A revision of the late Eocene snakeflies (Raphidioptera) of the Florissant Formation, Colorado, with special reference to the wing venation of the Raphidiomorpha Author Makarkin, Vladimir N. Author Archibald, S. Bruce text Zootaxa 2014 3784 4 401 444 journal article 46097 10.11646/zootaxa.3784.4.4 16f488d9-eb77-4ff5-a7ce-99eeb7875371 1175-5326 252218 D5E03502-7BD3-41F4-A4CF-5537B1462A23 Archiraphidia tranquilla ( Scudder, 1890 ) ( Figs 12–16 ) Raphidia (?) tranquilla Scudder, 1890 : 154 , 155; Pl. 14, Fig. 2 ; Scudder 1891 : 369 ; Handlirsch 1906–1908 [1907]: 907; Rohwer 1909 : 534 ; Cockerell & Custer 1925 : 296 . ? Archiraphidia tranquilla : Handlirsch 1910 : 104 . Archiraphidia ? tranquilla : Aspöck et al . 1991 : 538 , 665, Fig. 3050. “ Raphidia ” tranquilla : Engel 2002 : 22 . Raphidia tranquilla : Carpenter 1936 : 143 , 146, 150, Fig. 12 ; Oswald 1990 : 160 ; Engel 2003 : 125 ; Engel 2009 : 213 (Tabl. 1); Engel 2011 : 79 (Tabl. 2). Inocellia eventa Scudder, 1890 : 160 ; Handlirsch 1906–1908 [1907]: 907; Cockerell & Custer 1925 : 297 ; Carpenter 1936 : 143 , 145, 146, 149 (as a synonym of Inocellia tumulata ); Oswald 1990 : 161 (as a synonym of I. tumulata ); Engel 2002 : 22 (as a synonym of “ Raphidia ” tumulata ); Engel 2003 : 125 (as a synonym of R. tumulata ), syn. nov. ? Archiraphidia eventa : Handlirsch 1910 : 104 . Archiraphidia ? eventa : Aspöck et al. 1991 : 538 , 665. Diagnosis. May be distinguished from other species by forewing CuA1 forked [simple in A. tumulata ]; by short pterostigma [elongate in A.? somnolenta ]. Redescription. Specimen MCZ 246, holotype of Raphidia tranquilla ( Figs 12 , 13 ). Pterothorax poorly preserved; details not visible. Legs not preserved. Abdomen rather well preserved; with eight dark segments separated by broad pale intersegmental membrane between fourth to eighth segments. Apex of abdomen unclear, but surely male. Forewing long approximately 8.5 mm (estimated), 2.6 mm wide (length/width ratio 3.2). Costal space incompletely preserved, moderately broad. Distal part of ScP not preserved. Three preserved subcostal veinlets (branches of ScP) distad M+CuA fusion simple, widely spaced. In subcostal space, one preserved crossvein between ScP, RA well proximad origin of RP ; other crossvein closes pterostigma proximally. Pterostigma short, dark. RA with one branch, closing pterostigma distally; no branches within pterostigma detected ( Fig. 9 D); portion of RA distad pterostigma short, deeply curved. Two crossveins between RA, RP ; 3ra-rp located within pterostigma, in its proximal half. Anterior trace of RP forked apically, with two branches proximad 3ra-rp, one distad. RP 1 forked, one branch simple, other rather shallowly forked; RP 2 simple; RP 3 simple (left wing), shallowly forked (right wing). Two intraradial crossveins, between branches of RP 1, PR2 at its origin, and RP 2, RP 3. Three crossveins between RP , MA; 1r-m rather close to origin of RP ; 3r-m connecting RP 1, anterior branch of MA oblique. MA dichotomously forked; MP strongly zigzagged, pectinately branched; anterior trace, all two branches rather short, simple. Two long intramedian crossveins; 2im connects posterior branch of MA near its origin, MP. Proximal part of M, CuA mostly not preserved. Anterior trace of CuA simple, with one rather deeply forked branch. CuP, anal veins not preserved. FIGURE 12. Archiraphidia tranquilla (Scudder, 1890) . Holotype MCZ 246, the specimen as preserved. Scale bar = 4 mm. FIGURE 13. Archiraphidia tranquilla (Scudder, 1890) . Wing venation of the holotype MCZ 246. A, right forewing. B, right hind wing. C, left forewing. D, left hind wing. Scale bar = 4 mm (all to scale). Hind wing approximately 7.5 mm (estimated), 2.3 mm wide (length/width ratio 3.2). Costal space not preserved. ScP poorly preserved. In subcostal space, one crossvein detected, closing pterostigma proximally. Pterostigma slightly longer and narrower than in forewing. RA with one branch, which closes pterostigma distally; incorporated branch not detected; portion of RA distad pterostigma short, deeply curved. Two crossvein between RA, RP preserved; 3ra-rp within pterostigma, in its proximal half. Anterior trace of RP forked apically, with two branches proximad 3ra-rp, one distad. RP 1 deeply forked; anterior branch simple, posterior branch shallowly forked; RP 2, RP 3 simple. Three intraradial crossveins forming short gradate series, all between branches of RP (left wing); two intraradial crossveins between RP 1, origin of RP 2, and RP 2, RP 3 (right wing). Only one oblique distal crossvein between R, M detected (4r-m), connecting RP 1, anterior branch of MA. MA dichotomously (twice) forked. MP zigzagged, pectinately branched; anterior trace rather deeply forked; two branches simple, with their bases rather closely spaced (left wing), or one branch rather shallowly forked (right wing). One (2im) intramedian crossvein preserved. CuA incompletely preserved; anterior branch, one branch simple. 2m-cu connects MP, CuA. Basal portion of wing, CuP, anal veins not preserved. Specimen MCZ 4137, lectotype of Inocellia eventa ( Fig. 14 ). Male. Head long, details not clear. Prothorax relatively long, ca. 2.2 mm , 0.7 mm maximum width (caudally, in lateral view). Pterothorax, legs poorly preserved. Abdomen well preserved; with eight dark segments separated by broad pale intersegmental membrane between fifth to eighth segments. Apex of abdomen: Tergite 9 long (in lateral view); sternite 9 short; tergite 9, sternite 9 appear to form a ring; gonocoxites of segment 9 forms a large plate, its shape not entirely clear, probably somewhat elongate rectangular; gonapophyses of segment 9 (hypovalva) narrow anteriorly, appears broadened caudally. Segment 10 (ectoproct) appears nearly quadrate, without caudal elongation; gonostyli of gonocoxites 9 strongly curved apically upwards ( Fig. 15 ). Forewing approximately 9.5 mm long (estimated), 3.0 mm wide (length/width ratio 3.2). Costal space incompletely preserved, moderately broad. ScP relatively short; length from termination of ScP to pterostigma longer than length of pterostigma. Four preserved subcostal veinlets (branches of ScP) distad M+CuA fusion simple, widely spaced. In subcostal space, no crossvein between ScP, RA detected; crossvein closing pterostigma proximally straight. Pterostigma short, dark. RA with one branch, closing pterostigma distally; no branches within pterostigma detected ( Fig. 9 E); portion of RA distad pterostigma short, deeply curved. Two crossveins between RA, RP ; 2ra-rp located slightly distad termination of ScP; 3ra-rp short (very short in left wing), located within pterostigma, in its proximal half. Anterior trace of RP deeply forked apically, with posterior branch shallowly forked; two branches proximad 3ra-rp, one distad. RP 1 deeply forked, anterior branch simple, posterior shallowly forked. RP 2, RP 3 simple. Three (left wing), two (right wing) intraradial crossveins, between branches of RP . Three crossveins between RP , MA; 1rp-m rather close to origin of RP ; 3rp-m connecting RP 1, anterior branch of MA oblique. MA dichotomously (twice) forked. Anterior trace of MP strongly zigzagged, simple, pectinately branched; two short branches, one simple, other fused for long distance with anterior trace of CuA. Two long intramedian crossveins; 2im connects posterior branch of MA near its origin, MP. Proximal part of M, CuA mostly not preserved. Anterior trace of CuA simple, fused for long distance with basal-most branch of MP; with one forked branch. CuP simple. AA1 simple (left wing), very shallowly forked (right wing). AA2 incompletely preserved. Crossvein between AA1, AA2 (iaa) moderately long. FIGURE 14. Archiraphidia tranquilla (Scudder, 1890) (lectotype of A . eventa (Scudder, 1890) , MCZ 4137). A, the specimen as preserved (covered by Canada balsam). B, left forewing. C, left hind wing. D, right forewing. E, right hind wing. Scale bar = 4 mm (all to scale). Hind wing approximately 8.8 mm (estimated), 2.7 mm wide (length/width ratio 3.3). Costal space narrow; subcostal veinlets not preserved. ScP relatively short; length from termination of ScP to pterostigma longer than length of pterostigma. In subcostal space, one crossvein detected, closing pterostigma proximally. Pterostigma slightly longer and narrower than in forewing. RA with one preserved branch, which closes pterostigma distally; incorporated branch not detected; portion of RA distad pterostigma short, deeply curved. Three crossveins between RA, RP preserved; 2ra-rp somewhat distad termination of ScP; 3ra-rp within pterostigma, in its middle. Anterior trace of RP rather deeply forked apically (additional vein probably divided fork in right wing); with two branches proximad 3ra-rp, one distad. RP 1 deeply forked, each branch shallowly forked; RP 2, RP 3 simple. Three intraradial crossveins forming short gradate series, all between branches of RP . Only one oblique distal crossvein between R, M detected, connecting RP 1, anterior branch of MA. MA dichotomously (twice) forked. MP zigzagged; anterior trace rather deeply (right wing) or shallowly (left wing) forked; one rather deeply forked branch. One (distal) intramedian crossvein preserved (2im). CuA incompletely preserved; anterior trace, one branch simple. Basal and anal portion of wing not preserved. Specimen MCZ 253 , paralectotype of Inocellia eventa ( Fig. 16 ). Male. Head relatively short (ca. 1.8 mm including mouthparts), strongly narrowed caudally (in dorso-ventral view); antennae, palpi not preserved. Prothorax relatively long, slender, 2.0 mm long, ca. 0.4 mm (in anterior portion), ca. 0.5–0.6 mm maximum width (caudally). Abdomen poorly preserved, but surely male. FIGURE 15. Archiraphidia tranquilla (Scudder, 1890) (lectotype of A . eventa (Scudder, 1890) , MCZ 4137). Apex of abdomen (lateral view). gp9, gonapophyses of segment 9 (hypovalva); gs9, gonostyli of gonocoxites 9; gx9, gonocoxites of segment 9; S8, S9, sternites 8 and 9; T8, T9, T10, tergites 8 to 10. Scale bar = 400 µm. Forewing approximately 8.8 mm long (estimated), 2.8 mm wide (3.1). Costal space incompletely preserved, relatively broad. ScP relatively short; length from termination of ScP to pterostigma much longer than length of pterostigma. Five preserved subcostal veinlets (branches of ScP) distad fusion of M+CuA simple, widely spaced, except two which abnormally convergent towards ScP. In subcostal pace, one crossvein between ScP, RA; other crossvein closing pterostigma proximally nearly straight. Pterostigma short, dark. RA with two branches: one long, incorporated in pterostigma, other closing pterostigma distally ( Fig. 9 F); portion of RA distad pterostigma short, deeply curved. Anterior trace of RP forked apically: branches of fork divergent at right angle forming very broad, shallow forking; with two branches proximad 3ra-rp, one distad (wing apex poorly preserved). RP 1 forked, anterior branch simple, posterior rather deeply forked. RP 2 simple. RP 3 shallowly forked at least in right wing. Two crossveins between RA, RP ; 3ra-rp located within pterostigma, in its middle. Three intraradial crossveins between branches of RP ; distal-most crossvein long. Three crossveins between RP , MA; 1r-m rather close to origin of RP ; 3r-m connecting RP 1, MA oblique. MA dichotomously forked; MP strongly zigzagged, pectinately branched; anterior trace, all two branches rather short, simple. Two long intramedian crossveins; 2im connects posterior branch of MA and MP. M, CuA diverge at apparent R ( free stem of M+CuA absent). Anterior trace of CuA with one shallowly forked branch. CuP simple. One long intracubital crossvein (incompletely preserved). AA1 simple; AA2 incompletely preserved; crossvein between these (iaa) moderately short. Hind wing approximately 7.5 mm long, 2.5 mm wide (estimated 3.0 mm). Costal space, ScP poorly preserved. ScP short; length from termination of ScP to pterostigma much longer than length of pterostigma. In subcostal space, one crossvein detected, closing pterostigma proximally. Pterostigma narrower than in forewing. RA with two branches; one probably long, incorporated in pterostigma (only its basal part visible), other closing pterostigma distally ( Fig. 9 F); portion of RA distad pterostigma short, deeply curved. Two crossveins between RA, RP preserved; 2ra-rp located distad termination of ScP; 3ra-rp relatively short, within pterostigma (in middle portion). Anterior trace of RP forked apically: branches of fork divergent at right angle forming very broad, shallow fork; with two branches proximad 3ra-rp, one distad. RP 1 deeply forked; both branches rather deeply forked; RP 2, RP 3 incompletely preserved. Three-four intraradial crossveins between RP 1 to stem of RP (crossvein between RP 2, RP 3 abnormally doubled in left wing). M forked distad origin of RP ; mode of branching not visible. One (distal) intramedian crossvein (2im) preserved. Cu, anal veins not preserved. FIGURE 16. Archiraphidia tranquilla (Scudder, 1890) (paralectotype of A. eventa (Scudder, 1890) , MCZ 253). A, the specimen as preserved (covered by Canada balsam). B, right forewing, C, right hind wing. D, left forewing. E, left hind wing. Scale bar = 4 mm (all to scale). Type material. Holotype of A. tranquilla : MCZ 246 (Scudder’s number 4383), deposited in MCZ . A rather poorly preserved, incomplete male specimen in lateral aspect; all wings overlapping; head, prothorax and legs missing. Verbatim label data: [1] “Mus. Comparative Zoölogy, Cambridge, / No. 246 / Raphidia (?) tranquilla / Scud. / Florissant Miocene”; [2] 246 / type / Raphidia (?) / tranquilla Scud. / Florissant / Neuroptera : Raphididae [sic]”. The specimen has a red label glued to the rock that says “ Type .”, and black ink writing on the rock that says “246”. Types of A. eventa : lectotype (designated here) MCZ 4137 (Scudder’s number 9391), deposited in MCZ . An almost complete male specimen in lateral aspect; head poorly preserved. Verbatim label data: [1] “Mus. Comp. Zoöl. Cambridge, Mass. / No. 4137 / Inocellia eventa / Scud. / Florissant”; [2] 4137 / type / Inocellia / eventa Scud. / Florissant / Neuroptera : Inocellidae [sic]”. The specimen has a red label glued to the rock that says “ Type . / 4137”, and red ink writing on the rock that says “9391”. Paralectotype (designated here) MCZ 253 (Scudder’s number 8319), deposited in MCZ . An almost complete male specimen in dorso-lateral aspect; legs not preserved, right wings crumpled. Verbatim label data: [1] “Mus. Comp. Zoöl. Cambridge, Mass. / No. 253 / Inocellia eventa Scud. / Florissant Miocene”; [2] 253 / type / Inocellia / eventa Scud. / Florissant / Neuroptera : Inocelliidae ”. The specimen has a red label glued to the rock that says “ TYPE ”; black ink writing on the rock that says “253”, and red ink writing on the rock that says “8319”. Type locality. Florissant (precise collecting localities unknown). Remarks. The venation of these two specimens assigned to Archiraphidia eventa is so similar to that of the holotype of A . tranquilla that we do not doubt in their conspecifity. Archiraphidia tumulata and A. tranquilla are also very similar; the minor differences in morphologies noted in their diagnoses may be intraspecific variation. Archiraphidia tranquilla has only been figured by Scudder, once, with errors (1890), and neither specimens of A. eventa have been illustrated. Scudder (1890) described the latter species based on two these specimens (i.e., syntypes ), both of which are designated in MCZ as the type . We designate here a better preserved specimen (MCZ 4137) as the lectotype , and other (MCZ 253) as the paralectotype .