spur-like seta................................................................................. A. garleppi sp. nov.

Key to third instar larvae of Aspistomella View in CoL and Ulivellia View in CoL

1. Creeping welt of abdominal segment I (CW1) composed of three spinule rows ( Figs 21 A View FIGURE 21 , 47 A View FIGURE 47 )......... Aspistomella View in CoL … 2

- Creeping welt of abdominal segment I composed of five spinule rows ( Figs 58 B View FIGURE 58 , 60 A View FIGURE 60 ).................... Ulivellia View in CoL …4

2. Creeping welts CW3–CW8 with single row of large spinules in rows d ( Figs 21 C, D View FIGURE 21 ); posterior spiracles: edges of yellow spiracular slit stripes distally blurred, as in Fig. 9 A View FIGURE 9 ...................................... Aspistomella duo sp. nov.

- Creeping welts CW3–CW8 with double row of large spinules in rows d ( Figs 47 C, D View FIGURE 47 ), posterior spiracles: edges of yellow spiracular slit stripes sharp.............................................................................. 3

3. Posterior spiracles close together (sd/sw ratio: ca. 0.05), spiracular slits with 12–15 turns, yellow slit stripes narrow, as in Fig. 9 B View FIGURE 9 ............................................................................ Aspistomella tres sp. nov.

- Posterior spiracles: distance between spiracles larger (sd/sw ratio: 0.19), yellow spiracular slits with 3–4 turns, yellow slit stripes wide, as in Fig. 9 F View FIGURE 9 ................................................................. Aspistomella sp.

4. Creeping welts CW3–CW8 with single row of large spinules in rows d, central row c continuous ( Figs 58 D, E View FIGURE 58 ); posterior spiracles: spiracular slits with 5–6 turns, yellow slit stripes narrow proximally and wide distally, resembling the silhouette of a bowing person, as in Fig. 9 D View FIGURE 9 ........................................................ Ulivellia arcuata sp. nov.

- Creeping welts CW3–CW8 with double row of large spinules in rows d, central row c discontinuous ( Figs 60 C, D View FIGURE 60 ); posterior spiracles: spiracular slits with 8–11 turns, yellow slit stripes almost equally wide proximally and distally, as in Fig. 9 C View FIGURE 9 ..................................................................................... Ulivellia tenoris sp. nov.

Genus Aspistomella Hendel, 1909 View in CoL

Aspistomella Hendel 1909b: 265 View in CoL .

Type species: Aspistomella lobioptera Hendel, 1909 , by original designation.

Aspistomella View in CoL : Hendel 1910: 10, 47; Steyskal 1968: 54.14.

Paraphyola Hendel 1909b: 261 View in CoL , syn. nov.

Type species: Paraphyola angustifrons Hendel, 1909 , by subsequent designation of Hendel (1910: 46).

Paraphyola View in CoL : Hendel 1910: 40, 46; Steyskal 1968: 54.20.

Diagnosis. This genus has all the diagnostic characters of the tribe Lipsanini : head and body with silver, greenish or cyan metallic sheen, sparsely whitish microtrichose, ocellar seta strong; phallus bare or covered with fine microtrichia, apically without glans; hypandrium symmetrical, with deep phallic guide, two areas of trichoid sensilla (corresponding to the pregonites and postgonites); female abdominal tergites 4–6 without anteromedial apodemes; only two spermathecae present.

Aspistomella species can be easily distinguished from other Lipsanini species by the combination of the following characters: epistome high, at least 2× higher than clypeus; frontal and interfrontal setae well differentiated, longer than fine and short parafrontal setulae; crossvein r-m distal to the apex of vein R 1 or at most very slightly proximal of it. Aspistomella duo , A. heteroptera , A. lobioptera , A. pachitea , A. steyskali , and A. tres also differ by the very narrow pterostigma formed by the approximate, subparallel veins Sc and R 1; costal vein more or less thickened, often strongly arcuate and bearing thickened marginal setulae before the apex of the subcostal vein. Aspistomella duo , A. heteroptera , A. obliqua , A. sachavaca , and A. tres can be distinguished by the presence of setulose vein R 1, which is rarely found in other groups of Lipsanini .

From the monotypic genus Polyteloptera , the species here assigned to Aspistomella differ by having an apically rounded postpedicel and a gradually narrowing wing with the long ultimate section of vein M 1 ( P. apotropa has an antenna with a dorsally incised and apically pointed postpedicel and an apically broadened wing, with the short ultimate section of vein M 1).

The species of Aspistomella are distinguished from the superficially similar, elongate and moderately large metallic greenish flies of the genus Ulivellia , by the crossvein r-m distal to the apex of vein R 1 (at the level of vein Sc or basal to it in Ulivellia ).

Description

Adult. Body mostly dark brown or black, usually with a metallic sheen, sometimes faint greenish or cyan, partly obscured by sparse whitish or silvery white microtrichia ( Figs 10 A, B View FIGURE 10 , 14 A, E View FIGURE 14 , 24 A–F View FIGURE 24 , 25 A–C View FIGURE 25 , 34 A–C View FIGURE 34 , 38 D View FIGURE 38 , 44 A–C View FIGURE 44 , and others).

Head at most slightly higher than long. Frons narrow (1.05× longer than wide in A. pachitea and 1.1× A. schnusei ) to wide (0.65× in A. duo , 0.9× in A. lobioptera ), conspicuously narrowed towards vertex, reddish or brownish-yellow with black, often greenish shining vertical plates and ocellar triangle ( Figs 10 D View FIGURE 10 , 14 C View FIGURE 14 and others); frontal vitta smooth or with fine longitudinal wrinkles ( Fig. 15 G View FIGURE 15 , left), frontal plates with 4–8 short pro- or lateroclinate parafrontal setulae on each side ( Figs 13–18 View FIGURE 13 View FIGURE 14 View FIGURE 15 View FIGURE 16 View FIGURE 17 View FIGURE 18 ), 2–6 frontal setae and 1–3 interfrontal setae ( Figs 10 C, D View FIGURE 10 , 15 A View FIGURE 15 , 22 E View FIGURE 22 , 24 C View FIGURE 24 , 25 D View FIGURE 25 , 27 B View FIGURE 27 , 28 C View FIGURE 28 , 31 C View FIGURE 31 , 32 G View FIGURE 32 , 34 D, E View FIGURE 34 , 36 C View FIGURE 36 , 38 B, C View FIGURE 38 , 39 C View FIGURE 39 , 40 A View FIGURE 40 , 42 F View FIGURE 42 , 44 D View FIGURE 44 ); vertex and orbits with dense, proclinate, slightly thickened, short white microtrichia ( Fig. 15 G View FIGURE 15 , right). Vertical plates with 1 long orbital seta, and usually with a second vestigial reclinate seta anterior to it; ocellar triangle with ocelli forming an elongated acute triangle and a pair of usually very long ocellar setae, 4–10× longer than frontal setae ( Figs 14 B View FIGURE 14 , 22 D View FIGURE 22 , 25 C View FIGURE 25 , and others) or, in A. garleppi , 1–1.5× longer than frontal setae ( Fig. 24 D View FIGURE 24 ). Face dorsally flat, covered with white or black, dense, inclinate, curled and appressed microtrichia ( Figs 15 F View FIGURE 15 , 25 D View FIGURE 25 , 38 C View FIGURE 38 and others), and with ventral portion conspicuously produced antero-ventrally, forming a large, often metallically shining epistome, covered with sparse, thin, erect whitish microtrichia ( Figs 14 C View FIGURE 14 , 15 I View FIGURE 15 and others). Parafacial usually brownish-yellow, densely white microtrichose ( Fig. 15 H View FIGURE 15 ). Gena widened posteriorly, brownish, sparsely white microtrichose, bearing 1–2 strong genal setae and 4–6 moderately or very long, proclinate peristomal setulae ( Figs 25 C View FIGURE 25 , 27 B View FIGURE 27 , 28 B, C View FIGURE 28 ). Occiput black or dorsomedially and lateroventrally brown to yellow, densely white microtrichose. Scape short, dorsally with 6–8 marginal setulae; pedicel with rectangular incision; one long pedicellar seta and 20–25 setulae, the longest ventral, almost as long as pedicel width ( Fig. 15 D View FIGURE 15 and others). Postpedicel oval, 2.0–2.2× longer than wide, rounded at apex, densely microtrichose; arista 3-segmented ( Figs 15 D, E View FIGURE 15 ). Palp elongate, 4–4.5× longer than wide and almost twice as long as postpedicel, white microtrichose, with 15–25 ventral and ventromedial setae almost as long as palp width and 1–2 apical setae 1.5–2× longer than the rest ( Fig. 15 B View FIGURE 15 ). Clypeus much lower than epistome, convex, densely microtrichose, almost entirely hidden in oral cavity. Proboscis with prementum and labellum moderately large, with long, often slightly curled, thin setulae ( Fig. 15 B View FIGURE 15 ).

Thorax mostly black or brown, with faint bluish, greenish or golden sheen and moderately dense, thin, semierect white microtrichia not completely hiding shining cuticle. Postpronotal lobe with 1 seta and 5–15 setulae; proepisternum with 1 short proepiesternal seta (sometimes absent or indistinct) and 5–20 fine subequal setulae on proepiesternal ridge. Prosternum higher than wide, with distinct longitudinal suture and shallow ventral incision, sparsely microtrichose, without conspicuous setae. Mesonotum steel shining, with slight green or bluish sheen of cuticle not entirely obscured by sparse or moderately thick white microtrichia ( Figs 14 E View FIGURE 14 , 16 A View FIGURE 16 ); scutum setulose, except prescutellar area posterior to dorsocentral setae; dorsocentral setulae forming distinct row, with 8–18 poorly ordered rows of setulae between them; 1 postpronotal, 2 notopleural, 1 postsutural supra-alar, 1 intra-alar and 1 postalar, 2 dorsocentral and 1 acrostichal seta present, strong (in A. crucifera , A. lobioptera , A. lunata , A. obliqua , A. quinquincisa , A. sachavaca , A. steyskali , and A. teresensis ), weak (in A. schnusei ,) or absent (in A. angustifrons , A. duo , A. garleppi , A. heteroptera Hendel , A. pachitea , and A. tres ). Anepisternum setulose almost entirely anterior to vertical squama, with one strong seta and 5–12 shorter setae at posterior margin. Katepisternum with one strong posterodorsal seta. Scutellum subtriangular, slightly convex, smooth, with 2 pairs of scutellar setae, no setulae on disc and margins, sparsely microtrichose. Subscutellum matt, microtrichose. Mediotergite smooth, sparsely or densely microtrichose.

Wing moderately wide, 2.6–2.8× longer than wide (in A. lobioptera , A. lunata , A. enderleini , and A. steyskali ) to strongly elongate, 3.5–3.75× longer than wide (in A. tres ), largely dark with hyaline incisions or round spots or hyaline with darker crossbands ( Figs 5 View FIGURE 5 , 6 View FIGURE 6 ). Venation modified, pterostigma shorter than wide in A. duo , A. heteroptera , A. lobioptera , A. pachitea , A. steyskali , and A. tres ), and costal vein conspicuously bent and thickened before apex of subcostal vein, so that apical portion of costal cell conspicuously lobed with thick setulae on costal vein ( Figs 5 B–F View FIGURE 5 , 16 E, F View FIGURE 16 ), or, in other species, as long as or longer than wide. Vein R 1 dorsally with 3–10 setulae ( A. duo , A. heteroptera , A. obliqua , A. sachavaca , and A. tres ) to entirely bare. Crossvein r-m apical to R 1 apex in most species, except in line with R 1 apex in A. lunata , A. sachavaca , and A. schnusei or slightly basal of it in A. garleppi Cell r 4+5 slightly or conspicuously narrowed apically. Vein CuA Z-shaped, cell cua with moderately short (in most species) to long (in A. obliqua and A. sachavaca ) posteroapical lobe along vein CuP. Calypters white, with white cilia. Halter creamy white (in most species) to brown with black or brown knob (in A. duo and A. tres ).

Legs variably coloured; coxae usually yellow to brown, femora yellowish brown to entirely brown or black, tibiae and tarsi yellow to brown. Fore femur with 2 posterodorsal rows of setae; posteroventrally with row of 3–5 setae (in A. duo and A. tres ), often conspicuously thickened and spinulose, in A. pachitea without postero-ventral row; fore tibia antero-apically as well as basal fore tarsomere with brush of short pale setulae. Mid femur on anterior and posterior surfaces with row of moderately long, antero- and postero-ventrally directed setulae. Mid tibia apicoventrally with strong spur-like seta 1.5–3× longer than tibia width, in A. duo , A. obliqua and A. tres with additional seta ( Fig. 16 N View FIGURE 16 ) 0.7–1× longer than longest seta. Hind femur with 2–3 to 4–5 preapical setae dorsally.

Abdomen subshining brown to black, in A. heteroptera with syntergite 1+2 broadly yellow ( Figs 25 A, B View FIGURE 25 ); tergites and sternites with metallic silver, green or yellow sheen, sparsely white microtrichose ( Figs 10 G View FIGURE 10 , 16 C, D View FIGURE 16 , 34 B View FIGURE 34 , 36 G, H View FIGURE 36 ). Male and female abdomen with 5 visible tergites, of which tergites 3–5 subequal in length; female tergite 6 hidden beneath posterior margin of tergite 5. Sternites 3–5 not modified, subrectangular ( Figs 11 A View FIGURE 11 , 12 A View FIGURE 12 , 18 A View FIGURE 18 , 23 A View FIGURE 23 , 35 A View FIGURE 35 , 37 A View FIGURE 37 , 41 G View FIGURE 41 ).

Male postabdomen. Sternite 8 setulose, with 2 longer setulae. Epandrium moderately setulose. Surstyli of variable shape, often species-specific, fused with epandrium, usually wide basally, narrowed and curved mesally towards apex, often with small, mesoapically directed pimple-like process ( Figs 26 E View FIGURE 26 , 35 C View FIGURE 35 , 46 E View FIGURE 46 : pi p), in A. angustifrons , very narrow and curved ( Figs 11 C, F View FIGURE 11 ), in A. quinquincisa , blunt with smooth margins ( Figs 37 B–F View FIGURE 37 ), in A. enderleini , blunt with serrate posterior margin ( Figs 23 B–D View FIGURE 23 ), in A. lobioptera and A. steyskali , bilobate ( Figs 29 D–G View FIGURE 29 , 41 A–D View FIGURE 41 ). Cerci low, moderately protruding posteriorly, flattened and almost round in A. duo and A. tres ( Figs 16 B, D View FIGURE 16 , 46 A, B View FIGURE 46 : cerc), elongate and bilobate, partly connected in A. heteroptera , A. obliqua , and A. pachitea ( Figs 26 C, D View FIGURE 26 , 33 A, B, D View FIGURE 33 , 35 C, D View FIGURE 35 ), short, button-like in A. angustifrons ( Fig. 11 E View FIGURE 11 ), flat in A. enderleini , A. lobioptera , A. quinquincisa , and A. steyskali ( Figs 23 B, D View FIGURE 23 , 29 D–F View FIGURE 29 , 37 E, F View FIGURE 37 , 41 A–E View FIGURE 41 ), moderately long setulose. Subepandrial sclerite short, horseshoe-shaped, with ventral arms entirely integrated into surstylus, bearing a row of mesally directed setulae anteriorly ( Figs 23 B View FIGURE 23 , 29 D, E View FIGURE 29 ), without thickened tooth-like setulae (prensisetae), or with 3–4 sharp thickened subapical setae (in A. quinquincisa and A. steyskali ) ( Figs 37 E, F View FIGURE 37 , 41 A, B View FIGURE 41 ) or with 1–2 thickened and 2–3 fine setulae (in A. heteroptera ) ( Fig. 26 E View FIGURE 26 ). Hypandrium narrow, U-shaped, joined to basiphallus by thin membrane; phallapodeme with paired and rather long, sagitally flattened anterior arms, continuing into a pair of ventrally directed ridges or folds, forming a gutter-like phallic guide on both sides of basiphallus and basal part of distiphallus ( Figs 23 D–F View FIGURE 23 , 29 F View FIGURE 29 , 35 G View FIGURE 35 , 37 E, F View FIGURE 37 , 41 A, D, E View FIGURE 41 ); phallapodeme itself usually very short ( Fig. 46 C View FIGURE 46 ). Pregonites plate-like, almost symmetrical, antero-medially attached to phallapodeme and latero-ventrally to hypandrium by membrane, each bearing 4–6 setulae ( Fig. 33 E View FIGURE 33 ); in A. obliqua very strongly displaced and posterior to postgonites ( Figs 11 H View FIGURE 11 , 23 D View FIGURE 23 , 26 F View FIGURE 26 , 29 F View FIGURE 29 , 37 F View FIGURE 37 , 41 D View FIGURE 41 ). Postgonites symmetrical, reduced to oval sensillar plates, bearing 4–5 setulae located on membrane on lateral sides of phallic guide at level of anterior part of basiphallus ( Figs 11 H View FIGURE 11 , 23 D View FIGURE 23 , 26 F View FIGURE 26 , 29 F View FIGURE 29 , 33 E View FIGURE 33 , 37 F View FIGURE 37 , 41 D View FIGURE 41 ).

Phallus simple, band-like or tubular, moderately short in most species, usually 2–5× longer than epandrium ( Figs 11 B, C View FIGURE 11 , 17 K View FIGURE 17 , 23 B, C View FIGURE 23 , 26 A, B View FIGURE 26 , 33 B View FIGURE 33 , 35 C, D View FIGURE 35 , 37 B, C View FIGURE 37 , 41 A–F View FIGURE 41 , 46 A, B View FIGURE 46 ), in A. lobioptera very long, 6–8× longer than epandrium, with moderately large ring of basiphallus ( Figs 23 B, D View FIGURE 23 , 29 F View FIGURE 29 ) and simple distiphallus represented is a bare or short spinulose, densely trichose in A. lobioptera ( Fig. 29 C View FIGURE 29 ), flattened tube with a pair of narrow sclerotised stripes.

Ejaculatory apodeme usually with fan-shaped apical part 2–3 as long as moderately developed sperm pump ( Figs 17 J View FIGURE 17 , 23 C View FIGURE 23 , 26 G View FIGURE 26 , 29 B View FIGURE 29 , 35 E View FIGURE 35 , 37 C View FIGURE 37 ).

Female terminalia. Oviscape short, usually half as long as abdomen. Eversible membrane ( Figs 12 A View FIGURE 12 , 18 B View FIGURE 18 ) as long as aculeus, with 2 pairs of bare taeniae reaching its posterior 4/5; membrane finely microspinulose in anterior half. Aculeus ( Figs 12 B View FIGURE 12 , 18 C View FIGURE 18 ) narrow and moderately long, 8–11× longer than wide, with elongate oval cercal unit bearing 2–3 pairs of long setae (basal dorsal, subapical lateroventral and somewhat shorter apicodorsal) and several shorter setulae on ventral and latero-ventral sides; anal opening (slit) on dorsal side of cerci ( Figs 12 B View FIGURE 12 , 18 D View FIGURE 18 ); sternite 8 well sclerotised, with indistinct setulae on each side. Vagina ( Fig. 30 D View FIGURE 30 ) with simple, finger-like, apically rounded ventral receptacle as in all other Ulidiidae . Two spherical or somewhat elongated spermathecae ( Figs 12 C View FIGURE 12 , 18 E View FIGURE 18 , 30 B View FIGURE 30 , 43 E View FIGURE 43 ); short, invaginated “necks” at junctions of spermathecae with ducts.

Preimaginal stages

Egg known only for A. lobioptera , elongate oval, 4.5× longer than wide, slightly wider than aculeus, without stalks or tapering rear end ( Fig 30 C View FIGURE 30 ).

Third instar larvae. Aspistomella larvae are known from A. duo , A. tres and an unidentified third species. Larva whitish, maggot-like, medium-sized (7.5–12 mm long) and mostly smooth. Pseudocephalon with deep incision between cephalic lobes. Facial mask with 20–28 laterally bifurcated or double bifurcated furrows separating subparallel oral ridges, each with fringe of blunt scales ( Figs 19 C, D View FIGURE 19 ). Creeping welts on abdominal segments I–VII and caudal segment with spinules, anterior spiracles fan-shaped. Caudal segment posterodorsally truncate, with sclerotised, slightly elevated, rounded posterior spiracles. Each spiracular plate with three very long, sinuous spiracular slits, spiracular plate mostly dark, but yellowish along spiracular slits (= yellow slit stripes) ( Figs 9 A, B View FIGURE 9 ). Cephalopharyngeal skeleton ( Figs 7 A, B View FIGURE 7 ). Mouthhook with elongated base with semitransparent window, long and thin, ventrally curved apical tooth and moderately long and narrow, ventrally produced ventral apodeme, moderately short hypopharyngeal sclerite.

Puparium ( Fig. 9 G View FIGURE 9 ) elongate cylindrical, reddish brown, strongly tapering anteriorly and broadly truncate posteriorly; anterior spiracles, posterior spiracles and creeping welts as described for third instar larva.

Habitat. The larvae of Aspistomella duo , A. tres and A. sp. live in the water-filled internodes of young bamboo culms of Guadua angustifolia . The biology of Aspistomella is described in the Biology section of A. duo .

Remarks. The monophyly of the genus is doubtful, and the results of a morphology-based phylogenetic analysis neither strongly support nor refute it. Here we include in Aspistomella all species that do not have the diagnostic characters of Polyteloptera and / or Ulivellia . There is no evidence that they are nested within Aspistomella , and we consider them to be separate genera.

The diagnostic character proposed by Hendel (1909b, 1910) to separate Paraphyola from Aspistomella , namely a narrow wing with almost parallel anterior and posterior margins, shows a similar condition with a gradual transition to "moderately oval" wings in A. duo , A. crucifera , A. garleppi , A. obliqua , A. quinquincisa , A. schnusei , A. teresensis , and A. tres , so we prefer to place them all in Aspistomella together with Paraphyola angustifrons rather than separating them into two genera for ease of identification. As male genitalia are not examined in many species and molecular data are not available to provide more robust results, we consider the current generic classification to be tentative.

Steyskal (1968: 54.20) erroneously stated P. angustifrons to be the type species of Paraphyola by monotypy. However, Hendel (1909a) originally described two species, but did not designate any of them as the name-bearing type. It was only in the later published generic synopsis of the ulidiids in the series “Genera Insectorum” ( Hendel, 1910) that he clearly stated that his type species was Paraphyola angustifrons Hendel, 1909 . The latter is therefore the type species of the genus according to the later designation by Hendel (1910).

The unknown ulidiid larva found by DK in water-filled bamboo internodes probably belonged to Aspistomella , because CW1 had three rows of spinules (and not five as in Ulivellia ) and all other details of its creeping welts resembled A. tres . The spiracular slits of the unknown Aspistomella sp. larvae were clearly identifiable as belonging to a distinct species, because they had only a few turns and the yellow area above the spiracular slits was very wide compared to the larvae of other ulidiid species ( Fig. 9 F View FIGURE 9 ). Furthermore, the shape of the anterior spiracles was more angular than in Aspistomella duo and A. tres .

Kingdom

Animalia

Phylum

Arthropoda

Class

Insecta

Order

Diptera

Family

Ulidiidae

Loc

Aspistomella

Kovac, Damir, Kameneva, Elena P., Korneyev, Severyn V., Araújo, Alexandre Santos, Savaris, Marcoandre, Smit, John T., Schneider, Alexander, Schreiber, Robert & Korneyev, Valery A. 2024
2024
Loc

Aspistomella

Hendel, F. 1910: 10
1910
Loc

Paraphyola

Hendel, F. 1910: 40
1910
Loc

Aspistomella

Hendel, F. 1909: 265
1909
Loc

Paraphyola

Hendel, F. 1909: 261
1909
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