Archiinocellia protomaculata ( Engel, 2011 ) Archibald & Makarkin, 2021

Archibald, S. Bruce & Makarkin, Vladimir N., 2021, Early Eocene snakeflies (Raphidioptera) of western North America from the Okanagan Highlands and Green River Formation, Zootaxa 4951 (1), pp. 41-79 : 58-64

Archiinocellia protomaculata ( Engel, 2011) , comb. nov.

Figs 12–17

Material examined. Thirty-six specimens collected by David D. Kohls in 2004 to 2009 at three localities in Garfield County (Colorado, U.S.A.): (1) UCM locality 2005026 (Labandeira Site): specimens UCM 35717 (female); UCM 40659 (male); UCM 58068 (sex unknown); UCM 59090 (female); UCM 59509 (male); UCM 60297 (female); UCM 61231a (female); UCM 61231b (male); UCM 61231с (sex unknown); UCM 62026 (male); UCM 62523 (sex unknown); UCM 62525 (three specimens: male, female, and sex unknown); UCM 62687 (female); UCM 64319 (sex unknown); UCM 64727 (female); UCM 65146 (sex unknown); UCM 66134 (sex unknown); UCM 67986 (female); UCM 72989 (female); UCM 73247 (female); (2) UCM locality 2006032 (Donnell Site): UCM 40057 (male); UCM 42684 (male); UCM 42744 (female); UCM 45968 (female); UCM 45969 (female); UCM 45970 (female); UCM 53268 (female); (3) UCM locality 2007238 (Denson Site): UCM 41697 (male); UCM 42105 (female); UCM 42107 (female); UCM 42122 (sex unknown); UCM 54806 (male); UCM 55329 (female); UCM 56552 (female); the Parachute Creek Member of the Green River Formation; all are deposited in UCM collections.

Type locality and horizon. Probably Labandeira Site in the Parachute Creek Member of the Green River Formation , Garfield County, Colorado, U.S.A.; early Eocene (Ypresian) .

Diagnosis. May be distinguished from A. oligoneura by markedly narrower fore- and hind wings (see above).

Redescription (based on the holotype and material from UCM listed above): body length of male 8.1–9.2 mm (five specimens), female (without ovipositor) 8.7–9.1 mm (seven specimens). Head broadly oval, somewhat narrowed posteriorly (in dorsal view); dorsal maculation very distinct with extensive pale areas ( Fig. 15 ), especially beyond and around eyes, broad medial stripe (sometimes divided by a thin brown medial line: Fig. 15F ), and most areas before the eyes. Palpi brownish. Antennae: scapus brown to dark brown; flagellum distinctly paler, with slightly more than 40 segments. Ocelli not detected.

Pronotum dark brown, relatively short, 1.20–1.50 mm long; maximum width 0.57 mm in dorsal view, 0.65 mm in lateral view. Meso- and metathorax dark brown.

Legs pale except brown in proximal half of femora and in coxae ( Fig. 12 ).

Abdomen of male ( Fig. 13 ): Tergites, sternites dark, easily distinguished; T3 to T7, and S3 to S7 nearly equal, moderately long; T8 slightly shorter than T3 to T7, extending laterally notably more than these tergites; T9 short, extending laterally to ventral margin; ectoproct short, elongate in lateral view, ventrally appears narrowed; gonocoxites 9 not discernible (as in Fig. 2A ) or apparently weakly sclerotized (see Fig. 13B ). Abdomen of female ( Fig. 13C ): Tergites, sternites dark, easily distinguished; T8, T9 relatively short; ectoproct small, elongate in lateral view; ovipositor moderately long, 3.25–3.80 mm (0.27–0.30 body length).

Forewing relatively narrow, 7.4–8.0 mm long, 1.8–2.2 mm wide (length/width: 3.63–4.15, measured in five females). Costal space narrow, most dilated at proximal 1/6 wing length; at maximum four subcostal veinlets simple preserved (basal regions not preserved, probably six total), widely spaced. ScP very short, terminating on costal margin approximately at just basad mid-wing, strongly distad crossvein 2ra-rp; from termination of ScP to pterostigma approximately double length of pterostigma. Subcostal space: basal crossvein 1scp-r closer to divergence of M, CuA than to origin of RP; distal crossvein (2scp-r, forming basal margin of pterostigma) straight to incurved. Pterostigma elongate (4.9–5.1 times longer than width measured along to costal margin), lightly pigmented (from very pale to slightly brownish as preserved). Two branches of RA: one within pterostigma usually easily visible, strongly inclined, in middle of pterostigma; branch forming distal margin of pterostigma usually straight; no branch distad pterostigma; portion of RA distad pterostigma short. RA space with two crossveins forming two radial cells nearly equal in length: 2ra-rp usually relatively short, located about mid-way between pterostigma, termination of ScP; 3ra-rp located in distal part of pterostigma along RA. RP originating at nearly half (0.43–0.48) wing length, usually with four pectinate branches (rarely three): RP1 deeply forked (at about half or somewhat less its length), with anterior branch simple, posterior forked; other branches (RP2 to RP4) simple. One long intraradial crossvein rp1-rp2 between stems of RP, RP1. Three crossveins between RP, MA: 1r-m, 2r-m connecting stem of RP, MA; 3r-m connecting RP1, anterior branch of MA. M close to R (appears fused) for considerable distance, and then fused with CuA for short distance; forked well proximad origin of RP. MA deeply dichotomously forked twice. MP strongly zigzagged, anterior trace with two (rarely one) simple branches. Two intramedian crossveins form two doi, approximately equal in length (sometimes distal doi longer): 2im very long, joining posterior branch of MA near origin or at forking of MA. CuA fused with M for short distance; anterior trace CuA strongly zigzagged, fused with MP for long distance to form part of posterior margin of basal doi, with one long simple branch. CuP simple, aligned with Cu. Crossvein icu between CuA, CuP long. Crossveins between CuP, AA1, between AA1, AA2 short. AA1, AA2 simple; region of AA3 poorly discernible.

Hind wing 6.5–7.1 mm long, 1.7 mm wide (length/width: 3.82). Costal space narrow; at maximum three discernible simple subcostal veinlets, very widely spaced (slightly wider than in forewing). ScP very short, terminating at approximately mid-wing; from termination of ScP to pterostigma approximately twice length of pterostigma. One crossvein detected in subcostal space, 2scp-r, forming basal margin of pterostigma, straight to incurved. Pterostigma elongate (4.5–5.1 times longer than width measured along to costal margin). RA with two branches: one within pterostigma usually clearly visible, strongly inclined, located medially in pterostigma; branch forming distal margin of pterostigma usually straight; no branches distad pterostigma; portion of RA distad pterostigma short. Three crossveins between RA, RP: 1ra-rp located slightly proximad 2r-m (rarer probably distad: Fig. 17B ); 2ra-rp located far distad termination of ScP; 3ra-rp located just distad middle of pterostigma along RA. RP originating 0.33 wing length, with three to four pectinate branches, two proximad 3ra-rp, one to two distad. RP1 deeply forked, anterior branch simple, posterior forked; other branches (RP2 to RP4) simple. One intraradial crossvein rp1-rp2 between stems of RP, RP1. Three crossveins between R, M: 1r-m long, subparallel to R connecting to RP near its origin (its base unclear); 2r-m connecting stems of RP, MA; 3r-m connecting RP1, anterior branch of MA. M forked distad origin of RP. MA deeply dichotomously forked twice. MP: anterior trace zigzagged, simple, with three simple, pectinate branches. Two intramedian crossveins form two doi, proximal much shorter than distal. Two crossveins between M, Cu: basal 1m-cu relatively short, oblique, connecting M, CuA; 2m-cu connecting MP, CuA. Anterior trace of CuA simple distally, with one simple branch. CuP fused with AA1 (incompletely preserved).

Remarks. This species was described based on 19 specimens from the National Museum of Natural History ( Engel, 2011). The specimen mentioned and figured by Dayvault et al. (1995: Fig. 35) belongs to this species ( Engel, 2011), as does Aspöck’s (1998: Fig. 7 ) specimen (determination here by head maculation). We examined highquality photographs of 51 specimens in the UCM collections. Most are only partially visible pending preparation, as the rock in these beds does not split along fine, thin laminae bearing the fossil as is usually the case in Okanagan Highlands fossils, and portions often protrude into the matrix. We confidently assign 36 of these to A. protomaculata by their characteristic head maculation and/or wing venation. Five further fossils possibly belong to this species, but cannot be assigned with certainty by preservation: three females from the Labandeira Site (UCM 60298; UCM 62430; UCM 65597), a probable male (UCM 42748) from the Donnell Site, and a female (UCM 75788) from the UCM locality 2009063 (Claudia’s Place in Garfield County). Characters present on the remaining ten more poorly preserved specimens suggest that at least some of them might also belong to the species, including four from the Labandeira Site: UCM 61231d (sex unknown); UCM 61320 (male), UCM 62686 (sex unknown), UCM 62689 (sex unknown), UCM 68352 (sex unknown); two from the Donnell Site: UCM 39661 (sex unknown) and UCM 39662 (female); two from Claudia’s Place: UCM 75468 (sex unknown) and UCM 75790 (male); and one from the Denson Site: UCM 56842 (sex unknown).

The maculation of the head in Archiinocellia protomaculata is distinctive ( Fig. 14 ). Paler areas occupy most of the lateral part of its head, whereas there are paler streaks in some extant genera of both families; others are mostly dark, lacking reddish regions (see e.g., Albarda 1891). While the life colour of these pale regions of the head of A. protomaculata is unknown, lighter regions in the heads of extant species are reddish. Carpenter (1936) stated that the size of the reddish patches varies freely within Raphidiidae species (p. 9), but the colour patterning of the heads of A. protomaculata is strongly consistent among individuals, and so we consider it to be a good diagnostic character state of the species.

The maculation of the holotype head is typical of the species, but its wings are overlapping and many portions are obscured by the abdomen ( Fig. 12A ; Engel, 2011: Figs 1 , 2 ). The venation of both the fore- and hind wings ( Figs. 16A, B ), is generally similar to that of other examined specimens (e.g., Figs. 16C–D ; 17A–D ).

Wing venation shows little variation among the specimens examined. There are usually two branches of RP distad 3ra-rp, but in one specimen there is only one ( Figs. 17C–D ) in both the forewing and hind wing. The length of 3ra-rp varies from long (e.g., Fig. 17B–C ) to short (e.g., Figs. 16A, C ), also in both wings. In forewings, crossvein 2im usually joins the posterior branch of MA (e.g., Fig. 17C ), but may rarely be at its forking (e.g., Fig. 17A ), and while MP usually has two branches, it may rarely have one ( Engel, 2011: Figs 4 , 7 ). In hind wings, the size of the basal doi varies from relatively long ( Fig. 17D ) to short ( Fig. 17B ), and crossvein 2r-m may be placed relatively distally (distad 2ra-rp: Fig. 17D ; Engel, 2011: Fig. 7 ) to proximally (proximad 2ra-rp: Fig. 17B ). These variations are normal for the suborder, and in some species are greater (see e.g., Zelený, 1969).

Figures

FIGURE 1. Megaraphidia antiquissima sp. nov., holotype RBCM P1555, specimen as preserved. Scale bar = 5 mm.

FIGURE 2. Wing venation of Megaraphidia antiquissima sp. nov., holotype RBCM P1555. A, right forewing, B, left forewing; C, right forewing; D, left hind wing. Scale bar = 3 mm (all to scale).

FIGURE 4. Wing venation of Megaraphidia ootsa sp. nov., holotype RBCM P1556. A, right forewing, B, left forewing; C, right hind wing; D, left hind wing. Scale bar = 3 mm (all to scale).

FIGURE 7. Megaraphidia klondika sp. nov., specimen SR16-004-001-01.01. A, specimen as preserved; B, drawing. Scale bar = 3 mm (both to scale).

FIGURE 12. Archiinocellia protomaculata (Engel, 2011), females. A, holotype USNM 31487 (counterpart); B, UCM 42107. Scale bars = 5 mm.

FIGURE 13. Archiinocellia protomaculata (Engel, 2011), abdomen. A, UCM 61231b, male (lateral/slightly dorsal view); B, UCM 42684, male (lateral view); C, UCM 45968, female. e, ectoproct; gx9?, possible gonocoxites 9; ov, ovipositor; S2 to S7, second to seventh sternites; T3 to T9, third to ninth tergites. Scale bars = 1 mm.

FIGURE 14. Archiinocellia protomaculata (Engel, 2011).A, UCM 61231, rock with four specimens of A. protomaculata (a–c) and one undeterminable Raphidioptera (d); B, UCM 62523. Scale bars = 1 mm.

FIGURE 15. Archiinocellia protomaculata (Engel, 2011), head and prothorax (dorsal view). A, UCM 64727; B, UCM 62525; C, UCM 35717; D, UCM 45969; E, UCM 42122; F, UCM 62687. Scale bars = 1 mm.

FIGURE 16. Venation of Archiinocellia protomaculata (Engel, 2011), holotype USNM 31487. A, right forewing; B, left forewing; C, right hind wing. Scale bar = 3 mm (all to scale).

FIGURE 17. Venation of Archiinocellia protomaculata (Engel, 2011). A, right forewing of UCM 61231a (female); B, right hind wing of the same specimen; C, right forewing of UCM 62523; D, right hind wing of the same specimen. Scale bar = 3 mm (all to scale).